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EIGHTH EDITION

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lcs~

Inderbir Singh
G P Pal
HUMAN
EMBRYOLOGY
EIGHTH EDITION

INDERBIR SINGH
G.P. PAL
ft
© Inderbir Singh, 1976, 1979, 1981, 1985, 1986, 1996, 2001, 2007

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First published, 1976


Reprinted, 1977 (2 times)
Second edition, 1979
Third edition, 1981
Fourth edition, 1985
Fifth edition, 1986
Reprinted, 1987-1995 (9 times)
Sixth edition, 1996
Reprinted, 1997-2001 (7 times)
Seventh edition, 2001
Reprinted, 2002-2006 (14 times)
Eighth edition, 2007
Reprinted, 2009 (thrice)
Reprinted, 2010

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i Preface to the Eighth Edition

The eighth edition of HUMAN EMBRYOLOGY comes with several departures from earlier editions.
The book is now printed in a larger format. The drab black and white pages have given place to those
printed in full colour, and on much better paper. Paragraphs dealing with details that beginners can
skip are printed on a light blue background.
These cosmetic changes are accompanied by extensive revision of text to make it up to date. A new
chapter on 'Molecular Control of Development' has been added.
With printing in full colour, it has been possible to make a quantum jump in the quality of illustrations.
All illustrations are now in colour. Apart from addition of colour many old figures have been redrawn
and relabeled. Several new figures and tables have been added.
We are greatly indebted to Macmillan India Ltd. for their enthusiastic and wholehearted support
towards making this edition what it is.
We continue to be highly obliged to the numerous students and teachers who have written to us.
Their letters have been highly encouraging and useful.

June 2007 INDERBIR SINGH


G.P. PAL
Preface to the First Edition

This hook on human embryology has been written keeping in mind the requirements of undergraduate
medical students. The subject of embryology has traditionally been studied from imported textbooks
of anatomy or of embryology. Experience has shown that the treatment of the subject in most of these
books is way above the head of the average medical student in India. T h e difficulty has increased from
year to year as there has been, and continues to be, progressive deterioration in the standards of the
reaching of English in our schools and colleges. The combination of unfamiliar sophistications of
language and of an involved technical subject, has very often left the student bewildered.
In this book care has been taken to ensure that the text provides all the information necessary for an
intelligent understanding of the essential features of the development of various organs and tissues of
the human body. At the same time, several innovations have been used to make the subject easy to
understand.
Firstly, the language has been kept simple. Care has been taken not to compress too many facts into
an involved sentence. New words are clearly explained.
Secondly, simultaneous references to the development of more than one structure have been avoided
as far as possible. While this has necessitated some repetition, it is hoped that this has removed one of
rhe greatest factors leading to confusion in the study of this subject.
Thirdly, almost every step in development has been shown in a simple, easy to understand, illustration.
To avoid confusion, only structures relevant to the discussion are shown. As far as possible, the
drawings have been oriented as in adult anatomy to facilitate comprehension.
Fourthly, the chapters have been arranged so that all structures referred to at a particular stage
have already been adequately introduced.
In an effort of this kind it is inevitable that some errors of omission, and of commission, are liable
to creep in. To obviate as many of these as possible a number of eminent anatomists were requested to
read through the text. Their suggestions have greatly added to the accuracy and usefulness of this
book. Nevertheless, scope for further improvement remains, and the author would welcome suggestions
to this end both from teachers and from students.

Rohtak INDERBIR SINGH


January 1976
Contents

Preface to the Eighth Edition

Preface to the First Edition

1. Some Preliminary Considerations I

2. Spermatogenesis and Oogenesis 9

3. The Menstrual Cycle 25

4. Formation of Germ Layers M

5. Further Development of F.mbryonic Disc 46

6. The Placenta 59

7. Formation of Tissues of the Body 76

8. The Skin and its Appendages 9S

9. The Pharyngeal Arches 104

10. The Skeleton 116

11. Face, Nose and Palate 126

12. Alimentary System I: Mouth, Pharynx and Related Structures 138

13. Alimentary System II: Gastrointestinal Tract 148

14. Liver, Pancreas, Spleen; Respiratory System; Body Cavities 166


Contents

15. Cardiovascular System 191

16. Urogenital System 237

17. The Nervous System 268

18. Hypophysis Cerebri, Pineal, Adrenal 298

19. The Eye and Ear 302

20. Fate of the Germ Layers 319

21. Some General Considerations 322

22. Molecular Control of Development 330

Index 349
Chapter

Some Preliminary Considerations

HIGHLIGHTS cells resulting from a mitotic division are


similar to the parent cell, and have the same
• Embryology is the study of the development number of chromosomes (46).
of an individual before birth. • A special kind of cell division takes place in
• D u r i n g the first t w o m o n t h s we call the the testis and ovary for formation of gametes.
developing individual an embryo. After that It is called meiosis. The gametes resulting from
we call it a fetus. meiosis have t h e h a p l o i d n u m b e r of
chromosomes (23). The various gametes
• The testis is the male sex organ or male gonad.
formed do not have the same genetic content.
The ovary is the female sex organ or female
gonad. They produce gametes.
• Male gametes produced by the testis are called
spermatozoa. The process is called spermato-
genesis.
• Female gametes produced by the ovary are
called ova. The process is called oogenesis.
Spermatogenesis and oogenesis are together
called gametogenesis.
• Fertilization t a k e s place w h e n one WHAT IS EMBRYOLOGY?
spermatozoon enters an ovum. The fused ovum
and sperm form the zygote. Every individual spends the first nine months (266
• C h a r a c t e r s of p a r e n t s are t r a n s m i t t e d to days or 38 weeks to be exact) of its life within the
offspring through codes borne on strands of w o m b (uterus) of its mother. During this period it
DNA. Genes are made of such strands of DNA. develops from a small one-celled structure to an
They are located on chromosomes. organism having billions of cells. Numerous tissues
and organs are formed and come to function in
• A typical cell c o n t a i n s 46 c h r o m o s o m e s
perfect harmony. The most spectacular of these
(= diploid number).
changes occur in the first two months; the unborn
• A gamete contains 23 chromosomes {= haploid baby acquires its main organs and just begins to
number). be recognizable as h u m a n . D u r i n g these t w o
• The diploid number of chromosomes is restored m o n t h s we call the developing individual an
as a result of fertilization. embryo. From the third month until birth we call
it a fetus.
• Multiplication of cells takes place by cell
division. The usual method of cell division, Embryology is the study of the formation and
seen in most tissues, is called mitosis. Daughter development of the embryo (or fetus) from the
Human Embryology

moment of its inception up to the time when it is is referred to as the diploid (or double) number.
born as an infant. However, in spermatozoa and ova the number of
chromosomes is only half the diploid number, i.e.
twenty-three. This is called the haploid (or half)
Why a Medical Student should number. After fertilization the resulting zygote has
Study Embryology? twenty-three chromosomes from the sperm (or
1. This subject tells us h o w a single cell (the father), a n d t w e n t y - t h r e e from the o v u m (or
fertilized ovum) develops into a newborn, mother). The diploid number is thus restored.
containing numerous tissues and organs.
2. This knowledge helps us understand many
Autosomes and Sex Chromosomes
complicated facts of adult anatomy.
3. Embryology helps us understand why some The forty-six chromosomes in each cell can be
children are born with o r g a n s t h a t are divided into forty-four autosomes and two sex
abnormal. Appreciation of the factors res- chromosomes. T h e sex chromosomes may be of
ponsible for maldevelopment assists us in two kinds, X or Y. In a man there are forty-four
preventing, or treating such abnormalities. autosomes, one X-chromosome and one
Y-chromosome; while in a woman there are forty-
four autosomes and two X-chromosomes in each
Gonads and Gametes cell (Fig. 1.1). When we study the forty-four
The cells that carry out the special function of autosomes we find that they in fact consist of
reproduction are called gametes. The development twenty-two pairs, the two chromosomes forming
of a new individual begins when one male gamete a pair being exactly alike {homologous
{sperm or spermatozoon) meets and fuses with one chromosomes). In a w o m a n the two
female gamete (ovum or oocyte). The process of X-chromosutnes form another such pair; in a man
fusion of male a n d female g a m e t e s is called this pair is r e p r e s e n t e d by o n e X- a n d o n e
fertilization. The fused ovum and spermatozoon Y-chromosome. One chromosome of each pair is
form the zygote. The zygote later develops into derived from the mother and the other from the
an embryo and then into a fetus. father.
The male sex cells (spermatozoa) are produced
in the male gonads (testes) while the female sex Chromosome Structure
cells (ova) a r e p r o d u c e d in female g o n a d s
In a resting cell, chromosomes are not visible under
(ovaries). The formation of spermatozoa in the
a light microscope as their chromatin material is
testis is called spermatogenesis while the formation
highly dispersed. However, during cell division
of ova in the ovary is called oogenesis. The t w o
the chromatin network in the nucleus becomes
are collectively referred to as gametogenesis.
condensed into a number of chromosomes. The
To understand the structure of gametes and to
appearance of a typical chromosome is illustrated
study how they are formed, it is necessary to first
in Fig. 1.2. It is made up of t w o rod-shaped
review some facts regarding chromosomes and cell
structures or chromatids placed m o r e or less
division.
parallel to each other. The chromatids are united
to each other at a light staining area called the
SOME FACTS ABOUT CHROMOSOMES centromere (or kinetochore). Each chromatid has
two arms, one on either side of the centromere.
Haploid and Diploid Chromosomes Individual chromosomes differ from one another
The number of chromosomes in each cell is fixed in total length, in the relative length of the two
for a given species and in man it is forty-six. This arms and in various other characteristics; these
Some Preliminary Considerations

MAN

WOMAN

Fig. 1.1 Number of chromosomes in the somatic cells of a man and of a w o m a n .

Significance of Chromosomes
Satellite.
The entire human body develops from the fertilized

11
Secondary
constriction"*" ovum. It is, therefore, obvious that the fertilized
Short arm ovum contains all the information necessary for
of chromatid formation of the numerous tissues and organs of
Centromere—* the body, and for their orderly assembly and
function. Each cell of the body inherits from the
\ fertilized ovum, all the directions that are necessary
Long arm \ for it to carry out its functions throughout life.
of chromatid '
This tremendous volume of information is stored
within the chromosomes of each cell.

J u • Each c h r o m o s o m e bears on itself a very


large number of structures called genes.
• Genes are made up of a nucleic acid called
Fig. 1.2 Diagram to show the parts of a typical
deoxyribonucleic acid (or DNA) and all
chromosome.
information is stored in the molecules of
this substance.
differences enable us to identify each chromosome • Genes are involved in synthesis of proteins.
individually. Classification of chromosomes in this • Proteins are the most important constituents
way is called karyotyping. Karyotyping makes of our body. They make up the greater part
it possible for us t o detect a b n o r m a l i t i e s in of each cell and of intercellular substances.
c h r o m o s o m e number or in individual chromo- Enzymes, hormones and antibodies are also
somes. proteins.
Human Embryology

The nature and functions of a cell depend MITOSIS


o n the p r o t e i n s synthesized by it. It is,
therefore, not surprising that one cell differs M a n y cells of the body have a limited span of
from another because of the differences in functional activity, at t h e end of which they
the proteins that constitute it. undergo division into t w o d a u g h t e r cells. The
daughter cells in turn have their o w n span of
We n o w k n o w t h a t g e n e s c o n t r o l t h e activity, followed by another division. The period
d e v e l o p m e n t a n d f u n c t i o n i n g of c e l l s , by during which the cell is actively dividing is the
d e t e r m i n i n g w h a t t y p e s of p r o t e i n s will be p h a s e of m i t o s i s . T h e p e r i o d b e t w e e n t w o
synthesized within t h e m . T h u s genes play an successive divisions is called the interphase.
important role in the development of tissues and Mitosis is conventionally divided into a number
organs of an individual. of stages called prophase, metaphase, anaphase
T r a i t s ( c h a r a c t e r s ) of a n i n d i v i d u a l a r e a n d telophase. T h e sequence of events of the
d e t e r m i n e d by genes carried on his (or her) mitotic cycle is best understood starting with a
chromosomes. As we have seen half of these are cell in telophase. At this stage each chromosome
inherited from the father and half from mother. consists of a single chromatid (Fig. 1.3G). With
the progress of telophase, the chromatin of the
CELL DIVISION c h r o m o s o m e uncoils a n d e l o n g a t e s a n d the
c h r o m o s o m e can no longer be identified as
Multiplication of cells takes place by division of s u c h . H o w e v e r , it is b e l i e v e d t o r e t a i n its
pre-existing cells. Such multiplication constitutes identity during the interphase (which follows
an essential feature of embryonic development. telophase). This is shown diagrammatically in
Cell multiplication is equally necessary after the Fig. 1.3 A.
b i r t h of the i n d i v i d u a l for g r o w t h a n d for During a specific period of the interphase, the
replacement of dead cells. We have seen that DNA content of the chromosome is duplicated so
c h r o m o s o m e s within the nuclei of cells carry- that another chromatid identical to the original
genetic information that controls the development one is formed; the chromosome is n o w made up
and functioning of various cells and tissues; and, of two chromatids (Fig. I.3B). When mitosis begins
therefore, of the body as a whole. When a cell (i.e. d u r i n g p r o p h a s e ) the c h r o m a t i n of the
divides it is essential t h a t the entire genetic
chromosome becomes gradually more and more
information within it be passed on to both the
c o i l e d so t h a t t h e c h r o m o s o m e b e c o m e s
daughter cells resulting from the division. In other
r e c o g n i z a b l e as a t h r e a d - l i k e s t r u c t u r e t h a t
words, the daughter cells must have chromosomes
gradually acquires a rod-like appearance
identical in number (and in genetic content) to
(Fig. 1.3C). Towards the end of prophase, the two
those in the mother cell. This type of cell division
c h r o m a t i d s c o n s t i t u t i n g the c h r o m o s o m e
is called mitosis.
become distinct (Fig. 1.3D) and the chromosome
A different kind of cell division called meiosis n o w has the typical s t r u c t u r e i l l u s t r a t e d in
occurs during the formation of the gametes. This Fig. 1.2.
consists of two successive divisions called the first W h i l e t h e s e c h a n g e s a r e o c c u r r i n g in
and second meiotic divisions. The cells resulting chromosomes, a number of other events are also
from these divisions (i.e. gametes) differ from other taking place. T h e t w o centrioles separate and
cells of the b o d y in t h a t (a) the n u m b e r of
move to opposite poles of the cell. They produce
c h r o m o s o m e s is reduced to half the n o r m a l
a n u m b e r of microtubules that pass from one
number, and (b) the genetic information in the
c e n t r i o l e to t h e o t h e r a n d form a spindle.
various gametes produced is not identical.
Meanwhile the nuclear membrane breaks down
Some Preliminary Considerations

LATE INTERPHASE
DNAof each
chromosome has under-
gone duplication.

Fig. 1.3 Scheme to show the main stages of mitosis.

and nucleoli d i s a p p e a r (Fig. 1.3D). W i t h the longitudinally into two so that the chromatids now
formation of the spindle, chromosomes move to a become independent chromosomes.
position midway between the two centrioles (i.e. At this stage the cell can be said to contain
at the equator of the cell) where each chromosome forty-six pairs of chromosomes. One chromosome
becomes attached to microtubules of the spindle of each such pair now moves along the spindle to
by its centromere. This stage is referred to as cither pole of the cell (Fig. 1.3F). This is followed
metaphase (Fig. 1.3E). In the anaphase the by telophase in which the two daughter nuclei are
c e n t r o m e r e of e a c h c h r o m o s o m e s p l i t s formed by appearance of nuclear m e m b r a n e s .
Human Embryology

C h r o m o s o m e s gradually elongate and become


indistinct. Nucleoli reappear. T h e centriole is
duplicated at this stage or in early interphase 1. Chromosomes
become visible. Each
(Fig. 1.3G). chromosome is made
The division of the nucleus is accompanied by up of two chromatids
the division of the cytoplasm. In this process (but these cannot be
made out separately).
(cytokinesis) the organelles are presumably
duplicated and each daughter cell comes to have
a full complement of them.

MEIOSIS
2. Two homologous
As a l r e a d y s t a t e d m e i o s i s c o n s i s t s of t w o chromosomes come
successive divisions called the first and second to lie side by side
forming a bivalent.
meiotic divisions. During the interphase preceding
the first division, duplication of the D N A content
of chromosomes takes place as in mitosis. As a
result, another chromatid identical to the original
one is formed. Thus each chromosome is now made
up of two chromatids.
3. Four chromatids
First Meiotic Division are now distinct and
form a tetrad.
1. The prophase of the first meiotic division is
prolonged and is usually divided into a
number of stages as follows:

(a) Leptotene. T h e c h r o m o s o m e s b e c o m e 4. The two 'central'


visible (as in m i t o s i s ) . A l t h o u g h each chromatids become
chromosome consists of two chromatids, coiled on each other
so that they cross at
these cannot be distinguished at this stage a number of places.
(Fig. 1.4A). Only one such crossing
(b) Zygotene. We have seen that the forty-six is shown.
chromosomes in each cell in fact consist
of t w e n t y - t h r e e p a i r s ( t h e X a n d Y
chromosomes of a male being taken as a
pair). The t w o chromosomes of each pair 5. The chromosomes
come to lie parallel to each other, and are now separate. The
closely apposed. This pairing of chromo- 'central' chromatids
'break' at the points of
somes is also referred to as synapsis or crossing and unite
conjugation. The two chromosomes with the opposite
together constitute a bivalent (Fig. 1.4B). chromatid.
(c) Pachytene. T h e t w o chromatids of each
chromosome become distinct. The bivalent Fig. 1.4 Stages in the prophase of the first
now has four chromatids in it and is called meiotic division.
Some Preliminary Considerations

over. For sake of simplicity only one such


crossing is shown in Fig. 1.4D. At the site
where the chromatids cross they become

iX adherent; the points of adherence are called


chiasmata.
(d) D i p l o t c n e . T h e t w o c h r o m o s o m e s of a
bivalent n o w try to move a p a r t . In the
process, the chromatids involved in crossing
METAPHASE over 'break' at the points of crossing and
The nuclear membrane disappears. Spindle forms the 'loose' pieces become attached to the
and chromosomes are attached to it by their o p p o s i t e c h r o m a t i d . T h i s r e s u l t s in
centromeres. exchange of genetic material between these
chromatids. A study of Fig.l.4E will show
that each of the four chromatids of the
tetrad now has a distinctive genetic content.

2. This is followed by metaphase. As in mitosis


the forty-six chromosomes become attached
t o the spindle at the e q u a t o r , the t w o
chromosomes of a pair being close to each
ANAPHASE other (Fig. 1.5A).
One entire chromosome of the pair moves to either 3. The anaphase differs from that in mitosis
pole. Note thai the centromeres do not divide. in t h a t there is no splitting of the
centromeres. O n e encire c h r o m o s o m e of
each pair moves to each pole of the spindle
(Fig. 1.5B). The resulting daughter cells,
therefore, have twenty-three chromosomes,
each made up of two chromatids (Fig. 1.5C).
4. The anaphase is followed by the telophase
in which t w o daughter nuclei are formed.
TELOPHASE The division of the nucleus is followed by
Note that the chromosomes in each cell have been
reduced to the haploid number.
division of the cytoplasm.

Second Meiotic Division


Fig. 1.5 Metaphase (A), anaphase (B), and
telophase (C) of the first meiotic division. The first meiotic division is followed by a short
interphase. This differs from the usual interphase
a tetrad. There are t w o central a n d t w o in t h a t there is no duplication of DNA. Such
p e r i p h e r a l c h r o m a t i d s , o n e from e a c h duplication is unnecessary as chromosomes of cells
c h r o m o s o m e (Fig. 1.4C). An i m p o r t a n t resulting from the first division already possess
event n o w takes place. T h e two central two chromatids each (Fig. 1.5C).
c h r o m a t i d s (one belonging to each The second meiotic division is similar to mitosis.
chromosome of the bivalent) become coiled However, because of the crossing over that has
over each other so t h a t they cross at a occurred during the first division, the daughter
number of points. This is called crossing cells are not identical in genetic content (Fig. 1.6).
Human Embryology

spermatozoa and ova. It is, therefore, not


surprising t h a t no t w o persons (except
identical twins) are alike.

A NOTE ON CHRONOLOGY OF
EMBRYOLOGICAL EVENTS

In an earlier section it has been emphasized


that one of the main objectives of the study of
e m b r y o l o g y , by m e d i c a l s t u d e n t s , is t o
u n d e r s t a n d t h e c a u s a t i o n of c o n g e n i t a l
a n o m a l i e s . In this c o n n e c t i o n it has been
observed that if a growing embryo is exposed
Fig. 1.6 Daughter cells resulting from the second
to c e r t a i n a g e n t s (chemical o r p h y s i c a l ) ,
meiotic division. These are not alike
abnormalities in development can result. Such
because of the crossing-over during the
first meiotic division.
agents are called teratogens. The list of known
teratogens keeps increasing. It has also been
Significance of Meiosis observed that some particular organs are most
sensitive to teratogens when they are passing
Why are no two persons alike? through critical phases in their development.
T h i s p e r i o d of g r e a t e s t s u s c e p t i b i l i t y t o
1. In this kind ofcell division there is reduction
teratogens differs from organ to organ.
of the number of chromosomes from diploid
to haploid. At the time of fertilization the The importance of having a knowledge of
d i p l o i d n u m b e r (46) is r e s t o r e d . T h i s the timing of embryologies I events thus becomes
provides consistency of chromosome number obvious. In the early stages of development,
from generation to generation. the age of an embryo is reckoned in days. Later,
2. As stated earlier, the forty-six chromosomes when events are less d r a m a t i c , age can be
of a cell consist of twenty-three pairs, one expressed in weeks or months. However, the
chromosome of each pair being derived from exact age of an embryo is not always known.
the mother and one from the father. During An estimate can be made by observing the size
the first meiotic division the chromosomes of the embryo (expressed as C.R. length), or
derived from the father and those derived some other feature like the number of somites.
from the mother are distributed between the In most textbooks of embryology, there are
daughter cells entirely at random. numerous references to the timing of embryonic
3. This, along with the phenomenon of crossing events (most commonly in terms of C.R. length}.
over, results in thorough shuffling of the The disadvantage of doing so is that it adds yet
genetic material so that the cells produced one more complication to the understanding of
as a result of various meiotic divisions (i.e. an already intricate subject. Because of this
ova or spermatozoa) all have a distinctive r e a s o n r e f e r e n c e s t o t h e t i m e t a b l e of
genetic content. development have been kept to the minimum
in the main text of this book. However, a
4. A t h i r d step in this process of genetic
timetable of events is a d d e d at the end of
shuffling takes place at fertilization when
chapters wherever relevant.
there is a combination of randomly selected
Chapter

2 Spermatogenesis and Oogenesis

HIGHLIGHTS
A spermatozoon has a head, a neck, a middle
piece and a principal piece or tail (Fig, 2.1).
Stages of spermatogenesis are summarized in
Fig. 2.5.
S p e r m a t o z o a are derived from r o u n d e d
spermatids. The process of conversion of a
s p e r m a t i d to a s p e r m a t o z o o n is called
spermiogenesis {Fig.2.6).
Stages of oogenesis are s u m m a r i z e d in
Fig. 2.8.
An ovarian follicle is a rounded structure that
contains a developing ovum surrounded by
follicular cells. The follicle has a fluid filled
cavity (Fig. 2.12).
O v a r i a n follicles have a cellular covering
called the theca interna. The cells of the theea In this chapter we shall first study the structure of
interna produce oestrogens (Fig. 2.13). a m a t u r e spermatozoon {male sex cell) before
considering its formation in the testis.
The follicle gradually increases in size and
finally bursts to expel the ovum. This process
of shedding of the ovum is called ovulation. STRUCTURE OF A MATURE
SPERMATOZOON
The corpus luteum is formed by enlargement
and transformation of follicular cells, after A spermatozoon is a highly specialized, free swim-
shedding of the ovum {Fig. 2.17), The corpus ming, actively motile cell. The spermatozoon has
luteum secretes progesterone, which is essential a head, a neck, a middle piece and a principal piece
for maintenance of pregnancy. or to'/(Fig. 2.1). An axial filament passes through
the middle piece and extends into the tail. The
spermatozoon measures about 60 um in length.

The Head
The head of the human spermatozoon is piriform
in shape and measures 4 urn in length. It is derived
Chapter

2 Spermatogenesis and Oogenesis

HIGHLIGHTS
A spermatozoon has a head, a neck, a middle
piece and a principal piece or tail (Fig. 2.1).
Stages of spermatogenesis are summarized in
Fig. 2.5.
S p e r m a t o z o a are derived from r o u n d e d
spermatids. The process of conversion of a
s p e r m a t i d to a s p e r m a t o z o o n is called
spermiogettesis (Fig.2.6).
Stages of oogenesis are s u m m a r i z e d in
Fig. 2.8.
An ovarian follicle is a rounded structure that
contains a developing ovum surrounded hy
follicular cells. The follicle has a fluid filled
cavity (Fig. 2.12).
O v a r i a n follicles have a cellular covering
called the theca interna. The cells of the theca In this chapter we shall first study the structure of
interna produce oestrogens (Fig. 2.13). a mature spermatozoon (male sex cell) before
considering its formation in the testis.
The follicle gradually increases in size and
finally bursts to expel the ovum. This process
of shedding of the ovum is called ovulation. STRUCTURE OF A MATURE
SPERMATOZOON
The corpus Iuteum is formed by enlargement
and transformation of follicular cells, after A spermatozoon is a highly specialized, free swim-
shedding of the ovum (Fig. 2.17). The corpus ming, actively motile cell. The spermatozoon has
Iuteum secretes progesterone, which is essential a bead, a neck, a middle piece and a principal piece
for maintenance of pregnancy. or tot/(Fig. 2.1}. An axial filament passes through
the middle piece and extends into the tail. The
spermatozoon measures about 60 urn in length.

The Head
The head of the human spermatozoon is piriform
in shape and measures 4 pm in length. It is derived
Spermatogenesis and Oogenesis

Coarce petal "•>!•»•/ - ...*'. A v >?•**.


^\ . Fibrous sheath
;».?.'.'«.#:T't.\»-.'«V^4--
•• >..•.•' °°-6 ^ •:•:••! i

.' %•••#.;:;.•.•.•.:.•:.•-•?» <p


• " \ t ° i •%•.•"• «.••••.•/ '
Plasma membrane

Fig. 2.3 Transverse section across the principal Fig. 2.4 Microscopic structure of a seminiferous tubule,
piece (tail) of a spermatozoon to show
the arrangement of fibrils. which fits into a depression (implantation
fossa) in the head.
s u r r o u n d e d by a spiral sheath m a d e u p of 3. In addition to the doublets, the axial filament
mitochondria. contains nine coarser petal-shaped fibrils, one
The axial filament is actually composed of such fibril lying just outside each doublet.
several fibrils arranged as illustrated in Fig. 2 . 3 .
There is a pair of central fibrils, surrounded by SPERMATOGENESIS
nine pairs (doublets) arranged in a circle around
the central pair. The whole system of fibrils is In a h u m a n m a l e , the f o r m a t i o n of g a m e t e s
k e p t in p o s i t i o n by a s e r i e s of c o v e r i n g s . ( s p e r m a t o z o a ) t a k e s p l a c e o n l y d u r i n g the
Immediately outside the fibrils there is a fibrous reproductive period, which begins at the age of
sheath. In the region of the middle piece the fibrous puberty (12 to 16 years) and continues even through
s h e a t h is s u r r o u n d e d by s p i r a l l y a r r a n g e d old age. Spermatozoa are formed in the walls of
mitochondria. Finally, the entire spermatozoon is the seminiferous tubules of the testes. If we look at
enclosed in a plasma membrane. one of these tubules under a microscope, we find
that there are many cells of different sizes and
Some Further Details shapes (Fig. 2.4). Most of these represent stages in
1. T h e c h r o m a t i n in t h e h e a d of t h e the formation of spermatozoa, but some (called
spermatozoon is extremely condensed. This Sertoli cells) have only a supporting function. The
makes the head highly resistant to various v a r i o u s cell-stages in s p e r m a t o g e n e s i s are as
physical stresses. T h e chemical basis for follows (the number of chromosomes at each stage
condensation is the replacement of histones is given in brackets) (Fig. 2.5).
by protamines.
2. The basal body is made up of nine segmented 1. The spermatogonia (type A) or germ cells
rod-like structures. O n its proximal side (i.e. (44 + X + Y) divide mitotically, to give rise
towards the head of the spermatozoon) the to more spermatogonia of type A, and also
basal body has a convex articular surface to spermatogonia of type B (Fig. 2.5).
Human Embryology

4. Each secondary spermatocyte has


22 + X or 22 + Y chromosomes. It
SPERMATOGONIUM divides to form two spermatids. This
TYPE A is the second meiotic division and
this time there is no reduction in
chromosome number.
5. Each spermatid (22 + X or 22 + Y)
SPERMATOGONIUM g r a d u a l l y c h a n g e s its s h a p e t o
TYPEB V X+ Y become a spermatozoon. This
p r o c e s s of t r a n s f o r m a t i o n of a
circular s p e r m a t i d to a s p e r m a -
tozoon is called spermiogenesis.

Spermiogenesis

T h e p r o c e s s by w h i c h a s p e r m a t i d
b e c o m e s a s p e r m a t o z o o n is c a l l e d
spermiogenesis (or spermateleosis)
(Figs 2.2 and 2.6). The spermatid is a
more or less circular cell containing a
nucleus, Golgi apparatus, centriole and
mitochondria. All these components take
part in forming the spermatozoon. The
nucleus forms the head. The Colgi
a p p a r a t u s is t r a n s f o r m e d i n t o t h e
acrosomic cap. The centriole divides into
two parts that are at first close together:
SPERMATIDS the axial filament appears to grow out
of them. O n e centriole becomes spherical
transformed and comes to lie in the neck. According
into to some w o r k e r s , the o t h e r centriole
forms the basal body, but according to
some others it forms the annulus. The
SPERMATOZOA part of the axial filament between the
neck a n d t h e a n n u l u s , b e c o m e s
Fig. 2.5 Stages in spermatogenesis. Note the number of s u r r o u n d e d by m i t o c h o n d r i a , a n d
chromosomes at each stage.
together with these forms the middle
2. The spermatogonia (type B) (44 + X + Y) piece. The remaining part of the axial filament
enlarge, or undergo mitosis, to form primary elongates to form the principal piece or tail. Most
spermatocytes. of the cytoplasm of the spermatid is shed, but the
3. The primary spermatocytes {44 + X + Y) now cell m e m b r a n e persists as a covering for the
divide so t h a t each of t h e m forms t w o spermatozoon.
secondary spermatocytes. This is the first The process of spermatogenesis, including
meiotic division: it reduces the number of spermiogenesis, requires about two months for its
chromosomes to half. completion.
Spermatogenesis and Oogenesis

Head

Cell
complex membrane

Axial filament -
Membrane covering sperm -

Fig. 2.6 Stages in spermiogenesis. Also see Fig. 2.2.

of capacitation, the glycoprotein coat and seminal


Maturation and Capacitation
proteins lying over the surface of the spermatozoon
of Spermatozoa
are altered. S p e r m a t o z o a usually u n d e r g o
W h e n first formed in s e m i n i f e r o u s t u b u l e s , capacitation in the uterus or uterine tube, under
spermatozoa are immature. They are non-motile the influence of substances secreted by the female
and incapable of fertilizing an ovum. A current of genital tract. W h e n a s p e r m a t o z o o n comes in
fluid in seminiferous tubules carries spermatozoa contact with the zona pellucida, changes take
from the testis to the epididymis. Here they are place in the membranes over the acrosome and
stored and undergo maturation. As spermatozoa enable the release of lysosomal enzymes. This is
pass t h r o u g h the epididymis they u n d e r g o a called the acrosome reaction. Some enzymes help
process of maturation. Changes take place in in digesting the zona pellucida and in penetration
glycoproteins of the plasma membrane covering of the spermatozoa through it. Changes in the
the sperm h e a d . S p e r m a t o z o a acquire some properties of the zona pellucida constitute the zona
motility after maturation, but become fully motile reaction.
only after ejaculation when they get mixed up with
secretions of the p r o s t a t e gland and seminal
vesicles. Difference between
Spermatogenesis and Spermiogenesis
Spermatozoa acquire the ability t o fertilize an
ovum only after they have been in the female S p e r m a t o g e n e s i s is t h e c o m p l e t e p r o c e s s of
genital tract for some time. This final step in their formation of a spermatozoon from a spermato-
maturation is called capacitation. In the process gonium. It includes the first and second meiotic
Human Embryology

/•#-••»'."•"•'6 •••'.•:;.• 14'.::. .''*•*>-.


4J,.;**: ••.' .--.:.. . '.•• '• \ •.', '*'•»!• '
• **,<''Mi ,,,. ' . : "-—
••.->'••••'•••''•••*...:. . . : -.••
"V
• »*&KN1 &fe' •.'•'•• /.-. -•" •' • " " -
*'.• % ° $$ • •••• •••; M&

.»••*>• . 11 12 • .'• -^ • • :-'•


•"::$' ' ' ' ' ' '.' . ii'teiii ... '.'@l'6 •','•%'''•''

••••.;£:to.:*-*.*..-.t: ••»'*• "."•••

Fig. 2.7 Panoramic view of the structure of an ovary.

division and spermiogenesis. On the other hand,


spcrmiogcncsis is the process of transformation of
' 4 4
OOGONIUM | a rounded spermatid into a spermatozoon.
)
enlarges
10 form
I The female gonad is the ovary. It has an outer
PRIMARY /' 44 part called the cortex and an inner part, the
OOCYTE I
X+X medulla (Fig. 2.7). The cortex contains many large
round cells called oogonia. All the oogonia to be
First ^

1 meiotic
division
^@
used throughout the fertile life of a woman are
produced at a very early stage (possibly before
birth) and do not multiply thereafter.
SECONDARY /
OOCYTE 1 22+X
k First
polar body Ova are derived from oogonia as shown in
Fig. 2.8. Note how similar the process is to
spermatogenesis. However, there are important
1
Second
meiotic
division ^2+Xj differences as well.

c)
Second
OVUM ( polar body Differences between
Spermatogenesis and Oogenesis
1. Observe that whereas one primary
Fig. 2.8 Stages in oogenesis (Compare each stage spermatocyte gives rise to four spermatozoa,
with the corresponding one in Fig. 2.5). one primary oocyte forms only one ovum.
Spermatogenesis and Oogenesis

2. When the primary spermatocyte divides, its • If fertilization does not occur, the secondary
cytoplasm is equally distributed between the oocyte fails to complete the second meiotic
two secondary spermatocytes formed. division, and degenerates in about 24 hours
However, when the primary oocyte divides, after ovulation.
almost all its cytoplasm goes to the daughter In each menstrual cycle, 5 to 30 primary
cell, which forms the secondary oocyte. The oocytes start maturing, but only one of them
other daughter cell (first polar body}, receives reaches maturity and is ovulated. The
half the chromosomes of the primary oocyte, remaining degenerate.
but almost no cytoplasm. The first polar body • During the entire reproductive life of a
is, therefore, formed merely to get rid of female only around 400 ova are discharged
unwanted chromosomes. (out of 40,000 primary oocytes available).
Further Details
Formation of Ovarian Follicles
• In the late fetal period primary oogonia
enlarge to form primary oocytes. We have seen that ova develop from oogonia
" At the time of birth all primary oocytes are present in the cortex of the ovary. The oogonia
in the prophase of the first meiotic division. are surrounded by other cells that form the stroma.
Their number is about 40,000. These stromal cells form ovarian or Graafian
The primary oocytes remain in prophase and follicles that surround ova and protect them. The
do not complete their first meiotic division stages in the formation of a follicle are as follows:
until they begin to mature and are ready to
ovulate. 1. Some cells of the stroma become flattened
• The reproductive period of a female is bet- and surround an oocyte (Fig. 2.9). These
ween 12 to 50 years of age. With each mens-
trual cycle, a few primary oocytes (about 5
to 30) begin to mature and complete the first
meiotic division shortly before ovulation.
• The first meiotic division of a primary oocyte
produces two unequal daughter cells. Each
Follicular cell
daughter cell has the haploid number of
chromosomes (23). The large cell, which Stroma
receives most of the cytoplasm, is called the
secondary oocyte, and the smaller cell is
known as the first polar body.
• The secondary oocyte immediately enters the
second meiotic cell division. Ovulation takes Zona pellucida
place while the oocyte is in metaphase. The
secondary oocyte remains arrested in
metaphase till fertilization occurs. Follicular cells
become columnar.
• The second meiotic division is completed
only if fertilization occurs. This division Basement membrane
results in two unequal daughter cells. The
smaller daughter cell is called the second
polar body. The first polar body may also Figs 2.9 and 2.10 Early stages in the formation of
divide during the second meiotic division. ovarian follicles.
Human Embryology

stage of development are called primordial


follicles.
Follicular cells form 3. A h o m o g e n e o u s m e m b r a n e , the zona
membrana granulosa.
pellucida, appears between the follicular
cells and the oocyte (Fig. 2.10).
4. The follicular cells proliferate to fonn several
layers. T h e s e c o n s t i t u t e the membrana
granulosa (Fig. 2.11). The cells may now he
called granulosa cells.
5. A cavity (or antrum) appears within the
membrana granulosa. With its appearance
Fig. 2.11 Formation of membrana granulosa. a follicle is formed (follicle = small sac}
(Fig. 2.12).
6. The cavity of the follicle rapidly increases
in size. As a result, the wall of the follicle
(formed by the granulosa cells) becomes
r e l a t i v e l y t h i n . T h e o o c y t e n o w lies
eccentrically in the follicle, surrounded by
some granulosa cells that are given the name
cumulus oopboricus (or cumulus ovaricus).
T h e cells that attach it to the wall of the
follicle are given the name discus proligerus
Fig. 2.12 Formation of follicular cavity. (Fig. 2.13).
7. As the follicle e x p a n d s , the stromal cells
Theca externa surrounding the membrana granulosa
become condensed to form a covering called
the theca interna (theca = cover). The cells
\ ——Theca
' , : - \ •,. interna of the theca interna later secrete a hormone
called oestrogen; and they are then called
\ Membrana
"1 granulosa the cells of the thecal gland (Fig. 2.13).
8. Outside the theca interna some fibrous tissue

I / becomes condensed to form another covering


for the follicle called the theca externa
nm •V :' Cumulus (Fig. 2.13). The ovarian follicle is now fully
aT*Jp _ .-. ••;•/ oopnoncus formed.
o / •" Oocyte
Inter-dependence of Oocyte and
7*vl Zona pellucida Follicular Cells
Discus proligerus Borh the oocyte and surrounding follicular cells
Fig. 2.13 Fully formed ovarian follicle.
are dependent on each other for further develop-
ment. The follicular cells secrete meiotic inhibi-
flattened cells ultimately form the ovarian tory factors, which prevent primary oocytes
follicle and are, therefore, called follicular from maturing heyond the prophase of the first
cells. meiotic division. This effect can last for many
2. T h e f l a t t e n e d f o l l i c u l a r cells b e c o m e years. These inhibitory factors are transmitted
columnar (Fig. 2.10). Follicles up to this to the oocyte via gap junctions present between
Spermatogenesis and Oogenesis

Perivitelline space Zona peiiucida

\ /

Microvillus
of oocyte

Zona peiiucida

Fig. 2.14 Diagrams to show the intimate relationship of oocyte and follicular cells. Gap junctions are
present between microvilli of the t w o .

Ruptured follicle
Stroma and theca
become very thin here. Ovum

Fig. 2.15 Relationship of a growing ovarian follicle to the ovary.

microvilli of oocytes and of follicular cells. These compared to the thickness of the cortex of the ovary
junctions lie in the zona peiiucida (Fig. 2.14). (Fig. 2.15A). As it enlarges, it becomes so big that
Follicular cells are also responsible for it not only reaches the surface of the ovary, but
growth, metabolism and maturation of oocytes. also forms a bulging in this situation. Ultimately,
Conversely, oocytes are responsible for prolife- the follicle ruptures and the ovum is shed from the
ration and differentiation of follicular cells. ovary (Fig. 2.15D).
Factors produced in the oocyte help in formation Just before ovulation the follicle may have a
and maturation of Graafian follicles. diameter of 15 mm. The stroma and theca on this
side of the follicle become very thin. An avascular
area {stigma) appears over the most convex point
Ovulation
of the follicle. At the same time, the cells of the
The shedding of the ovum from the ovary is called cumulus oophoricus become loosened by
ovulation. The ovarian follicle is at first very small accumulation of intercellular fluid between them.
Human Embryology

The followingfactors may lead to ovulation: Structure of the Ovum


1. Ovulation occurs due to high concentration The ovum that is shed from the ovary is not fully
of LH (Luteinizing Hormone) in blood just mature. It is in fact a secondary oocyte that is
before ovulation (see Chapter 3). undergoing division to shed off the second polar
2. A high c o n c e n t r a t i o n of L H leads t o body (Fig. 2.8).
i n c r e a s e d a c t i v i t y of t h e e n z y m e At this stage the ovum has the appearance, as
collagenase. T h i s e n z y m e digests t h e illustrated in Fig. 2.16. N o t e that it is surrounded
collagen fibres surrounding the follicle. by the zona pellucida. Some cells of the corona
3. Increase in concentration of prostaglandins radiata can be seen sticking to the outside of the
causes contraction of smooth muscle in the zona pellucida. N o nucleus is seen, as the nuclear
wall of the ovary. membrane has dissolved for the second meiotic
4. T h e increased p r e s s u r e of fluid in the division. A spindle is, however, present. Between
follicular cavity is also a significant factor the cell membrane (or vitelline membrane) a n d
for ovulation to occur. the zona pellucida, a distinct perivitelline space is
5. However, the enzymatic digestion of the seen. The first polar body lies in this space. Note
follicular wall seems to be the main factor that the ovum is a very large cell and measures
responsible for ovulation. more than 100 um in diameter. In contrast, most
other cells of the body measure less than 10 um.
(One um is one thousandth of a
First polar body millimetre).

Cells of
Spindle for second corona radiata
meiotic division Fate of the Ovum
Let us see what happens to the
ovum that is shed from the ovary.
You already k n o w that the ovary
is c l o s e l y e m b r a c e d by t h e
fimbriated end of the uterine tube.
T h e o v u m is, therefore, easily
carried into the tube, partly by
the follicular fluid discharged
Fig. 2.16 Structure of o v u m at the time of ovulation.
from the follicle and partly by the
activity of ciliated cells lining the
tube. T h e o v u m slowly travels
t h r o u g h the t u b e t o w a r d s the
uterus, taking three to four days
t o d o so. If sexual intercourse
takes place at about this time, the
s p e r m a t o z o a d e p o s i t e d in the
vagina swim into the uterus and
into the uterine tube. One of these
s p e r m a t o z o a m a y fertilize the
Granulosa cells in Wall of follicle collapses Cells hypertrophy o v u m . If t h i s h a p p e n s , t h e
wall of follicle and becomes folded to form corpus luteum fertilized ovum begins to develop
into an embrvo. It travels to the
Fig. 2.17 Stages in the formation of the corpus luteum.
Spermatogenesis and Oogenesis

Small cells in wall of


follicle soon after its rupture

- Lutein pigment

Considerably enlarged,
cells of corpus luteum

Fig. 2.18 Transformation of follicular cells (A) to luteal cells (B).

uterus and gets implanted in its wall. On the other supplied with blood vessels for this purpose.
h a n d , if the o v u m (secondary oocyte) is not T h e o v a r i a n follicle itself has no blood
fertilized it dies in 12 to 24 hours. It passes through vessels, bur the surrounding theca interna is
the uterus into the vagina and is discharged. full of them. When the corpus luteum is
forming, blood vessels form the theca interna
invade it and provide it with a rich supply
C o r p u s Luteum
of blood.
T h e corpus luteum is an important structure. Its
m a i n f u n c t i o n is t o s e c r e t e t h e hormone T h e s u b s e q u e n t fate of the c o r p u s luteum
progesterone, but it secretes some oestrogen also. depends on whether the ovum is fertilized or not.
The corpus luteum is derived from the ovarian
follicle, after the latter has ruptured to shed the (a) If the ovum is not fertilized, the corpus luteum
ovum, as follows (Fig. 2.17): persists for about 14 days. During this period
it secretes progesterone. It remains relatively
1. When the follicle ruptures, its wall collapses small and is called the corpus luteum of
and becomes folded. menstruation. At the end of its functional
2. At this stage, the follicular cells are small life, it degenerates a n d forms a mass of
and rounded (Fig. 2.18A). They now rapidly fibrous tissue called the corpus albicans ( =
enlarge. As they increase in size their walls white body) (Fig. 2.19).
press against those of neighbouring cells so (b) If the o v u m is fertilized a n d p r e g n a n c y
that the cells acquire a polyhedral shape results, the corpus luteum persists for three
(Fig. 2.18B). Their cytoplasm becomes filled to four months. This is larger than the corpus
with a yellow pigment called lutein. They luteum of menstruation, and is called the
are now called luteal cells. The presence of corpus luteum of pregnancy.
this yellow pigment gives the structure a
yellow colour and that is why it is called the The corpus luteum of pregnancy may occupy
corpus luteum (= yellow body). Some cells one-third to half the total volume of the ovary.
of t h e t h e c a i n t e r n a a l s o e n l a r g e a n d The progesterone secreted by it is essential for the
contribute to the corpus luteum. maintenance of pregnancy in the first few months.
3. We have seen that the corpus luteum secretes After the fourth m o n t h , the corpus luteum is no
progesterone. This secretion has to be poured longer needed, as the placenta begins to secrete
into the blood like secretions of endocrine progesterone. Degeneration of the corpus luteum
g l a n d s . All e n d o c r i n e g l a n d s are richly in the early months of pregnancy is prevented by
Human Embryology

Primordial follicles

Follicles that do not


rupture degenerate;
the ovum and granulosa
cells dissappear.

One follicle matures


and sheds an ovum.

The cells of theca interna


persist as the 'Interstitial
gland'.

The ruptured follicle


becomes a corpus
luteum.

Corpus luteum
degenerates and Interstitial gland
becomes a corpus degenerates and
albicans. becomes a corpus albicans.

Fig. 2.19 Fate of ovarian follicles.

human chorionic gonadotropin (hCG) secreted by these follicles do not persist into the next ovarian
the trophoblast cells of the developing embryo. cycle, but undergo degeneration. The ovum and
T h e series of c h a n g e s t h a t begin with the granulosa cells of each follicle disappear. The cells
formation of an ovarian follicle and end with the of the theca interna, however, proliferate to form
degeneration of the corpus luteum constitute what the interstitial glands, also called the corpora
is called an ovarian cycle. An ovarian cycle has atretica (singular = corpus atreticum). These glands
an average duration of 28 days, with ovulation are believed to secrete oestrogens. After a period
occurring at mid-cycle, i.e. on the 14th day. of activity, each gland becomes a mass of scar
tissue indistinguishable from the corpus albicans
formed from the corpus luteum.
Fate of Ovarian Follicles
We have seen t h a t in each o v a r i a n cycle o n e
Ovarian Cycle and Hormones
follicle reaches maturity, sheds an o v u m , and
becomes a c o r p u s luteum. At the same time, The changes taking place during the ovarian cycle
several other follicles also begin to develop, but are greatly influenced by certain hormones pro-
do not reach maturity (Fig. 2.19). It is interesting duced by the hypophysis cerebri (see Chapter 3).
to note that, contrary to what one might expect, The hormones produced by the theca interna and
Spermatogenesis and Oogenesis

large nucleus or two nuclei. Two


Differences between male and female gametes
oocytes may be seen in one follicle.
Male gamete (sperm) Female gamete (ovum)
2. Chromosomal Abnormalities
Sperm ts small but long Ovum is a massive cell about
(about 60 um in length and 120 um in diameter. The gametes may be abnormal in
about 2 um in width).
chromosomal content as follows:
It is highly motife. It is immotile.

There is very little cytoplasm. The oocyte contains a large (a) During the first meiotic division,
This gives it high motility. amount of cytoplasm. the two chromosomes of a pair,
Shape is adapted for motility. Shape adapted to provide ample instead of separating at anaphase,
storage of nutrition for the embryo may both go to the same pole.
formed after fertilization.
(This is called non-disjunction.)
Spermatozoa are ot two All ova have (22+X) chromosomes.
chromosomal types The resulting gamete then has 24
(22+X)and(22+Y). chromosomes instead of the
normal 23 (Fig. 2.21).
Fig. 2.20 Differences between male and female gametes At fertilization by this gamete, the
zygote will, therefore, have 47
by the corpus luteum in turn influence other parts chromosomes; there being three identical
of the female reproductive system (notably the chromosomes instead of one of the normal
uterus), resulting in a cycle of changes referred to pairs. This is called trisomy. Depending
as the uterine or menstrual cycle. upon the particular chromosomes involved,
various abnormalities are produced.
Reproductive Period Trisomy of chromosome 21 results in a
condition called mongolism or Down's
In an individual the formation of gametes takes syndrome. In this condition the child has a
place only during the reproductive period which broad face, obliquely placed palpebral
begins at the age of puberty (10 to 14 years). In fissures, epicanthus, a furrowed lower lip,
women it ends between the ages of 45 and 50 years, and broad hands with a single transverse
but in men it may continue till the age of 60 years crease. Usually the patients are mentally
or more. retarded, and have anomalies of the heart.
The presence of an extra X or Y
Viability of Gametes chromosome can give rise to various
An ovum usually degenerates 24 hours after syndromes associated with abnormal
ovulation. However, at the most it may survive genital development, mental retardation
for two days. Similarly, sperms usually degenerate and abnormal growth. Some of these are:
48 hours after ejaculation, but may survive up to XXX (abnormal female); XXY (Klinefelter's
four days in female genital tract. syndrome: abnormal male); XYY
(abnormal male). In Klinefelter's syndrome
the subject is a male (because of the
ABNORMALITIES IN FORMATION presence of a Y chromosome). However,
OF GAMETES the testes are poorly developed leading to
1. Abnormalities of Form sterility and gynecomastia.

Spermatozoa may be too large (giant) or too Patients with XXX chromosomes show
small (dwarf). The head, body or tail may be two masses of sex chromatin in their cells
duplicated. The ovum may have an unusually and are sometimes referred to as 'super
Human Embryology

FIRST
MEIOTIC
DIVISION

SECOND
MEIOTIC
DIVISION

GAMETES

FERTILIZATION
with normal
gametes
containing
23 chromosomes

ZYGOTE

Trisomy Monosomy

Fig. 2.21 Effects of non-disjunction during the first meiotic division.

females'. However, there is nothing 'super' and menstruation is scanty. Mental


about them. In fact, their bodies show poor retardation is usual.
sexual development (i.e. they are infantile) (b) When both chromosomes of a pair go to
Spermatogenesis and Oogenesis

one gamete (as described above), the other represented by a single chromosome. This
gamete resulting from the division has only is called monosomy (Fig. 2.21).
22 chromosomes (instead of the normal 23); The best-known example of this is a
and at fertilization the zygote has only 45 female with only one X chromosome
chromosomes. Hence one pair is (Turner's syndrome). In this syndrome the

Fig. 2.22 Effects of non-disjunction during the second meiotic division.


Human Embryology

subject is always a female (because of and some of the genes are duplicated.
absence of a Y chromosome). There is The other chromosome will be shorter
agenesis of ovaries. Associated deformities than normal, some genes being
include mental retardation, skeletal missing.
abnormalities, and folds of skin on the sides (iv) A piece separating from a chromosome
of the neck (webbed neck). may get inverted before joining
Non-disjunction may also occur during the opposite chromosome (inversion).
second meiotic division as shown in Although the same genes are present,
Fig. 2.22. their sequence is disturbed.
(c) Such anomalies may affect more than one
(f) We have seen that during cell division the
pair of chromosomes. Alternatively, one
centromere splits longitudinally so that
pair may be represented by more than three each chromatid becomes a separate
chromosomes. When this happens with the chromosome. Sometimes the centromere
sex chromosomes, individuals with the splits transversely producing two dissimilar
constitution XXXY, XXXXY, XXYY, or chromosomes. One chromosome is made
XXXX may be produced. up of the short arms of both chromatids,
(d) Sometimes a gamete may have the diploid while the other is made up of the long arms.
number of chromosomes so that the zygote These are called isochromosomes.
will have 46 + 2*3 (i.e. 69) chromosomes.
Chromosomal errors of the type
This is called triploidy. Higher multiples
described above may also occur during
of 23 may also be seen. Such fetuses are segmentation of the ovum. This results in
generally born dead. a fetus having a mixture of cells with
(e) Abnormalities in the process of crossing normal and abnormal chromosomes. This
over can result in a number of chromosomal is called mosaicism. Such individuals may
abnormalities as follows: also show various abnormalities.
(i) Part of a chromosome may get attached
to a chromosome of a different pair 3, Gene Abnormalities (Gene Mutations)
(translocation). Genes are responsible for normal embryological
(ii) Part of a chromosome may be lost development. A change in the structure of a gene
(deletion). may occur at the time of gametogenesis. This
(iii) The two chromosomes of a pair may may give rise to birth defects. The change in the
break at unequal distances. When each structure or function of a gene is called gene
piece joins the opposite chromosome, mutation. At present many birth defects are
one chromosome is longer than normal known which are caused by gene mutations.
I3
Chapter

The Menstrual Cycle

HIGHLIGHTS
• The term menstrual cycle is applied to cyclical
changes that occur in the endometrium every
month. The most obvious feature is a monthly
flow of blood (menstruation).
• T h e m e n s t r u a l cycle is divided into the
following phases: postmenstrual, proliferative,
secretory, menstrual (Fig. 3.3).
• The menstrual cycle is also divided into the
follicular phase (in which changes are produced
mainly by oestrogens) and the luteal phase (in
which effects of progesterone predominate).
Both phases are of roughly equal duration.
• The main changes in the endometrium are:
(a) increase in thickness, (b) growth of uterine
glands, (c) changes in epithelial cells lining
the glands and (d) increase in thickness and
fluid c o n t e n t of t h e e n d o m e t r i a l s t r o m a The period of a woman's life in which she can
(Figs 3.4, 3.5). bear children is called the reproductive period.
• Just before onset of menstruation the blood T h e m o s t o b v i o u s feature of this period is a
supply to superficial parts of the endometrium monthly flow of blood from the uterus that is
is cut off (Fig. 3.6). This part is shed off and referred to as menstruation (or menses). The onset
rhere is bleeding. of menstruation (menarche) takes place at about
• The menstrual cycle is influenced by oestrogens, 12 years of age. Menstruation ceases to occur at
by progesterone, by the follicle stimulating about 45 years of age, and this is referred to as
h o r m o n e ( F S H ) , a n d by the l u t e i n i z i n g menopause.
hormone (LH). T h e m o n t h l y m e n s t r u a t i o n is the external
manifestation of a series of cyclic changes taking
place in the uterus. These changes constitute the
menstrual cycle. Simultaneously, cyclic changes
also take place in the ovaries, and these constitute
the ovarian cycle. The most important event in
the ovarian cycle is ovulation (Chapter 2).
Human Embryology

bleeding bleeding
\
Days
r One Menstrual Cycle -

Fig. 3.1 Diagram illustrating the definition of a menstrual cycle.

To understand the menstrual cycle it is (b) The stroma fills the interval between surface
necessary to know the structure of the uterus. The epithelium and myometrium. It contains
wall of the uterus is made up of three layers. numerous simple tubular glands (uterine
glands).
1. The outermost layer or perimetrium is made
(c) The arteries that supply the endometrium
up of peritoneum.
tend to run vertically towards the surface.
2. The main thickness of the wall is made up
Some of these run spirally and supply the
of smooth muscle. This is the myometrium.
whole thickness of the endometrium, while
3. The innermost layer (corresponding to
others that remain straight are confined to
mucous membrane) is called the endo-
metrium. It is this layer which undergoes the basal part.
changes during the menstrual cycle.
The constituents of the endometrium are as follows PHASES OF THE MENSTRUAL CYCLE
(tig- 3.2).
The menstrual cycle is usually divided into the
(a) The surface of the endometrium is covered following phases, on the basis of changes raking
by a lining epithelium. place in the uterine endometrium (Fig. 3.3):

Lining epithelium

Straight artery

Myometrium Arterial plexus


I
Fig. 3.2 Components of the uterine endometrium. These undergo changes
during each menstrual cycle.
The Menstrual Cycle

5 -
Cycle beams ? ''^fl
;V8
s\V
here.

28^W
O
if)

<
Q_ 26 1
•?° / %U11
<
LU
Corpus luteum
^U Secretory I12
Proliferative n I
begins to 24 \ U 13
degenerate. Jr14 m
23 N ^
^ ^ 1 5 Ovulation
21^^C= — ^ 1 6
20 17

Fig. 3.3 Phases of the menstrual cycle.

1. Postmenstrual into the phases mentioned above is, however,


2. Proliferative arbitrary. T h e changes are really continuous, and
3. Secretory or premenstrual may be summarized as follows:
4. Menstrual
1. The endometrium progressively increases in
The changes during rhe postmenstrual phase
thickness (Fig. 3.4}. In the postmenstrual
and during most of the proliferative phase take
p h a s e it is 0.5 to 1 m m t h i c k ; in t h e
place under the action of oestrogens produced by
the developing follicles in the ovary.
Hence this period is referred to as the
follicular phase of the menstrual cycle. Secretory
The follicular phase constitutes the first-
half of the menstrual cycle. Following
ovulation, the corpus luteum is formed
and starts secreting progesterone. During
the second-half of the menstrual cycle,
this h o r m o n e (along with oestrogens) m
p r o d u c e s s t r i k i n g c h a n g e s in t h e '-
e n d o m e t r i u m . As these changes take
place under the influence of the corpus
luteum, this half of the menstrua! cycle
Postmenstrual i
7

IT
is called the luteal phase. Just before the CM

o n s e t of t h e n e x t b l e e d i n g , t h e r e is
1
lowering of levels of both progesterone
and oestrogens, and it is believed that
this 'withdrawal' leads to the onset of
menstrual bleeding.
Fig. 3.4 Uterine glands at various phases of the
The division of the menstrual cycle menstrual cycle.
Human Embryology

proliferative phase it is 2 to 3 mm thick; surrounding the bodies of the uterine glands


a n d in the secretory phase its thickness becomes spongy. The deepest part of the
reaches 5 to 7 mm. stroma also remains compact. The stroma
The uterine glands grow in length. At first can, therefore, be divided into the following
they are straight (Fig. 3.4A), but gradually three layers (Fig. 3.6A).
become convoluted (Fig. 3.4B). Because of
(a) Stratum compactum
these convolutions, the glands acquire a
(b) Stratum spongiosum
' s a w - t o o t h e d 1 a p p e a r a n c e when seen in
(c) Stratum basale
l o n g i t u d i n a l s e c t i o n . T h e g l a n d s also
increase in diameter (Fig. 3.4C). The most During the secretory phase, these layers
basal p a r t s of uterine g l a n d s , however, become better defined. The endometrium
remain tubular and do not undergo these becomes soft and oedematous, because of
changes. the fluid secreted by the uterine glands.
D u r i n g the p o s t m e n s t r u a l phase the 5. The arteries of the endometrium are small
epithelium lining the glands is cuboidal to begin with. They grow in length during
(Fig. 3.5A). During the proliferative stage the proliferative phase. During the secretory
it becomes columnar (Fig. 3.5B). Glycogen phase, the arteries supplying the superficial
accumulates in the basal portion of the two-thirds of the endometrium become very
epithelial cell, pushing the nucleus nearer tortuous, and are called spiral arteries. The
the lumen (Fig. 3.5C). During the secretory a r t e r i e s t o t h e b a s a l t h i r d of t h e
phase the apical part of the cell is shed off endometrium (which does not participate in
as part of the secretion. The cell again the changes associated with the menstrual
becomes cubical, but the edge towards the cycle) remain straight and short.
lumen becomes irregular (Fig. 3.5D).
During the postmenstrual phase, the cells Towards the end of the secretory phase the
of the stroma are uniformly distributed and endometrium is thick, soft, and richly supplied
are compactly arranged (Fig. 3.2). As the with blood. The secretory activity of the uterine
endometrium increases in thickness (during glands not only makes the endometrium soft, but
the proliferative phase), the superficial part also provides nutrition to the e m b r y o . These
of the stroma remains compact, but the part changes are, therefore, an obvious preparation for

% B ^

Postmenstrual Secretory

Fig. 3.5 Changes in the epithelium of uterine glands during a menstrual cycle.
The Menstrual Cycle

Lining epithelium

. These layers
are shed off.

Lining epithelium is
reformed by proliferation
of cells lining the basal
parts of the uterine glands.

in
Secretory Endometrium Endometrium
endometrium at end of during the post-
menstruation menstrual phase

Fig. 3.6 Shedding off, and regeneration, of uterine endometrium, during


a menstrual cycle
SC = stratum c o m p a c t u m ;
SS = stratum spongiosum;
SB = stratum basale.

providing a suitable environment for the fertilized The mechanism for onset of menstrual bleeding
ovum, when it reaches the uterus. In the absence is as follows. A few hours before the onset of
of pregnancy, however, these measures are menstrual bleeding the spiral arteries get
abortive: the superficial parts of the thickened constricted so that blood supply to superficial
endometrium (stratum compactum and stratum parts of the endometrium is cut off. This
spongiosum) are shed off (Fig. 3.6B), and this is ischaemia leads to degeneration of the
accompanied by menstrual bleeding. endometrium and also damages the walls of
Menstrual bleeding causes the endometrium to the blood vessels themselves. Subsequently
be shed off bit by bit, and the blood along with when the arteries relax and blood again flows
shreds of endometrium flows out through the into the endometrium, it leaks out through the
vagina. At the end of menstruation, the damaged blood vessels. This leaking blood is
endometrium that remains is only 0.5 mm thick. responsible for gradual shedding of
It consists of the stratum basale along with the endometrium.
basal portions of the uterine glands (Fig. 3.6B).
The epithelium of these glands rapidly proliferates
and reforms the epithelial lining (Fig. 3.6C). Time of Ovulation in
Relation to Menstruation
The endometrial changes associated with the
menstrual cycle are confined to the body of the In a 28-day menstrual cycle, ovulation takes place
uterus. The cervical mucosa is not affected. at about the middle of the cycle (Fig. 3.3). The
Human Embryology

Fig. 3.7 Graph showing the morning temperature of a woman, on


various days of the menstrual cycle. There is a fall in
temperature at about the time of ovulation, followed by a rise.

period between ovulation and the next menstrual occur only if intercourse takes place during a
bleeding is constant at about 14 days, but the time period between four days before ovulation to
of ovulation does not have a constant relationship two days after ovulation. The remaining days
with the preceding menstruation. This is so because have been regarded as safe period as far as
the length of the menstrual cycle in women may prevention of pregnancy is concerned. This
vary from one m o n t h to another. Hence, it is forms the basis of the so-called rhythm-method
difficult to predict the date of the next ovulation of family planning.
from the date of menstruation unless the woman
has very regular menstrual periods. B. Where pregnancy is desired
There are many methods of finding out the Knowledge regarding the time of ovulation is
exact time of ovulation, but the one commonly also of importance in cases of sterility (difficulty
used is the temperature method. In this technique, in having children), where the couple can be
the woman's body temperature is recorded every advised to have intercourse during the days most
morning. When these temperatures are plotted on favourable for conception.
a g r a p h , we get a curve like t h a t s h o w n in
Fig. 3.7. The temperature is low during actual
Correlation between Ovarian and
menstruation. Subsequently it rises. At about the Uterine Cycles
middle of the cycle, there is a sudden fall in The ovarian and uterine cycles run parallel to
temperature followed by a rise. This rise is believed each other. Both are of 28 days duration. The
to indicate that ovulation has occurred. uterine cycle is dependent on the ovarian cycle.
The uterine endometrium shows cyclic changes,
CLINICAL CORRELATION which are dependent on the hormones secreted
by developing ovarian follicles a n d c o r p u s
Importance of Determining the Time of luteum of the ovary (Fig. 3.8).
Ovulation and 'Safe Period'
A. Where pregnancy is not desired HORMONAL CONTROL OF OVARIAN
AND UTERINE CYCLES
After ovulation, the ovum is viable {i.e..it can
be fertilized) for n o t m o r e t h a n t w o d a y s . These cycles are under the control of various hormones,
Spermatozoa introduced into the vagina die which can be briefly summarized here (Fig. 3.9).
within four days. Therefore, fertilization can The hypothalamus acts as a major centre for
The Menstrual Cycle

OVARIAN
CYCLE

Developing Mature Corpus Degenerating


follicles follicles luteum corpus luteum

FOLLICULAR PHASE LUTEAL PHASE


Menstrual Postmenstrual &
Secretory phase
phase proliferative phase

Fig. 3.8 Diagram showing correlation between ovarian and uterine cycles.

t h e c o n t r o l of r e p r o d u c t i o n . It secretes the pituitary (LH surge) about 24 to 36 hours before


Gorutdotropin-releasing hormones (GnRH), which ovulation (Fig. 3.10).
in turn control the secretion of gonadotrophs The LH surge leads to o v u l a t i o n ; and the
hormones by t h e a n t e r i o r p i t u i t a r y g l a n d Graafian follicle is transformed into the corpus
(adenohypophysis). luteum.
There are two g o n a d o t r o p h s hormones. They The LH stimulates the secretion of progesterone
are the follicle stimulating hormone (FSH) and by the corpus luteum. Though the secretion of
the luteinizing hormone (LH). progesterone predominates, some oestrogen is also
In the first-half of the menstrual cycle the C n R H produced. The combined action of oestrogen and
acts on the anterior pituitary to release FSH. The progesterone stimulates the endometrial glands to
FSH acts on the ovary and stimulates the formation secrete glycogen rich mucoid material (Fig. 3.9).
and maturation of ovarian follicles (Fig. 3.9). If fertilization does not occur the granulosa cells
T h e m a t u r i n g o v a r i a n follicles n o w s t a r t produce the protein inhibin, which acts on the
secreting oestrogens. The repair and proliferation anterior pituitary and inhibits the secretion of
of endometrium takes place under the influence of gonadotrophins. This leads to regression of the
oestrogens. The endometrial stroma progressively corpus luteum.
thickens, the glands in it elongate and the spiral Due to the regression of the corpus luteum there
a r t e r i e s begin t o g r o w t o w a r d s t h e surface is a fall in the b l o o d level of o e s t r o g e n a n d
epithelium. progesterone. The withdrawal of these hormones
The level of oestrogen rises to a peak about causes the endometrium to regress and triggers
two days before ovulation. This leads to sudden the onset of menstruation.
increase in the level of L H secreted by the anterior If fertilization occurs the corpus luteum does
Human Embryology

HYPOTHALAMUS
produces releasing factors

f
ADENOHYPOPHYSfS
produces

FSH LH

1
Formation & maturation Ovulation Formation of
of ovarian follicles corpus luteum

OVARY

0es,r
°9en Progesterone
Produced produced

Repair & Proliferative phase Secretory phase

ENDOMETRIUM

Fig. 3.9 H o r m o n a l control of ovarian and uterine cycles.

not regress. It continues to secrete progesterone


form of synthetic compounds). Better results are
and oestrogen. The secretory phase of endometrium
obtained when a small amount of oestrogen is
continues and menstruation does not occur.
also given.
In the most common variety of pill
CLINICAL CORRELATION (distributed by government agencies in India)
Use of Hormones for Contraception the progestin is norethisterone acetate (1 mg);
and the oestrogen is in the form of oestradiol
Ovulation in a woman (and by corollary, (50 ug). The pills are distributed in packets,
pregnancy) can be prevented by administering each packet containing 28 pills out of which
contraceptive pills. The most important 21 pills contain these hormones, and 7 pills do
ingredients of such pills are progestins (in the not (for use in the last 7 days). The use of pills
The Menstrual Cycle

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28

Concentration maximum
two days before ovulation

PROGESTERONE
Level increases to maximum in the
secretory phase under influence ot LH.

Fig. 3.10 Concentration of the hormones FSH, L H , estrogen and progesterone during a normal menstrual
cycle. O v u l a t i o n occurs because of a LH surge just before ovulation.

is started 5 days after onset of menstruation. because the progesterone in the pill prevents
They are taken continuously without any break the secretion of FSH and LH by the pituitary.
as long as contraception is desired. N o r m a l This interferes with the maturation of follicles
menstruation occurs during the 7 days in which and ovulation.
pills without hormones are being taken. If the Stoppage of pills reduces levels of these
pills are taken regularly there is a regular hormones in blood. It is this withdrawal that
menstrual cycle of 28 days duration. leads to menstrual bleeding. Contraceptive pills
Presence of progesterone in the preovulatory have almost 100 per cent success in suppressing
phase prevents occurrence of ovulation. This is maturation of follicles and ovulation.
Chapter

4 Formation of Germ Layers

a somatopleuric layer (in c o n t a c t w i t h


HIGHLIGHTS
trophoblast) and a splanchnopleuric layer (in
• Fertilization of the ovum takes place in the contact with yolk sac) (Fig. 4.7B).
ampulla of the uterine tube (Fig. 4.1). The
The trophoblast and underlying somatopleuric
fertilized ovum is a large cell. It undergoes a
mesoderm form a membrane called the chorion.
series of divisions (clevage) (Fig. 4.4).
The cells forming the wall of the amniotic
• When there are 16 cells, the ovum is called a
morula (Fig. 4.4). It has an inner cell mass cavity form the amnion.
c o v e r e d by an o u t e r layer of cells, the T h e a m n i o t i c cavity is n o w a t t a c h e d to
trophoblast (Fig. 4.5A). trophoblast by some mesoderm into which the
• Fluid partially separates the inner cell mass extra-embryonic coelom has not extended. This
from trophoblast. The morula now becomes a mesoderm forms the connecting stalk.
blastocyst (Fig. 4.5C). If we view t h e e m b r y o n i c disc from the
• The cells of the inner cell mass multiply, and ectodermal side we see that near one edge it
are rearranged to form an embryonic disc has a rounded area called the prochordal plate
having two germ layers. These layers are the (Fig. 4.9). Here ectoderm and endoderm are
ectoderm and endoderm. Later, a third germ not separated by mesoderm.
layer, the mesoderm, forms between ectoderm An elevation, the primitive streak, is also seen
and endoderm. on the embryonic disc (Fig. 4.10A}. A line
• A cavity appears on the ectodermal side of the drawn through the prochordal plate and the
disc. This is the amniotic cavity {Fig. 4.6C). primitive streak divides the embryonic disc into
Another cavity appears on the endodermal right and left halves.
side. This is the yolk sac. Cells multiplying in the primitive streak move
• At first the walls of the amniotic cavity and yolk i n t o the interval b e t w e e n e c t o d e r m a n d
sac are in contact with trophoblast (Fig. 4.6D). endoderm and form the mesoderm (third germ
They are soon separated from the latter by layer) (Fig. 4.12).
extra-embryonic mesoderm (Fig. 4.7A).
Caudal to the primitive disc we see a round
• A cavity, the extra-embryonic coelom appears area called the cloacal membrane. It is made
and splits the extra-embryonic mesoderm into up only of ectoderm and endoderm (Fig. 4.12).
Formation ot Germ Layers

pronucleus. Soon thereafter, the pronuclei lose


FERTILIZATION
their nuclear membranes. The 2 3 chromosomes
In Chapter 2 we have seen that while the ovarian of the female pronucleus and 2 3 of the male
follicle is g r o w i n g , t h e o o g o n i u m w i t h i n it pronucleus get mixed u p and form 2 3 pairs. These
undergoes maturation. The oogonium enlarges to 46 chromosomes undergo changes like those in a
form a p r i m a r y o o c y t e . T h e p r i m a r y oocyte typical mitotic division leading to the formation
undergoes the first meiotic division to shed off the of an embryo having two cells. Note that, strictly
first polar body and becomes a secondary oocyte speaking, there is no one-cell stage of the embryo.
(Fig. 4.2A). At the time of ovulation, the second
meiotic division is in progress and a spindle has Some details on t h e b i o c h e m i c a l c h a n g e s
formed for separation of the second polar body occurring during fertilization are worth noting.
(Fig. 4.2B). At this stage the ' o v u m ' enters the These are as follows.
mfundibulum of the uterine tube and passes into
1. T h e glycoprotein of the zona pellucida is
the ampulla (Fig. 4.1).
responsible for induction of the acrosomal
Fertilization of the ovum occurs in the ampulla r e a c t i o n ( C h a p t e r 2 ) . T h e release of
of the u t e r i n e t u b e . O u t of a few h u n d r e d a c r o s o m a l enzymes helps the sperm to
capacitated sperms, which surround the ovum, only penetrate the zona.
one pierces the zona pellucida and enters the ovum. 2. When a spermatozoon comes in contact
As soon as the spermatozoon enters the ovum the with the oocyte, plasma membranes of the
s e c o n d m e i o t i c d i v i s i o n ( w h i c h w a s so far two cells fuse. This, probably occurs at
incomplete! is completed, and the second polar receptor sites that are specific for a species.
b o d y is e x t r u d e d . T h e n u c l e u s of the o v u m Both the head and tail of the spermatozoon
becomes the female pronucleus. The head of the (excluding the plasma membrane) enter
spermatozoon (which it will be remembered is the cytoplasm of the ovum.
formed from the nucleus) separates from the middle 3. Alterations taking place in the plasma
piece and tail, and transforms itself into the male

Ampulla (6 cm)
Uterine part (1 cm) Isthmus (3 cm)

Fertilization
takes place here

Fig. 4.1 Path taken by the sperm (pink), and ovum (blue), for fertilization.
Human Embryology

- Zona peilucida
. Spindle of second
meiotic division
• Secondary oocyte

- First polar body " First polar body

Nucleus ot
Female pronucleus
mature ovum (From nucleus of ovum)

Vitelline membrane
Male pronucleus
(From head of sperm)

Spermatozoon
entering ovum

Fig. 4.2 Some stages in the maturation of the o v u m : (A) O v u m just before o v u l a t i o n ;
(B) O v u m at the t i m e of o v u l a t i o n ; (C) O v u m at the time of fertilization;
(D) O v u m just after fertilization.

membrane of the oocyte, and in the zona to formation of two cells, each having
peilucida, ensure that no other forty-six chromosomes (Fig. 4.3).
spermatozoon can enter the oocyte.
4. The zona peilucida is altered due ro release Male pronucleus
of l y s o s o m a l e n z y m e s by t h e p l a s m a
membrane of the oocyte. This process is
called the zona reaction.
5. As soon as a s p e r m a t o z o o n enters the Female pronucleus
ovum, the latter finishes its second meiotic
division and the second polar body is
formed.
6. Entry of the sperm leads to metabolic
Each chromosome
changes within the ovum that facilitate its duplicates itselt.
development into an embryo.
7. Each chromosome in the male and female
p r o n u c l e i is m a d e u p of o n l y o n e
Each daughter cell
c h r o m a t i d . Replication of D N A takes has 23 pairs of
place to form a second chromatid in each chromosomes - one of
each pair derived
c h r o m o s o m e . In the cell division t h a t from the spermatozoon
follows, each chromosome splits into two and the other from the ovum.
(as in mitosis). Meanwhile a spindle has
formed and one chromosome of each pair Fig. 4.3 Behaviour of chromosomes during
moves to each end of the spindle. This leads fertilization.
Formation of Germ Layers

8. From w h a t has been said above, it will be 22 + Y chromosomes. We speak of these as 'X-bear-
clear that as a result of fertilization ing", or 'Y-bearing', spermatozoa. An ovum can
be fertilized by cither type of spermatozoon. If the
fa) the diploid chromosome number (46)
sperm is X-bearing, the zygote has 44 + X + X
is restored; chromosomes and the offspring is a girl. If the
(b) determination of sex takes place; and sperm is Y-bearing the zygote has 44 + X + Y
(c) the fertilized ovum begins to divide chromosomes and the offspring is a boy.
into several cells (i.e. it undergoes
Thus the sex of a child is 'determined' at the
cleavage).
time of fertilization. It will now be clear that one
9. The important points to note at this stage chromosome of each of the 2 3 pairs is derived
are that from the mother and the other from the father.

(a) t h e t w o d a u g h t e r cells a r e still


surrounded by the zona pellucida; CLEAVAGE
(b) each daughter cell is much smaller
The two cells formed as described above undergo
than the ovum;
a series of divisions. One cell divides first so that
(c) as subsequent divisions occur, the cells
we have a '3-cell' stage of the embryo (Fig. 4.4R)
become smaller and smaller until they
followed by a '4-cell' stage (Fig. 4.4C), a '5-cell'
acquire the size of most cells of the
stage, etc. This process of subdivision of the ovum
body.
into smaller cells is called cleavage.
As cleavage proceeds rhe ovum comes to have
Test Tube Babies
16 cells. It now looks like a mulberry and is called
The so-called test tube babies are produced by the morula (Fig. 4.4D). It is still surrounded by
rhe Technique of in vitro fertilization (In vitro - rhe zona pcllucida. If we cut :i section across the
outside the body, as against in vivo - within morula, we see that it consists of an inner cell
the body). This technique is being increasingly mass that is completely surrounded by an outer
used in couples w h o are not able to achieve layer of cells. T h e cells of the outer layer will
fertilization in the normal way. later give rise to a structure called the trophoblast
G o n a d o t r o p i n s are a d m i n i s t e r e d t o the (Fig.4.5A).
woman to stimulate growth of follicles in the The inner cell mass gives rise to the embryo
ovary. Just before o v u l a t i o n , the o v u m is p r o p e r , a n d is, t h e r e f o r e , a l s o c a l l e d t h e
removed (using an aspirator) and is placed in a embryoblast. The cells of the trophoblast help to
suitable medium- Spermatozoa are added to provide nutrition to the embryo.
the m e d i u m . Fertilization and early Some fluid now passes into the morula from
development of the embryo take place in this the uterine cavity, and partially separates the cells
medium. The process is carefully monitored, of the inner cell mass from those of the trophoblast
and when the embryo is at the 8-cell stage it is (Fig. 4.5B). As the quantity of fluid increases, the
put inside the uterus. Successful implantation morula acquires the shape of a cyst. The cells of
takes place in about 20 per cent of such trials. the trophoblast become flattened, and the inner
cell mass comes to be attached to the inner side of
SEX DETERMINATION the trophoblast on one side only (Fig. 4 . 5 Q . The
morula has now become a blastocyst. The cavity
We know that all ova contain 22 + X chromo- of the blastocyst is called the blastocoele. The side
somes. However, we have seen that spermatozoa of the blastocyst to which the inner cell mass is
are of t w o types. Half of them have 22 + X attached is called the embryonic or animal pole,
chromosomes and the other half of them have while the opposite side is the abembryonic pole.
Human Embryology

can, therefore, invade and b u r r o w into


tissues with which they come in contact.
As the embryo travels down the uterine tube,
and the u p p e r m o s t p a r t of the uterine
cavity, it is prevented from 'sticking' to the
epithelium by the zona pellucida. During
this time it receives nutrition, partly from
the substances stored within the ovum (e.g.
yolk), and partly by diffusion from uterine
secretions. By the time a blastocyst is
formed, it is necessary for the embryo to
acquire additional sources of nutrition. This
is achieved when the blastocyst 'sticks' to
the uterine endometrium, and gets implanted
in it. However, before this can happen, it is
n e c e s s a r y for t h e z o n a p e l l u c i d a t o
disappear. The zona pellucida disappears
soon after the morula reaches the uterine
l u m e n . T h u s , the function of the zona
pellucida is to prevent implantation of the
blastocyst at an abnormal site.

Fig. 4.4 Some stages in segmentation of the fertilized T h e developing e m b r y o is genetically


o v u m ; (A) Two-cell stage; (B) Three-cell stage; different from the mother. This may evoke
(C) Four-cell stage; (D) M o r u l a . immunological reactions if embryonic and
maternal tissues come in contact. Presence
of zona pellucida (which lacks histocompa-
Function of the Zona Pellucida
tibility antigens), acts as a barrier that separates
The trophoblast has the property of being able to maternal tissues from the e m b r y o . After the
stick to the uterine (or other) epithelium and its d i s a p p e a r a n c e of z o n a p e l l u c i d a v a r i o u s
cells have the capacity to 'cat u p ' other cells. They immunosuppressive cytokinase and proteins are

Outer layer of cells


forming future trophoblast

Fig. 4.5 Formation of blastocyst.


Formation of Germ Layers

produced by the implanting embryo. This blocks form the second germ layer, the ectoderm.
the recognition of the embryo as a tissue foreign The embryo is now in the form of a disc
to the mother. having two layers.
A space a p p e a r s between the ectoderm
FORMATION OF GERM LAYERS (below) and the trophoblast (above). This
is the amniotic cavity (Fig. 4.6C), filled by
As the blastocyst develops further, it gives rise not amniotic fluid or liquor amnii. The roof of
only to the tissues and organs of the embryo but this cavity is formed by amniogenic cells
also to a number of structures that support the derived from the trophoblast, while its floor
embryo and help it to acquire nutrition. At a very is formed by the ectoderm.
early stage in development, the embryo proper Flattened cells arising from the endoderm
acquires the form of a three-layered disc. This is (or, according to some, from trophoblast),
called the embryonic disc (also called embryonic spread and line the inside of the blastocystic
area, embryonic shield, or germ disc). cavity. (This lining of flattened cells is called
The three layers that constitute this embryonic Heuser's membrane.) In this way, a cavity,
disc are: lined on all sides by cells of endodermal
origin, is formed. This cavity is called the
1. Endoderm {endo = inside) primary yolk sac (Fig. 4.6D).
2. Ectoderm (ecto = outside) The cells of the trophoblast give origin to a
3. Mesoderm (meso = in the middle)

These are the three germ layers. All tissues Inner cell mass

of the body are derived from one or more of


these layers (Chapter 21). Much of the student's
s t u d y of e m b r y o l o g y c o n c e r n s itself with
learning from which of these germ layers
particular tissues and organs develop. In the
further development of the blastocyst that we
will now consider, it is very important to have
a clear conception of the formation of germ Trophoblast
layers and of their fate.
We h a v e seen t h a t the b l a s t o c y s t is a
Amniogenic cells -
spherical cyst lined by flattened trophoblastic
cells, and that inside it there is a mass of cells, Amniotic cavity
the inner cell mass, attached eccentrically to
the trophoblast. Further changes are as follows:

1. S o m e cells of t h e i n n e r cell m a s s
differentiate (i.e. they become different
from others) into flattened cells, that
come to line its free surface (lower in
Fig. 4 . 6 A ) . T h e s e c o n s t i t u t e t h e
endoderm, which is thus the first germ
layer to be formed. Fig. 4.6 Differentiation of endoderm and ectoderm,
2. The remaining cells of the inner cell mass and the formation of the amniotic cavity and
become columnar (Fig. 4.6B). These cells the yolk sac.
Human Embryology

mass of cells called the extra-embryonic inside of the trophoblast, and the outside of
mesoderm (or primary mesoderm). These the amniotic cavity, is called the parietal
cells come to lie between the trophoblast or somatopleuric extra-embryonic
and the flattened endodermal cells lining mesoderm. (It is also referred to as the
the yolk sac, thus separating these from each chorionic plate.) The part lining the outside
other. These cells also separate the wall of of the yolk sac is called t h e visceral
the amniotic cavity from the trophoblast or splancbnopleuric extra-embryonic
(Fig.4.7A). mesodenn (Fig. 4.7B).
T h i s m e s o d e r m is c a l l e d ' e x t r a - From Fig. 4.7B it is clearly seen that the
e m b r y o n i c ' ' b e c a u s e it lies o u t s i d e the extra-embryonic coelom does not extend
embryonic disc. It does not give rise to any into t h a t p a r t of the e x t r a - e m b r y o n i c
tissues of the embryo itself. mesoderm which attaches the wall of the
Small cavities appear in the extra-embryonic a m n i o t i c cavity t o the t r o p h o b l a s t . T h e
mesoderm. Gradually these join together to developing embryo, along with the amniotic
form larger spaces and, ultimately, one large cavity and the yolk sac, is now suspended
cavity is formed. This cavity is called the in the e x t r a - e m b r y o n i c c o e l o m , and is
extra-embryonic coelom (Fig. 4.7B) (also attached to the wall of the blastocyst (i.e.
called the chorionic cavity). W i t h its trophoblast) only by this unsplit part of the
formation, the extra-embryonic mesoderm extra-embryonic mesoderm. This mesoderm
is split into two layers. The part lining the forms a structure called the connecting stalk.

Amniot c cavity
Future connecting
, stalk

Amniotic cavity
Embryonic disc ^- Amnion

^-Chorion
."<-i^m
\ \ Secondary
yolk sac

J. W— Trophoblast

~"~—"-1 -. v
Splanchnopleuric
and
Extra-embryonic Primary yolk sac
mesoderm Somatopleuric
extra-embryonic
"^-—__A--^ mesoderm
Extra-embryonic coelom B

Fig. 4.7 Formation of extra-embryonic mesoderm and extra-embryonic coelom. Note carefully, the
composition of the amnion, and of the chorion.
Formation of Germ Layers

Formation of Chorion and Amnion: At this the lining cells. These cells are no longer
stage, two very important membranes are flattened but become cubical (Fig. 4.7B).
formed. One is formed by the parietal extra- 9. At this stage, the embryo proper is a circular
embryonic mesoderm (on the inside] and the disc composed of two layers of cells: the
overlying trophoblast (on the outside); this upper layer (towards amniotic cavity) is the
is called the chorion (Fig. 4.7B). The other ectoderm, the cells of which are columnar,
is t h e amnion w h i c h is c o n s t i t u t e d by while the lower layer (towards yolk sac) is
amniogenic cells forming the wall of the the e n d o d e r m , made up of cubical cells.
amniotic cavity (excluding the ectodermal There is no indication yet of a head- or tail-
floor). These cells are derived from the end of the embryonic disc (Fig. 4.8).
trophoblast. We have already seen that the 10. However, we soon see that, at one circular
amnion is covered by the parietal extra- area near the margin of the disc, the cubical
e m b r y o n i c m e s o d e r m , and that the cells of the endoderm become columnar. This
connecting stalk is attached to it. area is called the prochordal plate. T h e
T h e c h o r i o n a n d a m n i o n play an a p p e a r a n c e of t h e p r o c h o r d a l p l a t e
important role in child birth (parturition) determines the central axis of the embryo
and we will refer to these again. (i.e. enables us to divide it into right and
With the appearance of the extra-embryonic left h a l v e s ) , a n d a l s o e n a b l e s us t o
mesoderm, and later of the extra-embryonic distinguish its future head and tail ends
coelom, the yolk sac becomes much smaller (Fig. 4.9).
than before and is now called the secondary I I. Soon after the formation of the prochordal
yolk sac. T h i s a l t e r a t i o n in size is plate some of the ectodermal cells lying
accompanied by a change in the nature of along the central axis, near the tail-end of

Endodermal
Embryonic cells here b e c o m e
disc s e e n in columnar.
surface view.

Centra! axis
is now a p p a r e n t .
Tail e n d
Columnar ectoderm
Head end

B
Cubical endoderm
P r o c h o r d a l plate

Fig. 4.8 Embryonic disc before appearance of a Fig. 4.9 Embryonic disc after establishment of a
central axis. (B) represents a section central axis. (B) represents a section
along the axis XY shown in (A). along the central axis.
Human Embryology

the disc, begin to proliferate, and form an plate, the ectoderm and endoderm remain
elevation t h a t bulges into the a m n i o t i c in c o n t a c t . In l a t e r d e v e l o p m e n t , t h e
cavity. This elevation is called the primitive ectoderm and endoderm mostly persist as a
streak (Figs 4.10A, B). The primitive streak lining epithelium. On the other hand, the
is at first a rounded or oval swelling, but bulk of the tissues of the body is formed
with elongation of the embryonic disc it predominantly from mesoderm. As there is
becomes a linear s t r u c t u r e lying in the no mesoderm in the prochordal plate, this
central axis of the disc. region remains relatively thin, and later
12. The cells that proliferate in the region of forms the bucco-pharyttgeal membrane.
the primitive streak pass sideways, pushing 14. The primitive streak gradually elongates,
t h e m s e l v e s b e t w e e n the e c t o d e r m a n d along the central axis of the embryonic disc.
endoderm (Fig. 4.111. These cells form the The disc also elongates and becomes pear-
intra-embryonic mesoderm (or secondary shaped {Fig. 4.12).
mesoderm) which is the third germ layer. 15. We have seen that when the embryonic disc
The process of formation of the primitive is first formed it is suspended {along with
streak, and of intra-embryonic mesoderm a m n i o t i c cavity a n d yolk sac) from the
by the streak, is referred to as gastrnlation. t r o p h o b l a s t by t h e c o n n e c t i n g s t a l k
13. T h e intra-embryonic mesoderm spreads (Figs 4 . 7 , 4 . 1 3 ) . To begin w i t h , t h e
throughout the disc except in the region of connecting stalk is very broad compared to
the prochordal plate. Note that the the size of the embryo. As the embryonic
mesoderm extends cranial to rhe prochordal disc enlarges in size, and also elongates,
plate, and here mesoderm from the two sides the connecting stalk becomes relatively
becomes c o n t i n u o u s across the midline small, and its attachment becomes confined
(Fig. 4.1 2). In rhe region of rhe prochordal to the region of the rail-end of the embryonic

— Prochordal piate

Primitive streak
(Ectodermal cells Mesodermal cells spread
proliferate here.) sidewards from
primitive streak.
Primitive streak
j
B
B

Fig. 4.10 Appearance of primitive streak. (B) is a Fig. 4.11 Formation of intra-embryonic meso-
section along axis XY shown in (A). derm. (B) is a section along axis KL in (A).
Formation of Germ Layers

^ Prochordal plate

/ I J \ , Developing notochord
Primitive streak —•—Jl 1 \ ^ - " ' \ Intra-embryonic
^~T> 1 *-\^ mesoderm

\ ^~—1 l^s\/-f— Cloacal membrane

Intra-embryonic nkLj-VT Extra-embryonic


into connecting stalk ,....„.* \
fl'M . - mesoderm in
*Z^ connecting stalk

Fig. 4.12 Spread of intra-embryonic mesoderm. Note that the mesoderm comes to lie
between ectoderm and endoderm in all parts of the embryonic disc except at
(1) the prochordal plate, (2) the cloacal membrane, and (3) the region of the
notochord.

disc (Fig. 4 . 1 3 ) . Some i n t r a - e m b r y o n i c region of the primitive streak and form the
mesoderm arising from the primitive streak, mesoderm. Other cells push the hypoblast aside
passes backwards into the connecting stalk and form the endoderm; while those that remain
(Figs 4.12, 4.13). As it does so, it leaves an form the ectoderm. Thus, according to this view,
area caudal to the primitive streak, where all the three germ layers are derived from the
ectoderm and endoderm remain in contact epiblast. It will be o b v i o u s t h a t the main
(i.e. mesoderm does not separate them). This d i f f e r e n c e f r o m c l a s s i c a l d e s c r i p t i o n is
r e g i o n is, t h e r e f o r e , s i m i l a r t o t h e regarding the formation of endoderm.
prochordal plate, a n d forms the cloacal According t o some workers, the prochordal
membrane (Fig. 4.12). plate and the neural crest also form some intra-
embryonic mesoderm.
ALTERNATIVE VIEW O F
F O R M A T I O N O F G E R M LAYERS USE O F STEM CELLS IN T H E
T R E A T M E N T O F DISEASES
The preceding account of the formation of germ
layers conforms to classical descriptions. Some By now readers must have realized that the cells
authorities describe the process differently, the of i n n e r cell m a s s h a v e t h e p o t e n t i a l t o
main points of difference being as follows: differentiate into three different germ layers
T h e layer of c o l u m n a r cells (Fig. 4.7A> (ectoderm, endoderm and mesoderm). Since all
(described above as ectoderm) is defined as the the cells, tissues and organs of the body are
epiblast. The layer of cuboidal cells (described formed from these three layers, the cells of the
above as endoderm) is defined as the hypoblast. inner cell mass are called embryonic stem cells.
Some cells of the epiblast migrate to the T h e s e s t e m cells c a n be m a i n t a i n e d a n d
Human Embryology

Primitive streak

-Extra-embryonic mesoderm

- Extra-embryonic coelom

Amniotic cavity - Connecting stalk


' (Note intra-embryonic
mesoderm passing
Intra-embryonic^ into it)
mesoderm

Extra-embryonic
coelom

Fig. 4.13 Diagram showing the attachment of the connecting stalk to the caudal end of the
embryonic disc. Note the cells of the intra-embryonic mesoderm passing into the
connecting stalk.

propagated in an undifferentiated state, in on 'therapeutic stem cell cloning'. In this


culture, in laboratories. If these cells are exposed procedure the nucleus of patient cell is
to certain specific growth factors, in culture, introduced into the embryonic stem cell.
the stem cells can form various types of adult These cells are then allowed to grow in any
cells, e.g. neurons, muscle cells, blood cells, tissue of the patient. As the tissues arising
cartilage cells, etc. It has been observed that from the stem cells are now genetically
when these stem cells are introduced into the identical to those oi the patient, rejection is
tissues of a living person, the local environment avoided.
helps these stem cells to differentiate into cells Though the embryonic stem cells are most
similar to those of the tissue in which they are suitable for therapeutic purposes, stem cells can
placed. This technique has tremendous also be isolated from some adult tissues, e.g.
potential for treatment of various diseases. Some bone marrow, brain and skeletal muscle.
of these are Parkinson's disease, Alzheimer However, adult stem cells are difficult to culture
disease, diabetes, myocardial infarction, blood in laboratories and have less potential to
diseases, severe burns, osteoporosis, spinal cord differentiate in adult tissues.
injury, to name but a few. However, in this As the human embryos are needed for stem
procedure the complication of immune rejection cell research, some authorities object to it on
is always present as the genetic constitution of ethical grounds. The main objections are that
stem cell is different from that of patient. To it is against nature and it treats the embryo
overcome this problem scientists arc working with disrespect.
Formation of Germ Layers

TIMETABLE OF EVENTS DESCRIBED IN the beginning of what is termed the embryonic


THIS CHAPTER period (3rd to 8th week). Most congenital
anomalies are produced by teratogens acting
Development of the embryo from fertilization
during this period.
up to the formation of the bilaminar disc is
described as the pre-organogenesis period, as
no organs are as yet recognizable. These events Age (in Days) Developmental Events
take place in the first 14 days of pregnancy. 2 Embryo is at two-cell stage
Anomalies produced by teratogens acting
3 Morula is formed
during this period usually result in death of the
embryo. These anomalies are, therefore, seldom 4 Blastocyst is formed
seen in babies reaching full term. 8 Bilaminar disc is formed
Establishment of the primitive streak and 14 Prochordal plate and
formation of intra-cmbryonic mesoderm mark primitive streak is seen
the onset of gastrulation. Gastrulation begins 16 Intra-embryonic mesoderm
in the third week and most of it will be is formed/disc is now three
considered in Chapter 5. The third week marks layered.
Chapter

5 Further Development of
Embryonic Disc

HIGHLIGHTS undergoes folding at the cranial and caudal


e n d s . T h e s e are the head and tail folds
The cranial end of the primitive streak enlarges (Fig. 5.7). Lateral folds also appear. As a result
to form the primitive knot (Fig. 5.1). of these folds the endoderm is converted into a
Cells of the primitive knot multiply and pass tube, the gut. It is divisible into foregut, midgut
cranially to form a rod-like structure reaching and bindgut.
up to the p r o c h o r d a l p l a t e . T h i s is the • After formation of the head fold the gut is closed
notochordal process. cranially by the prochordal plate, which is now
The notochordal process undergoes changes called the buccopharyngeal membrane. Caudally,
that convert it first into a canal and then into the gut is closed by the cloacal membrane.
a p l a t e , a n d finally back into a r o d - l i k e • The umbilical cord d e v e l o p s from the
structure. This is the notocbord. connecting stalk. It contains the right and left
Most of the notochord disappears. Remnants umbilical arteries, the left umbilical vein, and
r e m a i n as the nucleus pulposus of each remnants of the vitello-intestinal duct and yolk
intervertebral disc. sac. The ground substance of the umbilical cord
A w i d e s t r i p of e c t o d e r m o v e r l y i n g the is made up of Wharton's jelly derived from
notochord becomes thickened and forms the mesoderm. The cord is covered by amnion.
neural plate (Fig. SAC) from which the brain • The allantoic diverticulum arises from the yolk
and spinal cord develop. sac before formation of the gut (Fig. 5.10). After
Intra-embryonic mesoderm shows three f o r m a t i o n of the tail fold, it is seen as a
subdivisions (Fig. 5.4D). The mesoderm next diverticulum of the hindgut.
t o the m i d d l e line is called t h e paraxial • The pericardial cavity is derived from part of
mesoderm. It undergoes segmentation to form the intra-embryonic coelom that lies cranial
somites. The mesoderm in the lateral part of to the prochordal plate (Fig. 5.11). The deve-
the embryonic disc is called the lateral plate l o p i n g h e a r t lies v e n t r a l to t h e cavity
mesoderm. A cavity called the infra-embryonic (Fig. 5.12). After formation of the head fold
coelom appears in it and splits the mesoderm the pericardial cavity lies ventral to the foregut;
into a somatopleuric layer (in contact with and the developing h e a r t is d o r s a l to the
ectoderm) and a splanchnopleuric layer (in pericardial cavity (Fig. 5.13).
contact with endoderm) (Fig. 5.5D). A strip of • The septum transversum is made of intra-
mesoderm between the lateral plate mesoderm embryonic mesoderm that lies cranial to the
and the p a r a x i a l m e s o d e r m is called the pericardial cavity (Figs 5 . 1 1 , 5 . 1 2 ) . After
intermediate mesoderm. formation of the head fold it lies caudal to the
The intra-embryonic coelom later forms the pericardium and heart (Fig. 5.13). The liver
pericardial, pleural and peritoneal cavities. and the diaphragm develop in relation to the
T h e e m b r y o n i c disc, which is at first flat, septum transversum.
Further Development of Embryonic Disc

the streak is called the primitive knot,


FORMATION OF THE NOTOCHORD
primitive node or Benson's node (Figs 5.1 A,
The notochord is a midline structure, that develops 5.2A).
in the region lying between the cranial end of the 2. A depression appears in the centre of the
p r i m i t i v e s t r e a k a n d t h e c a u d a l end of the primitive knot. This depression is called the
prochordal plate (Figs 4.12, 5 . 1 , 5.2). During its blastopore (Figs 5.IB, 5.2B).
development, the notochord passes through several 3. Cells in the primitive knot multiply and pass
stages that are as follows: cranially in the middle line, between the
ectoderm and endoderm, reaching up to the
The cranial end of the primitive streak caudal margin of the prochordal plate. These
becomes thickened. This thickened part of cells form a solid cord called the notochordal

Prochordal plate Prochordal Primitive Primitive


plate
i
Primitive knot

-Primitive streak Cloacal membrane

Cloacal membrane Blastopore

B I^H

Notochordal process

Notochordal canal

Fig. 5.2 Sections through the embryonic disc


along the axis KL shown in Fig. 5.1 A to
Fig. 5.1 Formation of primitive knot (A), illustrate the formation of the primitive
blastopore (B) and notochordal process knot (A), blastopore (B), notochordal
(C). Note that the notochordal process is process (C) and notochordal canal (D).
deep to ectoderm and that its position is Stages A, B and C correspond to those
shown diagrammatically. of Fig. 5.1.
Human Embryology

process or bead process (Figs 5.1C, 5.2C, Some details of the process of formation of the
5.3A). The cells of this process undergo notochord are as follows:
several stages of rearrangement (Figs 5 . 1 ,
5.2) ending in the formation of a solid rod 1. After the formation of the blastopore, its
called the notochord. cavity extends into the n o t o c h o r d a l
p r o c e s s , a n d c o n v e r t s it i n t o a t u b e
As the embryo enlarges the notochord elongates called the notochordal canal (Figs 5.2D,
considerably and lies in the midline, in the position 5.3B).
to be later occupied by the vertebral column. 2. T h e cells f o r m i n g t h e floor of t h e
However, the notochord does not give rise to the notochordal canal become intercalated in
vertebral column. Most of it disappears, but parts (i.e. become mixed up with) the cells of
of it persist in the region of each intervertebral the e n d o d e r m (Fig. 5 . 3 C ) . T h e cells
disc as the nucleus pulposus. forming the floor of the notochordal canal
now separate the canal from the cavity of
the yolk sac.
3. The floor of the notochordal canal begins
to break d o w n . At first there are small
MMMMm (MfflSQMfflB openings formed in it, but gradually the
UZUJ-UJ* TXXXJJXD whole canal comes to communicate with
QDUJJJJ-XLU
the yolk sac (Fig. 5.3D). The notochordal
Notochordal process canal also communicates with the amniotic
Intra-embryonic cavity through the blastopore. Thus, at this
mesoderm stage, the amniotic cavity and the yolk
sac a r e in c o m m u n i c a t i o n with e a c h
other.
axtxamammw ffffflfflTffllfflffl-ffl 4. Gradually the walls of the canal become
flattened so that instead of a rounded canal
we have a flat plate of cells called the
Notochordal canal Notochordal plate
notochordal plate (Fig. 5.3E).
5. However, this process of flattening is soon
C F reversed and the notochordal plate again
becomes curved, to assume the shape of a
tube (Figs 5.3F, G). Proliferation of cells
of this tube converts it into a solid rod of
cells. This rod is the definitive (i.e. finally
G formed) notochord. It gets completely
separated from the endoderm.

The notochord is present in all animals that


Definitive notochord belong to the phylum Chordata. In some of them,
e.g. Amphioxus, it persists into adult life and
Fig. 5.3 Transverse sections through the forms the central axis of the body. In others,
embryonic disc (along the axis XY shown i n c l u d i n g m a n , it a p p e a r s in t h e e m b r y o
in Fig. 5.1C) to illustrate stages in the but only small remnants of it remain in the
formation of the notochord. adult.
Further Development of Embryonic Disc

F O R M A T I O N O F THE NEURAL TUBE


and (b) a caudal tubular part that forms the
spinal cord.
The details of the formation of the neural tube 4. In early embryos, the developing brain
will be studied later. For the time being, it may be forms a large conspicuous mass, on the
noted that: dorsal aspect.

1. The neural tube gives rise to the brain and The process of formation of the neural tube is
the spinal cord. referred to as neurulation.
2. The neural tube is formed from the ectoderm
overlying the notochord and, therefore, S U B D I V I S I O N S O F INTRA-
extends from the prochordal plate to the EMBRYONIC MESODERM
primitive knot (Fig. 5.4C).
3. The neural tube is soon divisible into: (a) a We have seen that the intra-embryonic mesoderm
cranial enlarged part that forms the brain, is formed by proliferation of cells in the primitive

Prochordal plate

Neural plate

Intermediate
mesoderm
Primitive streak

Paraxial mesoderm Neural plate


Ectoderm Extra-embryonic
Lat. plate mesoderm Lat. plate mesoderm mesoderm

m m ly.v.
termediate mesoderm

Notochord mesoderm

Fig. 5.4 Subdivisions of intra-embryonic mesoderm. 'B' and 'D' are transverse sections across axes XY
(in A) and MN (in C) respectively. Note that the notochord and mesoderm are deep to the
ectoderm. Their position is shown diagrammatically.
Human Embryology

streak and that it separates the ectoderm and the coelom (Fig. 5.5C}. With the formation of the intra-
e n d o d e r m , e x c e p t in t h e following r e g i o n s : embryonic coelom, the lateral plate mesoderm
(a) prochordal plate fb) cloacal membrane, and splits into:
(c) in the midline caudal to the prochordal plate,
as this place is occupied by the notochord. 1. Somatopleuric or parietal, intra-embryonic
Cranial to the prochordal plate, the mesoderm mesoderm that is in contact with ectoderm.
of the two sides meets in the midline (Fig. 4.12). 2. Splanchnopleuric, or visceral, intra-
At the edges of the embryonic disc, the intra- embryonic mesoderm that is in contact with
e m b r y o n i c m e s o d e r m is c o n t i n u o u s with the endoderm (Figs 5.5B, D}.
extra-embryonic mesoderm (Fig. 5AD). The intra-
embryonic mesoderm now becomes subdivided T h e i n t r a - e m b r y o n i c c o e l o m gives rise t o
into three parts (Fig. 5.4): pericardial, pleural, and peritoneal cavities. Their
development will be considered later. For the time
(A) T h e m e s o d e r m , o n e i t h e r side of t h e being note that the pericardium is formed from
notochord, becomes thick and is called the that part of the intra-embryonic coelom that lies,
paraxial mesoderm. in the midline, cranial to the prochordal plate.
(B) M o r e laterally, the m e s o d e r m f o r m s a T h e heart is formed in the s p l a n c h n o p l e u r i c
t h i n n e r l a y e r c a l l e d t h e lateral plate mesoderm forming the floor of this part of the
mesoderm. coelom (Fig. 5.6). This is, therefore, called the
( Q Between these two, there is a longitudinal cardiogenic area (also called cardiogenic plate,
strip called the intermediate mesoderm. heart-forming plate). Cranial to the cardiogenic
The paraxial mesoderm now becomes area (i.e. at the cranial edge of the embryonic
segmented into cubical masses called somitomeres disc} the s o m a t o p l e u r i c and splanchnopleuric
which give rise to somites (also called metameres mesoderm are continuous with each other. The
or primitive segments) (Figs 5.4C, D). The first mesoderm here does not get split, as the intra-
somites are seen on either side of the midline, a embryonic coelom has not extended into it. This
little behind the prochordal plate. More somites unsplit mesoderm forms a structure called the
are formed caudally, on cither side of the deve- septum transversum (Fig. 5.6).
loping neural tube. The somites have an interesting
history which is considered in Chapter 7.
YOLK SAC AND
FOLDING OF EMBRYO
FORMATION OF THE
INTRA-EMBRYONIC COELOM The early history of the yolk sac has been traced
in Chapter 4 (Figs 4.6, 4.7}. We have seen that the
While the p a r a x i a l m e s o d e r m is u n d e r g o i n g primary yolk sac is bounded above by cubical
segmentation, to form the somites, changes are endoderm of the embryonic disc and elsewhere by
also occurring in the lateral plate mesoderm. Small flattened cells lining the inside of the blastocystic
cavities appear in it. These coalesce (come together} cavity. With the formation of the extra-embryonic
t o form o n e l a r g e cavity, c a l l e d the intra- mesoderm, and later the extra-embryonic coelom,
embryonic coelom. The cavity has the shape of a the yolk sac becomes much smaller; it comes to
horseshoe (Figs 5.5A, C). There are two halves of be lined all round by cubical cells; and it is then
the cavity (one on either side of the midline} which called the secondary yolk sac.
are joined together cranial to the prochordal plate. The changes that now take place will be best
At first, this is a closed cavity (Fig. 5.5A) but soon understood by a careful study of Fig. 5.7. N o t e
it comes to communicate with the extra-embryonic the following:
Further Development of Embryonic Disc

Intra-embryonic coelom Neural groove

Splanchnopleuric intra- Somatopleuric intra-


embryonic mesoderm embryonic mesoderm

Intra-embryonic
mesoderm

Intra-embryonic coelom
opens into extra-embryonic
coelom here.

Amniotic cavity

Intra-embryonic
coelom

Extra-embryonic
coelom

Fig. 5.5 Intra-embryonic c o e l o m . (B) and (D) are sections across axes XY in (A) and (C) respectively.
(E) shows the relationship between the intra-embryonic and the extra-embryonic coeloms.

There is progressive increase in the size of and tail ends. These are called the head and
the embryonic disc. tail folds.
The head and tail ends of the disc (X, Y), With the formation of the head and tail folds,
however, remain relatively close together. parts of the yolk sac become enclosed within
H e n c e , the increased length of the disc the embryo. In this way, a tube lined by
causes it to bulge upwards into the amniotic endoderm is formed in the embryo. This is
cavity. the primitive gut, from which most of the
With further enlargement, the embryonic gastrointestinal tract is derived. At first, the
disc becomes folded on itself, at the head gut is in wide communication with the yolk
Human Embryology

bindgut; while the intervening


Somatopleuric p a r t is c a l l e d t h e midgut
Amiotic cavity , intra-embryonic (Fig. 5.7E). The communica-
S mesoderm
Prochordal di 3C T , tion with the yolk sac becomes
Ectoderm i
Intra-embryonic progressively narrower. As a
i - " coelom
result of these changes, the
- — Future septum yolk sac becomes small and
\ transversum i n c o n s p i c u o u s , a n d is n o w
~\~~~. Splanchnopleuric termed the definitive yolk sac
/ t ' intra-embryonic
mesoderm
(also called the umbilical
Endoderm Yolk sac
vesicle). The narrow channel
c o n n e c t i n g it to the gut is
Fig. 5.6 M i d l i n e section through cranial end of the embryonic disc to called the vitello-intestinal
show the relationship of the pericardial cavity to other duct (also called vitelline duct,
structures. yolk stalk or omphalomesen-
teric duct). This duct becomes
sac. T h e part of the gut cranial to this elongated and eventually disappears.
communication is called the foregut, the part As the head and tail folds are forming,
caudal to the communication is called the similar folds are also formed on each side.

Amniotic cavity

O *Q
Midgut
Foregut

Definitive yolk sac

Fig. 5.7 Formation of head and tail folds and establishment of the gut.
Further Development of Embryonic Disc

Embryonic Connecting
Extra-embryonic disc stalk
mesoderm
Amniotic
cavity

Trophoblast A m n j0tic Extra-embryonic


cavity

Fig. 5.8 Stages in the establishment of the umbilical cord.

These are the lateral folds. As a result, the is a circular aperture which may n o w be
embryo comes to be enclosed all around by called the umbilical opening.
ectoderm except in the region through which 6. As the embryonic disc folds on itself, the
the vitello-intestinai duct passes. Here, there amniotic cavity expands greatly, and comes
Human Embryology

to surround the embryo on all sides. In this 2. M e s o d e r m ( e x t r a - e m b r y o n i c ) of t h e


way, the embryo now floats in the amniotic c o n n e c t i n g s t a l k . T h i s m e s o d e r m gets
fluid, which fills the cavity. converted into a gelatinous substance called
Wharton's jelly. It protects blood vessels in
CONNECTING STALK the umbilical cord.
3. Blood vessels that pass from the embryo to
While discussing the f o r m a t i o n of the e x t r a - placenta.
embryonic coelom, we have seen that with the 4. A small part of the extra-embryonic coelom.
formation of this cavity, the embryo (along with
rhe amniotic cavity and yolk sac) remains attached This tube of amnion, and the structures within
t o t h e t r o p h o b l a s t only by e x t r a - e m b r y o n i c it, constitute the umbilical cord (Fig. 5.9). This
mesoderm into which the coelom does not extend cord progressively increases in length to allow free
(Figs 5.8A - C). This extra-embryonic mesoderm movement of the e m b r y o within the amniotic
forms the connecting stalk. We shall see later that cavity. At the time of birth of the child (i.e. at full
the t r o p h o b l a s t , a n d the tissues of the uterus, term), the umbilical cord is about half a metre
together form an important organ, the placenta, long, and about 2 cm in diameter. It shows marked
which provides the growing embryo with nutrition torsion, which is probably due to fetal movements.
and with oxygen. It also removes waste products An umbilical cord that is either too short or too
from the embryo. The importance of the connecting
long can cause problems during delivery of the
stalk is obvious when we see that this is the only
fetus.
connecting link between the e m b r y o and the
placenta.
As the embryo grows, the area of attachment
of the connecting stalk to it becomes relatively Umbilical vein
smaller. Gradually this attachment is seen only
Remnants of
n e a r the c a u d a l e n d of t h e e m b r y o n i c disc vitello-intestinal duct
(Figs5.8D,E). With the formation of the tail fold,
Remnants of allantois
the attachment of the connecting stalk moves (with
the tail end of the embryonic disc) to the ventral
aspect of the embryo. It is now attached in the
region of the umbilical opening (Fig. 5.8E). Umbilical artery
By now, blood vessels have developed in the
embryo, and also in the placenta. These sets of
Fig. 5.9 Section through umbilical cord.
blood vessels are in communication by means of
arteries and veins passing through the connecting
stalk. At first, there are two arteries and two veins
ALLANTOIC DIVERTICULUM
in the connecting stalk, but later the right vein
disappears (the left vein is 'left*'). Before the formation of the tail fold, a small
It is clear from Fig. 5.8F that, at this stage, the e n d o d e r m a l diverticulum called the allantoic
amnion has a circular attachment to the margins diverticulum arises from the yolk sac near the
of the umbilical opening and forms a wide tube in caudal end of the embryonic disc (Fig. 5.10A).
which the following lie: This diverticulum grows into the mesoderm of the
connecting stalk. After the formation of the tail
1. Vitello-intestinal duct and remnants of the fold, p a r t of t h i s d i v e r t i c u l u m is a b s o r b e d
yolk sac. into the hindgut. It now passes from the ventral
Further Development of Embryonic Disc

Allantoic diverticulum Hindgut

Amniotic /.
cavity / | Amniotic cavity

Extra-embryonic
coelom Mesoderm of ^_^_^m
A connecting stalk
1 B
Fig. 5.10 Allantoic diverticulum, and its relationship to the connecting stalk.

side of the hindgut into the connecting stalk the cavity (Fig. 5.13). The pericardium
(Fig. 5.10B). We will refer to it again while enlarges rapidly, and forms a conspicuous
considering the development of the urinary bulging on the ventral side of the embryo
bladder. (Fig. 5.14).
The septum transversum, which was the
EFFECT OF HEAD AND TAIL FOLDS most cranial structure in the embryonic disc
ON POSITIONS OF (Fig. 5.11), now lies caudal to the heart
OTHER STRUCTURES (Fig. 5.13). At a later stage in development,
the diaphragm and liver develop in relation
Just before the formation of the head and tail folds, to the septum transversum.
the structures in the embryonic disc are oriented,
as shown in Fig. 5.11. A median (midline) section Future Septum transversum
across the disc, at this stage, is shown in Fig. 5.12.
From the cranial to the caudal side, the structures Future Pericardial cavity
seen in the midline are (a) the septum transversum,
Prochordal plate
(b) the developing pericardial cavity and the heart,
(c) the prochordal plate, (d) the neural plate, Neural plate
(e) the primitive streak, and (f) the cloacal Position of
intra-embryonic
membrane. Note that the primitive streak is now coelom
inconspicuous. After folding, the relative positions
Somite
of these structures change to that shown in
Figs 5.13 and 5.14. The important points to note
here are as follows:
Developing spinal cord
1. With the formation of the head fold, the
developing pericardial cavity comes to lie Primitive streak
on the ventral side of the embryo, ventral Cloacal membrane
to the foregut. The heart, which was
developing in the splanchnopleuric
mesoderm in the floor of the pericardial Fig. 5.11 Embryonic disc showing the neural plate
cavity (Fig. 5.12), now lies in the roof of and related structures.
Human Embryology

Amniotic cavity Notochord Primitive streak

Neural plate

Prochordal plate

Fig. 5.12 Embryonic disc and related structures just before the formation of the
head and tail folds.

Foregut Amniotic cavity Notochord

Allantoic
diverticulum

Cloacal membrane

Fig. 5.13 Formation of head and tail folds. Also see Fig. 5.14.
Further Development of Embryonic Disc

Midgut Amniotic cavity

Foregut

Cloacal membrane

Septum transversum Allantoic diverticulum

Fig. 5.14 Later stage in the formation of the head and tail folds. Note the changing relationships of septum
transversum, pericardium, buccopharyngeal membrane, cloacal membrane and allantois.

3. The region of the prochordal plate now forms distal end of the hindgut is closed by the
the buccopharyngeal, or oral membrane, cloacal membrane. At first, this is directed
w h i c h c l o s e s t h e f o r e g u t c r a n i a 11 y. caudally (Fig. 5.13), but later it comes to
When this m e m b r a n e breaks d o w n , face ventrally (Fig. 5.14).
the foregut communicates with the
exterior. We have traced the development of the embryo
4. The most cranial structure of the embryo is to a stage when the rudiments of the nervous
now the enlarged cranial part of the neural system, the heart and the gut have been formed.
tube, which later forms the brain (Fig. 5.13). We are now in a position to trace the development
This enlarges enormously (Fig. 5.14). There of individual organ systems in detail. Before we
are n o w t w o big bulgings on the ventral do this, however, we must study the development
aspect of the embryo. Cranially, there is the of the placenta.
developing brain, and a little below it there
is the bulging pericardium (Fig. 5.14). In
Some Additional Points of Interest
between these t w o , there is a depression
called the stomatodaeum or stomodaeum, 1. In later life, r e m n a n t s of the primitive
t h e f l o o r of w h i c h is f o r m e d by t h e streak may give rise to peculiar tumours
buccopharyngeal membrane. that contain tissues derived from all three
5. Towards the tail end of the embryo, the germ layers. These t u m o u r s are seen in
primitive streak is n o w an inconspicuous the sacral region and are called
structure, that gradually disappears. The sacrococcygeal tumours.
Human Embryology

2. Experiments have shown that the of somites. In the head region, cranial to
formation of the neural tube is induced by somites, somitomeres give origin to some
the notochord. mesenchyme.
3. Somitomeres are not confined to the region 4. Wharton's jelly is rich in proteoglycans.

TIMETABLE OF EVENTS DESCRIBED IN THIS CHAPTER


• (in Days) Developmental Events
15 Primitive streak appears.
Definitive yolk sac is formed.
17 Notochordal process appears.
Heart tube is seen in cardiogenic area.
Allantoic diverticulum is seen.
19 Intra-embryonic mesoderm is being formed.
Connecting stalk can be distinguished.
21 Neural groove is seen.
Head fold begins to form.
23 Closure of the neural tube is seen.
Chapter

The Placenta
|6
which is a syncytium {cytoplasm with nuclei,
but no cell boundaries).
A developing e m b r y o gets attached to the
• The first-formed villi are called primary villi.
u t e r i n e e n d o m e t r i u m . T h i s is called
They consist of a central core of c y t o t r o -
implantation.
phoblast covered by syncytiotrophoblast.
In human beings the embryo gets buried in the
substance of the endometrium. This type of • Secondary villi have three layers. From inside
implantation is called interstitial implantation. o u t these are e x t r a - e m b r y o n i c m e s o d e r m ,
cytotrophoblast and syncytiotrophoblast.
After implantation the endometrium is called
the decidua. • In tertiary villi, blood capillaries are formed
in the extra-embryonic mesoderm.
The placenta is formed partly from embryonic
structures and partly from the decidua. It is • Villi are surrounded by an intervillous space
responsible for t r a n s p o r t of n u t r i e n t s and which contains maternal blood. As the placenta
enlarges, septa grow into the intervillous space
oxygen to the fetus, and for removal of waste
dividing the placenta into lobes. The fully
products.
formed placenta is about six inches in diameter
T h e essential elements of the placenta are and about 500 g in weight.
chorionic villi. The villi are surrounded by
maternal blood. Fetal blood circulates through • The placenta is normally attached to the upper
capillaries in villi. part of the body of the uterus. A placenta
attached lower down is called placenta praevia.
The maternal and fetal blood are separated by It can cause problems during child birth.
a very thin placental membrane (or barrier).
All substances passing from mother to fetus • The embryo is surrounded by three large cavities.
(and vice versa) traverse this membrane. These are the a m n i o t i c cavity, t h e e x t r a -
e m b r y o n i c coelom, and the uterine cavity.
The fetal tissue that takes part in forming the Enlargement of the amniotic cavity obliterates
placenta is chorion. It consists of trophoblast the extra-embryonic coelom, leading to fusion
(one layer of cells) resting on extra-embryonic of amnion and chorion. Further enlargement
mesoderm. of a m n i o t i c cavity o b l i t e r a t e s the uterine
Proliferation of cells of the trophoblast leads cavity. Fused a m n i o n a n d c h o r i o n (called
to formation of two layers: cytotrophoblast, membranes) bulge into the cervical canal
which is cellular and syncytiotrophoblast, (during child birth) and help to dilate it.
Human Embryology

IMPLANTATION
- Biasiocyst

After the ovum is shed from the ovary, it travels


through the uterine tube t o w a r d s the uterus. If - Uterine gland
fertilization occurs, segmentation of the ovum - Stroma
begins. By the time the fertilized ' o v u m ' reaches
the uterus, it has already become a morula. The
morula is still surrounded by the zona pellucida,
which prevents it from 'sticking 1 to the wall of the
Fig. 6.1 Relationship of blastocyst to uterine
uterine tube. The cells lining the surface of the
endometrium.
morula, constitute the trophoblast. The trophoblast
has t h e p r o p e r t y of a t t a c h i n g itself t o , a n d f e r t i l i z a t i o n . T h e t r o p h o b l a s t of the h u m a n
invading, any tissue it comes in contact with. Once blastocyst invades the endometrium of the uterus.
the zona pellucida disappears, the cells of the The blastocyst burrows deeper and deeper into
trophoblast stick to the uterine endometrium. This the uterine mucosa till the whole of it comes to lie
is called implantation (Fig. 6.1). In h u m a n s , within the thickness of the endometrium (Fig. 6.2).
i m p l a n t a t i o n begins on the 6 t h day after This is called interstitial implantation (Fig. 6.3).

Trophoblast

OffifflffiffifflHI^ $XU1T
Stroma
(Decidua)

Fig. 6.2 Stages in implantation of blastocyst.


The Placenta

DECIDUA
Decidua basalis
Amnion After the implantation of the embryo,
Choroin the uterine endometrium is called the
decidua. When the morula reaches the
uterus, the endometrium is in the
secretory phase. After implantation, the
features of the endometrium, which
are seen during the secretory phase of
the menstrual cycle, are maintained
Extra-embryonic and intensified. The stromal cells
coelom enlarge, become vacuolated, and store
Uterine lumen glycogen and lipids. This change in the
stromal cells is called the decidual
Myometrium
reaction.
Fig. 6.3 Manner of implantation in the human uterus. This The portion of the decidua where the
type of implantation is interstitial. Various other placenta is to be formed (i.e. deep to the
types occur in other mammals. developing blastocyst) is called the
decidua basalis (Fig. 6.4). The part of
The process of implantation is aided by proteo- the decidua that separates the embryo from the
lytic enzymes produced by the trophoblast. The uterine lumen is called the decidua capsularis,
uterine mucosa also aids the process. The while the part lining the rest of the uterine
trophoblastic cells which are situated over the cavity is called the decidua parietalis. The
inner cell mass, start penetrating the epithelium decidua basalis consists predominantly of large
of the endometrium. The implantation results decidual cells which contain large amounts of
due to the mutual interaction between tropho- lipids and glycogen (that presumably provide
blastic cells and endometrium. This interaction a source of nutrition for the embryo). The
is mediated by the receptors present on uterine decidua basalis is also referred to as the
epithelium and the secretion of 'L-selectin1 and decidual plate, and is firmly united to the
'integrins' by the trophoblast cells. chorion.
At the end of pregnancy, the decidua
Myometrium is shed off, along with the placenta and
membranes. It is this shedding off which
Decidua basalis
gives the decidua its name (c.f.
Chorion deciduous trees).

Decidua capsularis
FORMATION OF
CHORIONIC VILLI
Decidua parietalis
The essential functional elements of the
Uterine cavity
placenta are very small finger-like
processes or villi. These villi are
surrounded by maternal blood. In the
substance of the villi, there are
Fig. 6.4 Subdivisions or' decidua. capillaries through which the fetal blood
Human Embryology

1m
circulates. Exchanges between the maternal and
fetal circulations take place through the tissues
forming the walls of the villi (Fig. 6.5).
The villi are formed as offshoots from the surface
of the trophoblast. As the trophoblast, along with
.Capillary carrying the underlying extra-embryonic mesoderm,
fetal blood
constitutes the chorion, the villi, arising from it,
s Intervillous space are called chorionic villi.
containing
maternat blood The chorionic villi are first formed all over the
trophoblast and grow into the surrounding decidua
Blood vessels in
extra-embryonic
(Fig. 6.6A). Those villi related to the decidua
1 L^> mesoderm capsularis are transitory. After some time these
degenerate. This part of the chorion becomes
Fig. 6.5 Scheme to show that fetal b l o o d s m o o t h a n d is called t h e chorion laevae. In
circulating through capillaries of villi is i
contrast, the villi that grow into the decidua basalis
close relation to maternal b l o o d in the
intervillous space.
undergo considerable development. Along with the
tissues of the decidua basalis these villi form a
disc-shaped mass which is called the placenta
(Fig. 6.6B).The part ofthe chorion that helps form
Decidua basalis
the placenta is called the chorion frondosum.
T h e essential features of the f o r m a t i o n of
- Chorionic villi chorionic villi are as follows. The trophoblast is
at first m a d e u p of a single l a y e r of cells
(Fig. 6.7A). As these cells multiply, two distinct
layers are formed (Fig. 6.7B). The cells that are
nearest to the decidua (i.e. the most superficial
cells) lose t h e i r cell b o u n d a r i e s . T h u s , o n e
continuous sheet of cytoplasm containing many-
n u c l e i is f o r m e d . Such a t i s s u e is c a l l e d a
syncytium. Hence, this layer o f t h e trophoblast is
called the syncytiotrophoblast or plasmodio-
trophoblast. Deep to the syncytium, the cells of
the trophoblast retain their cell walls and form
the second layer called the cytotrophoblast (also
called Langhan's layer). The cytotrophoblast rests
on extra-embryonic mesoderm. All these elements
(syncytium, cytotrophoblast and mesoderm) take
Chorion laevae part in forming chorionic villi.
T h e following three stages in formation of
chorionic villi are seen:

Fig. 6.6 Two stages in the formation of chorionic (a) Primary villi consist of a central core of
v i l l i . Note their relationship to the c y t o t r o p h o b l a s t c o v e r e d by a layer of
decidua. In {B) note that the villi over the s y n c y t i o t r o p h o b l a s t . Adjoining villi are
decidua capsularis have disappeared. separated by an intervillous space.
The Placenta

Details of the process of


villus f o r m a t i o n a r e as
-Trophoblast follows:
_ Extraembryonic 1. The syncytiotrophoblast
mesoderm
grows rapidly and
becomes thick. Small
- Syncytiotrophoblast cavities (called lacunae)
a p p e a r in t h i s l a y e r
(Fig. 6.7C). Gradually,
B the lacunae increase in
- Cytotrophoblast
size. At first they are
irregularly arranged
(Fig. 6.7D), but gradu-
ally they c o m e t o lie
r a d i a l l y (Figs 6 . 8 A ,
6.9) a r o u n d the blasto-
Lacunae in thickened cyst. T h e lacunae are
" syncytiotrophoblast s e p a r a t e d from one
another by partitions of
syncytium, which are
called trabecular The
lacunae gradually com-
municate with each
other, so that eventually
o n e l a r g e s p a c e is
formed. Each trabe-
D c u l e is now surrounded
- Lacunae enlarge all around by this lacu-
nar space (Fig. 6.8B).
The syncytiotrophoblast
(in which these changes
are o c c u r r i n g ) g r o w s
Fig. 6.7 Early stages in formation of chorionic villi:
into the endometrium.
(A) Cytotrophoblast in contact with decidua;
(B) Syncytiotrophoblast formed; As the endometrium is
(C) Lacunae appear in syncytiotrophoblast; e r o d e d , s o m e of its
(D) The lacunae enlarge. blood vessels are open-
ed up, and blood from
them fills the lacunar
(b) Secondary villi show three layers: outer space (Fig. 6.10).
syncytiotrophoblast, an intermediate layer Each t r a b e c u l u s is, initially, m a d e up
of cytotrophoblast, and an inner layer of entirely of syncytiotrophoblast (Fig. 6.10).
extra-embryonic mesoderm N o w the cells of the cytotrophoblast begin
(c) Tertiary villi arc like secondary villi except to multiply and grow into each trabeculus
t h a t t h e r e are b l o o d c a p i l l a r i e s in t h e (Fig. 6.11 A). T h e trabeculus thus comes
mesoderm. to have a central core of cytotrophoblast
Human Embryology

4. Extra-embryonic meso-
A derm invades the centre of
each primary villus
(Fig. 6.12A). The villus

mf¥TlOE ^>Lacunar space


now has a core of meso-
derm (Fig. 6.12B) covered
._ by cytotrophoblast and by
syncytium. This structure
is called a secondary villus.
Trabeculae
5. Soon thereafter, blood
vessels can be seen in the
B mesoderm forming the
0 Lacunar space
completely
core of each villus. With
their appearance, the
* surrounding

• i each trabeculus villus is fully formed and


is called a tertiary villus
(Fig. 6.13). The blood
Fig. 6.8 Formation of chorionic v i l l i . The trabeculae are n o w •
vessels of the villus estab-
regularly arranged. (B) is a transverse section across (A) in the lish connections with the
plane XY. circulatory system of the
embryo. Fetal blood now
circulates through the villi,
while maternal blood
-Decidua basalis circulates through rhc
intervillous space.
.— Cytotrophoblast 6. From Figs 6.11A, 6.12A
and 6.13A it will be seen
Connecting stalk
that the cytotrophoblast,
that grows into the tra-
Syncytiotrophoblast beculus (or villus) does not
containing radially
arranged trabeculae penetrate the entire
thickness of syncytium
Extra-embryonic coelom and, therefore, does not
come in contact with the
Decidua capsulars decidua. At a later stage,
however, the cytotropho-
Fig. 6.9 Figure showing radial arrangement of trabeculae and blast emerges through the
lacunae around the blastocyst, syncytium of each villus.
The cells of the cytotro-
covered by an outer layer of syncytium. It phoblast now spread out to form a layer
is surrounded by maternal blood, filling that completely cuts off the syncytium from
the lacunar space. The trabeculus is now the decidua. This layer of cells is called
called a primary villus (Fig. 6.11) and the the cytotrophoblasticshell (Fig. 6.14). The
lacunar space is now called the intervillous cells of this shell multiply rapidly and the
space. placenta increases in size.
The Placenta

The villi that are first formed (as


. Uterine described above) are attached on the
blood vessel feta! side (Fig. 6.15) to the embryonic
mesoderm and on the maternal side to
the cytotrophoblastic shell. They are,
Maternal blood
in lacunar space therefore, called anchoring villi. Each
a n c h o r i n g villus consists of a stem
{truncus chorii); this divides into a
number of branches {rami chorii) which
in t u r n divide i n t o finer b r a n c h e s
iramuli chorii). The ramuli are
attached to the cytotrophoblastic shell.
Lacunar spaces The anchoring villi give off numerous
- seen in A' form one
continuous space branches which grow into the
fifled with maternal blood. i n t e r v i l l o u s s p a c e as free villi
(Fig. 6.16). N e w villi also sprout from
the chorionic side of the intervillous
space. Ultimately, almost the whole
Fig. 6.10 Uterine b l o o d vessels in the decidua open into the intervillous space becomes filled with
lacunar space and fill it w i t h maternal b l o o d . (B) is a villi. As a result, the surface a r e a
transverse section through trabeculae. a v a i l a b l e for e x c h a n g e s b e t w e e n
maternal and fetal circulations becomes
enormous.

These, newly formed, villi at first


consist only of syncytiotrophoblast.
They are subsequently invaded by
Syncytiotrophoblast
cytotrophoblast, mesoderm, and
Maternal blood blood vessels, and pass through the
in lacunar space
stages of primary, secondary and
Cytotrophoblast tertiary villi, as described above.
Extra-embryonic
mesoderm
FURTHER DEVELOPMENT
OF THE PLACENTA
• Primary villus
The placenta now becomes subdivided
- Syncytiotrophoblast
into a number of lobes, by septa that
„ Core of
cytotrophoblast grow into the intervillous space from
the maternal side (Fig. 6.17). Each such
Maternal blood lobe of the placenta is often called a
in lacunar space maternal cotyledon. If the placenta is
viewed from the maternal side, the
Fig. 6.11 Cells of cytotrophoblast grow into the
bases of the septa are seen as grooves
syncytiotrophoblast of each trabeculus. The (Fig. 6.18) while the cotyledons appear
trabeculae are now called primary v i l l i . as c o n v e x a r e a s b o u n d e d by t h e
Human Embryology

g r o o v e s . T h e n u m b e r of l o b e s
- Decidua generally varies between 15 and 20.
Each lobe c o n t a i n s a n u m b e r of
-Secondary villus
anchoring villi and their branches.
O n e such villus a n d its b r a n c h e s
- Syncytiotrophoblast
constitute a fetal cotyledon. The fully
- Cytotrop he-blast
formed placenta has 6 0 - 1 0 0 such
_ Extra-embryonic fetal cotyledons. The placenta n o w
mesoderm
forms a compact mass and is disc-
Maternal blood in shaped (Figs 6.17, 6.18).
intervillous space At full term (9 months after onset
of p r e g n a n c y ) the placenta has a
diameter of 6 to 8 inches and weighs
- Secondary villus
about 500 g. After the birth of the
- Syncytiotrophoblast child, the placenta is shed off along
with the decidua. The maternal
surface (formed by the decidual plate)
- Cytotrophoblast
is r o u g h a n d is s u b d i v i d e d i n t o
Extra-embryonic
mesoderm c o t y l e d o n s . T h e fetal s u r f a c e
(chorionic plate) is lined by amnion.
Fig. 6.12 Extra-embryonic mesoderm grows into the
It is smooth and is not divided into
cytotrophoblastic core of each primary villus,
cotyledons. The umbilical cord is
converting it into a secondary villus.
attached to this surface.

Decidua
Placental Membrane
Tertiary villus In the placenta, maternal blood circu-
lates through the intervillous space
Intervillous space
and fetal blood circulates through
Syncytiotrophoblast
blood vessels in the villi. The mater-
Cytotrophoblast
nal and fetal blood do not mix with
Capillary in extra-
each other. They are separated by a mem-
embryonic mesoderm
brane, made up of the layers of the
wall of the villus (Fig. 6.19A). These
Maternal blood in (from the fetal side) are as follows:
intervillous space
(a) the endothelium of fetal blood
v e s s e l s , a n d its b a s e m e n t
Tertiary villus membrane.
Syncytiotrophoblast
(b) surrounding mesoderm (con-
nective tissue).
(c) cytotrophoblast, a n d its base-
Cytotrophoblast
Capillaries in extra-embryonic ment membrane.
mesoderm (d) syncytiotrophoblast.

Fig. 6.13 Blood capillaries invade the extra-embryonic mesoderm of These structures c o n s t i t u t e the
each secondary villus thus converting it into a tertiary villus. placental membrane or barrier. All
The Placenta

Maternal blood vessels


of the membrane is increased,
in decidua by d i s a p p e a r a n c e of t h e
Cytotrophoblastic shell cytotrophoblastic layer from

I II II UE
m o s t villi, a n d by c o n s i -
Syncvtiotrophoblast d e r a b l e t h i n n i n g of t h e
Maternal blood in
connective tissue (Fig 6.19B).
intervillous space
This membrane, which is at
Cytotrophoblast
first 0 . 0 2 5 m m t h i c k , is
Capillary in villus r e d u c e d ro 0 . 0 0 2 m m .
Fetal blood vessels in However, towards the end of
extra-embryonic mesoderm pregnancy, a fibrinoid
d e p o s i t a p p e a r s on the
Fig. 6.14 Formation of cytotrophoblastic shell. Note that after
membrane, and this reduces
formation of this shell the syncytiotrophoblast is no longer in
contact w i t h maternal tissues.
its efficiency.

Functions of Placenta
Maternal surface
1 1. The placenta enables the
a - ^ ^ ^ ^ ^ ^ ^ ^ n _
t r a n s p o r t of o x y g e n ,
water, electrolytes and
nutrition (in the form of
carbohydrates, lipids,
f W Ramus ^ - ^ ^ JFRamuli y\g J^. polypeptides, amino acids
^ ^ M chorii ^k and vitamins) from
^M Truncus Mi Intervillus mp ^^L maternal to fetal blood. A
chorii ""•"' space ~~~f~^. ^ ^ full term fetus takes up
about 25 ml of oxygen per
m i n u t e from m a t e r n a l
b l o o d . Even a s h o r t
Fetal surface i n t e r r u p t i o n of oxygen
Umbilical cord
s u p p l y is fatal for t h e
fetus.
Fig. 6.15 Arrangement of anchoring villi and intervillous spaces within
the placenta. Note the subdivisions of each anchoring illus. 2. It a l s o p r o v i d e s for
e x c r e t i o n of c a r b o n
i n t e r c h a n g e s of o x y g e n , n u t r i t i o n a n d w a s t e dioxide, urea and other
products take place through this membrane. waste products produced by the fetus into
the maternal blood.
The total area of this membrane varies from 4 3. Maternal antibodies (IgG, gamma globulins
t o 14 square metres. It is interesting to note or I m m u o g l o b u l i n s ) reaching the fetus
that this is equal to the total absorptive area of through the placenta give the fetus immunity
the adult intestinal tract. As in the gut, the against some infections (e.g. diphtheria and
effective absorptive area is greatly increased measles).
by the presencejpf numerous microvilli on the 4. The placenta acts as a barrier and prevents
surface of the syncytiotrophoblast. many bacteria and other harmful substances
In the later part of pregnancy, the efficiency from reaching the fetus. However, most
Human Embryology

diethylstilbestrol) easily cross the placenta


and can have adverse effects on the fetus
(including carcinoma in later life).
5, While permitting the exchange of several
substances between the maternal and fetal
blood, it keeps these blood streams separate,
thereby, preventing antigenic reactions
between them.
6. The placenta synthesizes several hormones.
These are probably produced in the
syncytiotrophoblast.
Progesterone secreted by the placenta is
essential for maintenance of pregnancy after
the fourth month (when the corpus luteum
degenerates).
Oestrogens (mainly estriol) produced by
Fig. 6.16 Free villi arising from an anchoring villus.
the placenta reach maternal blood and
viruses (including poliomyelitis, measles and promote uterine growth and development
rubella) and some bacteria can pass across of the mammary gland.
it. Drugs taken by the mother may also enter
the fetal circulation and can produce CLINICAL CORRELATION
congenital malformations. Human chorionic gonadotropin (hCG) produced
As a rule, maternal hormones do not by the placenta is similar in its actions to
reach the fetus. However, synthetic luteinizing hormone of the hypophysis cerebri.
progestins and synthetic oestrogens (e.g. Gonadotropins are excreted through maternal

Maternal vessel opening Intervillous space


into intervillous space 0ne cotyledon-,
Septum
\ [ ... .1 i Anchoring villus

Umbilical cord

Fig. 6.17 Structure of a fully formed placenta. Each lobe (labelled cotyledon) contains a number of
anchoring villi but only one is shown here for the sake of simplicity.
The Placenta

urine where their presence is used as a test to


detect a pregnancy in its early stages.

Somatomammotropin (bCS) h a s an a n t i -
insulin effect on the mother leading to increased
plasma levels of glucose and amino acids in
t h e m a t e r n a l c i r c u l a t i o n . In t h i s w a y it
increases availability of these materials for the
fetus. It also enhances glucose utilization by
the fetus.

Circulation of Blood through


the Placenta
Blood flow t h r o u g h l a c u n a r spaces in the
syncytiotrophoblast begins as early as the 9th
day of pregnancy. Thereafter, the maternal
blood in the intervillous spaces is constantly in
circulation. Blood enters the intervillous space
Fig. 6.18 Placenta after shedding, through maternal arteries that open into the
the maternal aspect. space. The pressure of blood drives it right up
to the c h o r i o n i c p l a t e . Blood from the
intervillous spaces is drained by veins that also
Layers of barrier open into the same spaces.
Syncytiotrophoblast In the fully funned placenta, the intervillous
Cytotrophoblast spaces contain about 150 ml of blood which is
Mesoderm
replaced in 15 to 20 seconds {i.e. three to four
Endothelium of
times per minute).
fetal blood vessel

Fetal blood in capillary


' Maternal blood around villus NORMAL SITE OF IMPLANTATION
OF THE OVUM

The uterus can be divided into an upper part,


Layers of barrier consisting of the fundus and the greater part of the
Syncytiotrophoblast b o d y , a n d a l o w e r p a r t , c o n s i s t i n g of t h e
Endothelium of lower part of the body and the cervix. These are
fetal capillary c a l l e d t h e upper uterine segment, and the
lower uterine segment, respectively. It is the
upper uterine segment that enlarges during
pregnancy.
Fig. 6.19 Diagrams of placental barrier: (A) in early The placenta is normally attached only to the
part of pregnancy, and (B) in later part. upper uterine segment (Fig. 6.21).
Human Embryology

— Upper uterine
segment

Vagina

Fig 6.21 Upper and lower uterine


segments and their normal
relationship to the placenta.

Intervillus space
Umbilical cord

Fig. 6.20 Scheme to show h o w maternal blood


circulates through the placenta. Villi are ABNORMAL SITES OF
not drawn for the sake of simplicity. IMPLANTATION OF THE OVUM
Abnormal Implantation
within the Uterus
The attachment of the placenta may extend
partially, or completely, into the lower uterine
segment. T h i s condition is called placenta
praevia. It causes difficulty during childbirth
and may cause severe bleeding. Various degrees
of placenta praevia may be recognized, as given
below:

1. First degree: The attachment of the placenta


extends into the lower uterine segment, bur
does not reach the internal os (Fig. 6.22A).
2. Second degree: The margin of the placenta
reaches the internal os, but does not cover
it (Fig. 6.22B).
3. Third degree: T h e edge of the placenta
covers the internal os, but when the os
Third dilates during childbirth, the placenta no
degree
longer occludes it (Fig. 6.22C).
4 Fourth degree: T h e placenta completely
covers the internal os, and occludes the os
Fig. 6.22 Types of placenta praevia. even after it has dilated (Fig. 6.22D).
The Placenta

2. lobed, w h e n it is d i v i d e d i n t o l o b e s
(Fig.6.24B);
3. diffuse, when chorionic villi persist all round
the blastocyst: the placenta is thin and does
not assume the shape of a disc (Fig. 6.24C);
4. placenta succenturiata, when a small part
of the placenta is separated from the rest of
it (Fig. 6.24D);
5. fenestrated, when there is a hole in the disc
(Fig. 6.24E); and
6. ctrcumvallate, when the peripheral edge of
Fig. 6.23 Abnormal sites of implantation: the placenta is covered by a circular fold of
(1) Normal site, (2) Placenta praevia, decidua (Fig. 6.24F).
(3) Interstitial tubal implantation,
(4) Tubal implantation, (5) A b d o m i n a l The umbilical cord is normally attached to the
implantation, (6) O v a r i a n implantation. placenta near the centre (Fig. 6.25A). However,
this attachment may be:
Implantation outside the Uterus
When the ovum gets implanted at any site outside
the uterus, this is called an ectopic pregnancy. This
may be as follows:

1. Tubal pregnancy: T h e b l a s t o c y s t gets


i m p l a n t e d in t h e u t e r i n e t u b e . Such a
pregnancy cannot go on to full term, and
may result in rupture of the tube. After
r u p t u r e , the b l a s t o c y s t may a c q u i r e a
secondary implantation in the abdominal
c a v i t y (Fig. 6 . 2 3 ) , g i v i n g rise t o an
abdominal pregnancy.
2. Interstitial tubal implantation: The
blastocyst may get implanted in the part of
the uterine tube passing through the uterine
wall.
3. Implantation in the ovary: Fertilization and
implantation may occur while the ovum is
still in the ovary.

Other Anomalies of Placenta


Instead of being shaped like a disc, the placenta
may be: Fig. 6.24 Anomalies of placenta: (A) Bidiscoidal;
<B) Lobed, (C) Diffuse, (D) Placenta
1. bidiscoidal, when it consists of two discs succenturiata, (E) Fenestrated placenta,
(Fig. 6.24A); {F)Circumvallate placenta.
Human Embryology

Fig. 6.25 Variations in attachment of umbilical cord t o placenta: (A) N o r m a l , (B) M a r g i n a l . (C) Furcate,
(D) Velamentous insertion.

Amniotic cavity

Decidua / /*
parietalis / /
Decidua basalts

Fig. 6.26 Relationship of amniotic cavity, extra-embryonic coelom and uterine


cavity. For description see text.
The Placenta

1. marginal, when the cord is attached at the MUTUAL RELATIONSHIP OF


margin of the placenta (Fig. 6.25B) (this type AMNIOTIC CAVITY, EXTRA-
of placenta is called Battledore placenta); EMBRYONIC COELOM AND
or UTERINE CAVITY
2. furcate, when blood vessels divide before
reaching the placenta (Fig. 6.25C}. We have so far considered the fetal membranes
3. When blood vessels are attached to amnion, (amnion and chorion), and the placenta, mainly
where they ramify before reaching the in relation to the fetus. Let us now see their
placenta (Fig. 6.25D), the condition is relationships to the uterine cavity. These are
referred to as velanientous insertion. important, as they help us to understand some

Decidua capsulars Umbilical cord


(green)

Chorion / \ \ Decidua
(pink) / / \ \ ^ basalis

^ A \ V— Placenta
/ 1

(
s
Amniotic cavity j f 1 J
Amnion X.
(blue)

Uterine / / Decidua
cavity / / parietatis

N. > / / Note that amnion


I T~"""^ / and chorion have fused
to obliterate the
1 J A\ extra-embryonic coelom.

Fig. 6.27 Relationship of amniotic cavity and uterine cavity after obliteration
o f the extra-embryonic c o e l o m .
Human Embryology

aspects of the process of childbirth. The changing membrane. From Fig. 6.27 it will be seen that the
relationships will be best u n d e r s t o o d by first wall of the amniotic cavity is now formed by (i)
reviewing Figs 4.6, 4.7 and 4.13 and then by amnion, (ii) chorion, and (Hi) decidua capsularis,
studying Figs 6 . 2 6 - 6 . 2 8 . all three being fused t o one another.
In Fig. 6.26 we see three cavities, namely, the Further expansion of the amniotic cavity occurs
uterine cavity, the extra-embryonic coelom, and at the expense of the uterine cavity. Gradually,
the amniotic cavity. The outer wall of the extra- the decidua capsularis fuses with the decidua
embryonic coelom is formed by chorion and the parietalis, and the uterine cavity is also obliterated
inner wall by a m n i o n . As the amniotic cavity (Fig. 6.28). Still further expansion of the amniotic
enlarges, the extra-embryonic coelom becomes cavity is achieved by enlargement of the uterus.
smaller and smaller. It is eventually obliterated, H n l a r g e m e n t of t h e a m n i o t i c c a v i t y is
by fusion of a m n i o n and c h o r i o n . T h e fused accompanied by an increase in the a m o u n t of
chorion and amnion form the amniochorionic amniotic fluid.

Decidua capsularis Decidua parietalis


(dark green; (light green)

Uterine cavity has


been obliterated by 'Membranes'
fusion of decidua (fused amnion, chorion,
capsularis and and decidua capsularis)
decidua parietalis lie over the internal os.

Fig. 6.28 A m n i o t i c cavity after obliteration of the extra-embryonic c o e l o m


and uterine cavity.
The Placenta

At the time of parturition (childbirth), the fused transferred through the placenta to maternal
amnion and chorion (amniochorionic membrane) blood. When the fetal kidneys start working
(along with the greatly thinned out decidua the fetus passes urine into the amniotic fluid.
capsularis), constitute what are called the This does not cause harm because fetal urine is
membranes. As the uterine muscle contracts, made up mostly of water (metabolic wastes
increased pressure in the amniotic fluid causes these being removed from blood by the placenta and
membranes to bulge into the cervical canal. This not through the kidneys).
bulging helps to dilate this canal. The bulging In some cases hydramnios is associated with
membranes can be felt through the vagina and atresia of the oesophagus, which prevents
are referred to as the bag of waters. Ultimately swallowing of amniotic fluid by the fetus.
the membranes rupture. Amniotic fluid flows out Oligamnios is sometimes associated with renal
into the vagina. After the child is delivered, the agenesis as no urine is added to the amniotic
placenta and the membranes, along with all parts fluid.
of the decidua, separate from the wall of the uterus
and are expelled from it.
TIMETABLE OF SOME EVENTS
DESCRIBED IN THIS CHAPTER
AMNIOTIC FLUID
Age (in Days) Developmental Events
Amniotic fluid provides support for the delicate 8th day Trophoblast differentiates
tissues of the growing embryo or fetus. It allows into cytotrophoblast and
free movement and protects the fetus from external
syncytiotrophoblast.
injury. It also avoids adhesion of the fetus to
amnion. As pregnancy advances the quantity of 9th day Lacunae appear in rhe
this fluid increases, till at full term it is about one syncytium.
litre. 11 th day Embryo gets completely
The condition in which there is too much implanted in the
amniotic fluid (over 1500 ml) is called endometrium.
hydramnios; and when the fluid is too little it is 13th day Primary villi are formed.
called oligamnios. Both conditions can cause 16th day Secondary and tertiary villi
abnormalities in the fetus. They can also cause are seen.
difficulties during childbirth. 2nd month Villi are seen all around the
trophoblast.
There is constant exchange of water between 4th month A definitive placenta is
the amniotic fluid and maternal blood, the water formed.
being completely replaced every three hours.
Some time in the fifth month the fetus begins to Full term Placenta is shed about half
swallow amniotic fluid. This fluid is absorbed an hour after birth of the
(through the gut) into fetal blood and is baby.
Chapter

Formation of Tissues of the Body

HIGHLIGHTS ossification. Secondary centres (of variable


number) appear for bone ends. The part of bone
• Eptthelia may o r i g i n a t e from ectoderm, ossified from a secondary centre is called an
endoderm or mesoderm. epiphysis.
• Epithelia lining external surfaces of the body, • In growing bone the diaphysis and epiphysis are
and terminal parts of passages opening to the separated by the epiphyseal plate (which is
outside are derived from ectoderm. made up of cartilage). Growth in length of a
• Epithelium lining the gut and of organs that bone takes place mainly at the epiphyseal plate.
d e v e l o p as d i v e r t i c u l a of t h e g u t , is
• T h e p o r t i o n of d i a p h y s i s a d j o i n i n g the
endodermal in origin.
epiphyseal plate is called the metaphysis.
• Epithelium lining most of the urogenital tract
is derived from mesoderm. In some parts it is • Somites undergo division into three p a r t s .
endodermal in origin. These are: (a) the dermatome which forms the
dermis of the skin; (b) myotome which forms
• Mesenchyme is made up of cells that can give skeletal muscle; and (c) sclerotome which helps
rise to cartilage, bone, muscle, blood and to form the vertebral column and ribs.
connective tissues.
• Skeletal muscle is derived partly from somites
• Blood cells are derived from mesenchyme in and partly from mesenchyme of the region.
bone marrow, liver, and spleen. Lymphocytes
are formed mainly in lymphoid tissues. • M o s t smooth muscle is derived from
mesenchyme related to viscera, and blood
• M o s t bones are formed by endochondral
vessels.
ossification, in which a cartilaginous model is
first formed and is later replaced by bone. • Cardiac muscle is derived from mesoderm
Some bones are formed by direct ossification related to the developing heart.
of membrane (intramembranous ossification).
• Neurons and many neuroglial cells are formed
• An area where ossification starts is called a in the neural t u b e . T h e myelin sheaths of
centre of ossification. In the case of long bones peripheral nerves are derived from Schwann
the shaft (or diaphysis) is formed by extension cells, while in the central nervous system they
of ossification from the primary centre of are derived from oligodendrocytes.
Formation of Tissues of the Body

The human body is made up of many types of external acoustic meatus and outer surface
tissue. These are k n o w n as basic tissues of the of tympanic membrane.
body. They are as follows: 3. Epithelium of some parts of the mouth, lower
part of anal canal, terminal part of male
Epithelial Tissue urethra, parts of female external genitalia.
Epithelium consists of cells arranged in the form
Some Epithelia derived from Endoderm
of continuous sheets. Epithelia line the external
and internal surfaces of the body and of body 1. Epithelium of the entire gut except part of
cavities. t h e m o u t h a n d a n a l c a n a l (lined by
ectoderm).
Connective Tissue 2. Epithelium of auditory tube and middle ear.
Connective tissue proper includes loose connective 3. Epithelium of respiratory tract.
tissue, dense connective tissue and adipose tissue. 4. Epithelium over part of urinary bladder,
Blood, cartilage and bone are special connective urethra and vagina.
tissues.
Some Epithelia derived from Mesoderm
Muscular Tissue 1. Tubules of kidneys, ureter, trigone of urinary
bladder.
This is of three types: striated, cardiac and smooth.
2. Uterine tubes, uterus, part of vagina.
Nervous Tissue 3. Testis and its duct system.
4. Endothelium lining the heart, blood vessels
This tissue consists of neurons (nerve cells), nerve
and lymphatics.
cell processes (axons and dendrites) and cells of
5. M e s o t h e l i u m l i n i n g t h e p e r i c a r d i a l ,
neuroglia.
peritoneal and pleural cavities; and cavities
In this chapter we shall study the formation of
of joints.
these basic tissues.

GLANDS
EPITHELIA
Almost all glands, both exocrine and endocrine,
An epithelium may be derived from ectoderm, develop as diverticula from epithelial surfaces
endoderm or mesoderm. In general, ectoderm gives (Fig. 7.1A}. T h e gland may be derived from
rise to epithelia covering the external surfaces of elements formed by branching of one diverticulum
the body; and some surfaces near the exterior. (e.g. p a r o t i d ) or may be formed from several
Endoderm gives origin to the epithelium of most diverticula (e.g. lacrimal gland, prostate). The
of the gut; and of structures arising as diverticula opening of the duct (or ducts) is usually situated
from t h e g u t (e.g. t h e liver a n d p a n c r e a s ) . at the site of the original outgrowth. In the case of
Mesoderm gives origin to the epithelial lining of endocrine glands (e.g. thyroid, anterior part of
the greater part of the urogenital tract. hypophysis cerebri) the gland loses all contact with
the epithelial surface from which it takes origin.
Some Epithelia derived from Ectoderm The diverticula are generally solid to begin with
1. Epithelium of skin, hair follicles, sweat (Figs 7.1 A, B) and are canalized later (Fig. 7.1C).
glands, sebaceous glands, and m a m m a r y The proximal parts of the diverticula form the
glands. duct system. The distal parts of the diverticula
2. Epithelium over cornea and conjunctiva, form the secretory elements (Fig. 7.1D).
Human Embryology

T
Solid downgrowth from epithelium Branching of downgrowth

Canalization of ducts Formation of acini from terminal


parts of ducts

Fig. 7.1 Stages in the development of a typical gland.

Depending on the epithelium from which they


take origin, glands may be of ectodermal {e.g.
sweat glands, mammary glands), endodermal (e.g.
pancreas, liver), mesodermal (e.g. adrenal cortex),
or of mixed origin (e.g. prostate).

MESENCHYME
We have seen above that a small proportion of
m e s o d e r m a l cells give rise to e p i t h e l i a . T h e
r e m a i n i n g cells, t h a t m a k e up t h e bulk of
mesoderm, get converted into a loose tissue called
mesenchyme (Fig. 7.2). Mesenchymal cells have
the ability to form many different kinds of cells
that in turn give rise t o various tissues (Fig. 7.3).
Chondroblasts arising from m e s e n c h y m a l Fig. 7.2 Mesenchymal cells. Note the delicate
cells form c a r t i l a g e , osteoblasts form b o n e , cytoplasmic processes j o i n i n g the cells
myoblasts form muscle, while lymphoblasts and t o one another.
Formation of Tissues of the Body

- -M~ Osteocyte

Precursors of
celts of blood
Lymphoblast

Fig. 7.3 Derivatives of mesenchymal cells.


Human Embryology

haemocytoblasts form various cells of blood. In the third week of embryonic life, formation
Mesenchymal cells also give rise to endothelial of blood vessels and blood cells is first seen in the
cells from which blood vessels and the primitive w a l l of t h e y o l k s a c , a r o u n d t h e a l l a n t o i c
heart tubes are formed. However, after all these diverticulum and in the connecting stalk. In these
tissues have been formed many mesenchymal cells situations, clusters of mesodermal cells aggregate
are still left and they give rise to cells of various to form blood islands. These mesodermal cells are
types of connective tissue. then converted to precursor cells (haemangioblasts)
which give rise t o blood vessels and blood cells
(Fig. 7.4). Cells, which are present in the centre of
CONNECTIVE TISSUE
the blood island, form the precursors of all blood
As the name suggests, connective tissue serves as cells [haematopoietic stem cells). Cells at the
a connecting system binding, s u p p o r t i n g a n d periphery of the island form the precursors of blood
strengthening all other body tissues together. vessels (angioblasts).
Connective tissue consists of three components,
i.e. cells, fibres and ground substance.
The fibres and ground substance are Cubical cells lining the
s y n t h e s i z e d b y t h e c e l l s of t h e secondary yolk sac
connective tissue. Splanchnopleuric
extra-embryonic
Formation of Loose mesoderm
Connective Tissue Mesodermal cells
forming blood island
At the site of f o r m a t i o n of loose
connective tissue the mesenchymal
cells get converted into fibroblasts.
Fibroblasts secrete the g r o u n d
substance and synthesize the collagen, Epithelium lining
the yolk sac
r e t i c u l a r a n d clastic fibres. Some
m e s e n c h y m a l cells p r e s e n t in the
developing connective tissue also get Mesodermal cells of
blood island become
converted into histiocytes, mast cells, haemangioblasts
plasma cells and fat cells (Fig. 7.3).

FORMATION OF BLOOD
Blood is a specialized fluid connective
tissue, which acts as a major transport
system within the body. The formation Endothelial cells
of the cells of blood begins very early
in embryonic life (before somites have
appeared) and continues throughout
life. Blood formation is specially rapid
in the embryo to provide for increase
in blood volume with the growth of Fig. 7.4 Formation of blood cells and b l o o d vessel from a
the embryo. b l o o d island.
Formation of Tissues of the Body

Blood cells arising in the blood islands of the mesoderm surrounding the developing aona. These
yolk sac are temporary. They are soon replaced stem cells first form colonies in the liver.
by permanent stem cells, which arise from the In the late embryonic period the formation of

T-CELL
/ X T-LYMPHOCYTES
PRECURSOR
/
[
TOTIPOTENT \
HAFMAI ] „
PLEURIPOTENT
LYMPHOID
*-
STEM CELLS
B-CELL
\ (THSC) / (PLSC) B-LYMPHOCYTES
PRECURSOR


/ PLEURIPOTENA
HAEMAL
\ STEM CELL /
\ (PHSC) /

Colony forming unit


Through
for mast cells MAST CELLS &
*- various
and basophils BASOPHILS
(CFU-Ma/CFU-Bas)

Colony forming unit Throug


various
) EOSINOPHILS
for eosinophils
(CFU-Eo) staqes

Colony forming unit Throug 1


for neutrophils - various NEUTROPHILS
Colony forming unit » (CFU-G) stages
for neutrophils
and monocytes Throuc h
(CFU-GM)
• Colony forming unit
for monocytes - variou MONOCYTES
(CFU-M)

Burst forming unit


for erythrocytes Colony forming unit
for erythrocytes •> variou ERYTHROCYTES
(BFU-E)
(CFU-E)

Colony forming unit


» for megakaryoblasts
(CFU-Meg)
—**| MEGAKARYOBL/ S T | — • •MEGAKARYOCYTE 1 PLATELETS

Fig. 7.5 Scheme showing the terms applied to precursors of various blood cells. CFU = Colony Forming
Unit. BFU = Burst Forming Unit. Note the other abbreviations used for other precursor cells.
Human Embryology

b l o o d s t a r t s in t h e liver, w h i c h r e m a i n s an very obvious. In some situations, the intercellular


important site of blood cell formation till the sixth substance is permeated by elastic fibres forming
month of intrauterine life. elastic cartilage.
A l m o s t n e a r t h e m i d d l e of p r e n a t a l life, Mesenchymal cells surrounding the surface of
definitive haematopoietic stem cells from the liver the developing cartilage form a fibrous membrane,
migrate to colonize the bone marrow. At the time the perichondrium.
of the birth, blood formation is mainly in the bone
marrow. Here totipotent haemal stem cells give BONE
rise t o p l e u r i p o t e n t lymphoid stem cells a n d
pleuripotent haemal stem cells (Fig. 7.5). These To u n d e r s t a n d t h e f o r m a t i o n of b o n e , it is
stem cells form colony forming units (CFU). Cells necessary for students to know its normal structure.
of o n e p a r t i c u l a r c o l o n y f o r m i n g u n i t a r e This can be read up from the author's TEXTBOOK
committed to differentiate only into one line of
blood cells, i.e. erythrocytes, megakaryocytes, The unit of bone structure is called a lamellus.
granulocytes, monocytes, macrophages and
lymphocytes (Fig. 7.5). In the case of erythrocytes
stem cells divide so rapidly that they seem to burst.
They are therefore called burst forming units (BFU).
Their daughter cells then form colony forming
units.
In the adult, the main sites of blood formation
are bone marrow, lymph nodes, thymus and spleen. Bone acquires thickness by stacking of
lamellae over one another
As stated above the precursors of the various
types; of blond cells; :ire generally regarded as being
oi mesodermal in origin. However, blood forming
cells differentiating in relation to the wall of the
yolk sac a n d p r o b a b l y in the liver, may be
endodermal in origin.

FORMATION OF CARTILAGE

Cartilage is formed from mesenchyme. At a site Between adjoining lamellae there are spaces
called lacunae. These spaces are occupied by
where cartilage is to be formed, mesenchymal cells
cells of bone (osteocytes).
b e c o m e c l o s e l y p a c k e d . T h i s is c a l l e d a
mesenchymal condensation. The mesenchymal
cells then become rounded and get converted into
cartilage forming cells or chondroblasts (Fig. 7.3).
U n d e r t h e i n f l u e n c e of c h o n d r o b l a s t s , t h e
intercellular substance of cartilage is laid down.
Some chondroblasts get imprisoned within the
substance of this developing cartilage and arc
called chondrocytes. Some fibres also develop in
the intercellular substance. In hyaline cartilage,
collagen fibres are present, but are not seen easily. Fig. 7.6 Scheme to show that bone is made up of
In fibrocartilage, collagen fibres are numerous and lamellae.
Formation of Tissues of the Body

OF HUMAN HISTOLOGY. Some features are


shown in Figs 7.6 - 7.8.

Cells of Bone
Three main types of cells present in bone are as
follows:
Osteocytes are cells that are seen in mature
bone.
Capillaries in bone marrow
Osteoblasts are bone forming cells. These
cells are, therefore, seen wherever bone is
Fig. 7.7 Osteocytes placed amongst lamellae of bone. being laid down. They have abundant

Bony trabeculus

Marrow cavity

^^in •*r' "-JU. mkM


- Haversian canal

;-*:
- Haversian system
•us i ^ j y .

B
Fig. 7.8 (A) Structure of spongy bone. (B) Structure of compact bone.
Human Embryology

of the embryo, bone formation is preceded


Osteoblasts
iayinc down bone
by the formation of a cartilaginous model
that closely resembles the bone to be
IV formed. This cartilage is subsequently
replaced by (not converted into) bone. This
±__+_z-± •>.i ;» ^•_Z- kind of bone formation is called
endochondral ossification. Bones formed
in this way are, therefore, called cartilage

~w&^ \
Osteoclasts
^
(/~~ bones. In some situations (e.g. the vault
of the skull) formation of bone is not
preceded by formation of a cartilaginous
model. Instead, bone is laid down directly
in a fibrous membrane. This is called
removing bone intramembranous ossification and these
bones are called membrane bones. These
Fig. 7.9 Relationship of osteoblasts and osteoclasts to include the bones of the vault of the skull,
developing bone. the mandible and the clavicle.

basophilic cytoplasm and are arranged in


regular rows, looking very much like an Endochondral Ossification
epithelial lining (Fig. 7.9).
Osteoclasts are, on the other hand, The essential steps in the formation of bone by
responsible for bone removal. They are large endochondral ossification are as follows:
multinucleated cells and are seen in regions
where bone is being absorbed (Fig. 7.9). (a) At the site where the bone is to be formed,
the mesenchymal cells become closely
packed to form a mesenchymal
Formation of Bone condensation (Figs 7.10A, B).
All bone is of mesodermal origin. The process of (b) Some mesenchymal cells become
bone formation is called ossification. In most parts chondroblasts and lay down hyaline

'• :'*•"' '-•"-•"•' .'.-.'•:

f |I |
1 ill
SQB

&&&$

A B C D
Fig. 7.10 Endochondral ossification: Formation of cartilaginous m o d e l .
P = Perichondrium; ECC - Enlarged cartilage cells.
Formation of Tissues of the Body

cartilage (Fig. 7.IOC). Mesenchymal cells


on the surface of the cartilage form a
membrane called the perichondrium. This
membrane is vascular and contains
osteogenic cells.
(c) The cells of the cartilage are at first small
and irregularly arranged. However, in the
area where bone formation has to begin,
the cells enlarge considerably (Fig. 7.10D).
(d) The intercellular substance between the
enlarged cartilage cells becomes calcified,
under the influence of an enzyme called
alkaline phosphatase, which is secreted by
the cartilage cells. The nutrition to the cells
is thus cut off and they die, leaving behind
empty spaces called primary areolae
(Figs 7.11A, B).
(e) Some blood vessels of the perichondrium
(which may be called periosteum as soon
as bone is formed) now invade the calcified
cartilaginous matrix. They are accompanied
by osteogenic cells. This mass of vessels and
cells is called the periosteal bud. It eats away
much of the calcified matrix forming the
walls of the primary areolae, and thus
creates large cavities called secondary
areolae (Fig. 7.11C).
(f) The walls of the secondary areolae are
formed by thin layers of calcified matrix
that have not been dissolved. The osteogenic
cells become osteoblasts and arrange Fig. 7.11 Endochondral ossification: Formation of
themselves along the surfaces of these bars, primary and secondary areolae.
or plates, of calcified cartilaginous matrix
(Fig.7.12A). The calcified matrix of cartilage only acts as a
(g) These osteoblasts now lay down a layer of support for the developing trabeculae and is not
ossein fibrils embedded in a gelatinous itself converted into bone.
intercellular matrix. This material is called At this stage the ossifying cartilage shows a
osteoid (Fig. 7.12B). It is calcified and a central area (1 in Fig. 7.13A) where bone has been
lamellus of bone is formed (Fig. 7.12C). formed. As we move away from this area, we see
(h) The osteoblasts now lay down another layer the following:
of osteoid over the first lamellus. This is
also calcified. Thus two lamellae of bone (a) a region where the cartilaginous matrix has
are formed. Some osteoblasts that get caught been calcified and surrounds dead and dying,
between the lamellae form osteocytes. As cartilage cells (2 in Fig. 7.13A);
more lamellae are laid down, bony (b) a zone of hypertrophied cartilage cells, in
trabeculae are formed (Fig. 7.12D). an uncalcified matrix (3 in Fig. 7.13A); and
Human Embryology

Bar of calcified matrix

Osteoid (layer 1) -

1st lamellus

2nd (amellus-

Osteoid (layer 2)

Osteoid (layer 3) -

" Osteocyle —

total effect is that the ossifying cartilage


progressively increases in size.
Fig. 7.12 Endochondral ossification; Stages in the
formation of bony lamellae.
Development of a Typical Long Bone
(c) normal cartilage {4 in Fig. 7.13A) in which We may now consider how a long bone develops.
there is considerable mitotic activity.
1. A mesenchymal condensation is seen in the
If we see the same cartilage a little latter limb bud in the region where the bone is to
(Fig. 7.13B), we find that ossification has now be formed (Figs 7.14A, B).
extended into zone 2, and simultaneously the 2. The mesenchymal condensation is converted
matrix in zone 3 has become calcified. The deeper into a cartilaginous model. This model
cells of zone 4 have hypertrophied, while the more closely resembles the bone to be formed. It
superficial ones have multiplied to form zone 5. is covered by perichondrium (Fig. 7.14C)
In this way, formation of new cartilage keeps pace which has a superficial fibrous layer and a
with the loss due to replacement by bone. The deeper layer that has osteogenic cells.
Formation oi Tissues of the Body

/ •* Cartilaginous model

yti 1 Primary centre


Loose mesenchyme A gtf.i of ossification

! _
I
/ Cartilaginous model
has grown larger.

Mesenchymal
condensation flEK/l Rone formation has
B M f i j ^ — — extended towards the
ffifll ends from primary centre.

«§£ Periosteal collar


WHH has also elongated

Cartilage formed
by conversion of
mesenchymal cells
to cartilage cells

Fig. 7.15 Formation of a typical long bone:


Primary centre ot ossification and
periosteal collar.

This is accompanied by enlargement of the


Fig. 7.14 Formation of a typical long bone: cartilaginous model (Fig. 7.15).
Establishment of cartilaginous m o d e l . 5. Soon after the appearance of the primary
centre and onset of endochondral
3. Endochondral ossification starts in a small ossification in it, the perichondrium (which
area of the shaft as described above. This may now be called periosteum) becomes
area is called the primary centre of active. The osteogenic cells in its deeper
ossification (Fig. 7.15). layer lay down bone on the surface of the
4. Gradually, bone formation extends from the cartilaginous model by intramembranous
primary centre towards the ends of the shaft. ossification. This periosteal bone
Human Embryology

completely surrounds the cartilaginous shaft (Fig. 7.16A). At varying times after birth,
and is, therefore, called the periosteal collar secondary centres of e n d o c h o n d r a l
(Fig. 7.15). It is first formed only around ossification appear in the cartilages forming
the region of the primary centre but rapidly the ends of the bone (Fig. 7.16B). These
e x t e n d s t o w a r d s t h e e n d s of t h e centres enlarge until the ends become bony
cartilaginous model. The periosteal collar (Fig. 7 . 1 6 C ) . M o r e than one secondary
acts as a splint and gives strength to the centre of ossification may appear at either
cartilaginous model, at the site where it is end. The portion of bone formed from one
weakened by the formation of secondary secondary centre is called an epiphysis.
areolae. We shall see that most of the shaft
of the bone is derived from this periosteal For a considerable time after birth, the bone of
collar and is, therefore, intramembranous the diaphysis and the bone of the epiphysis are
in origin. s e p a r a t e d by a p l a t e of c a r t i l a g e called the
At about the time of birth, the developing epiphyseal cartilage, or epiphyseal plate. This is
bone consists of a part called the diaphysis formed by cartilage into which ossification has
(or shaft) (that is bony, and has been formed not extended either from the diaphysis or from the
by e x t e n s i o n of the p r i m a r y c e n t r e of epiphysis. We shall see that this plate plays a vital
ossification}; and ends that are cartilaginous role in growth of the bone.

Metaphysis

Diaphysis —*-

Marrow
cavity

Fig, 7.16 Formation of a typical long b o n e : Secondary centres of ossification.


Formation of Tissues of the Body

As the shaft increases in diameter, there is a


Growth of a Long Bone
corresponding increase in the size of the m a r r o w
A growing bone increases both in length and in cavity. This cavity also extends towards the ends
thickness. of the diaphysis but does not reach the epiphyseal
We have seen that the periosteum lays down a plate. Gradually, most of the bone formed from
layer of bone a r o u n d the shaft of the cartilaginous the primary centre (i.e. of endochondral origin) is
model. This periosteal collar gradually extends removed, except near the ends, so that the wall of
over the whole length of the diaphysis. As more the shaft is made up purely of periosteal bone
layers of bone are laid down over it, the periosteal f o r m e d by the p r o c e s s of i n t r a m e m b r a n o u s
bone becomes thicker and thicker. However, it is ossification.
neither necessary nor desirable for it to become To understand how a bone grows in length, we
t o o thick. Hence, osteoclasts c o m e t o line the will now have a closer look at the epiphyseal plate.
internal surface of the shaft and remove bone from Depending on the arrangement of its cells, three
this aspect. As bone is laid down outside the shaft, zones can be recognized (Fig. 7.18).
it is removed from the inside. The shaft thus grows
in diameter, and at the same time, its wall does (a) Zone of resting cartilage: Here, the cells
not become too thick (Fig. 7.17). The osteoclasts are small and irregularly arranged.
also remove the trabeculae lying in the centre of (b) Zone of proliferating cartilage: Here, the
the b o n e t h a t were formed by e n d o c h o n d r a l cells are larger and are undergoing repeated
ossification. In this way, a marrow cavity is formed. mitosis. As they multiply, they come to be

Layer 2 removed
Periosteal bone

Endochondral bone

Layer 3

Layer 4 deposited

Layer 3 removed

Layer 4

Layer 5 deposited
Layer 3 deposited

Fig. 7.17 Formation of a typical long bone: Increase in thickness. Note that the shaft is ultimately made up
almost entirely of periosteal bone formed by the process of intramembranous ossification.
Human Embryology

Size after
some growth

1 Bone has grown


Initial size I by this length.

Bone of
Formed by
epiphysis
!• growth
of cartilage

Epiphyseal This part of


plate Y cartilage has
become bone.

Bone of
diaphysis

Fig. 7.19 G r o w t h in length of bone at epiphyseal


cartilage.

continues to extend into it until the whole of


Endochondral
the epiphyseal plate is converted into bone. The
bone bone of the diaphysis and epiphysis then
„ Marrow
becomes continuous. This is called fusion of
cavity epiphysis.
Periosteal bone
Metaphysis
Fig. 7.18 Structure of epiphyseal cartilage. The portion of diaphysis adjoining the
epiphyseal plate is called the metaphysis. It is a
arranged in parallel columns, separated by region of active bone formation and, for this
bars of intercellular matrix. reason, it is highly vascular. The metaphysis does
(c) Zone of calcification: Here, the cells become not have a marrow cavity. Numerous muscles and
ligaments are usually attached to the bone in this
still larger and the matrix becomes
region. Even after bone growth has ceased, the
calcified.
calcium-turnover function of bone is most active
in the metaphysis, which acts as a storehouse of
Next to the zone of calcification, there is a zone calcium. This region is frequently the site of
where cartilage cells are dead and the calcified infection.
matrix is being replaced by bone. Growth in length
of the bone takes place by continuous
Interstitial and Appositional Growth
transformation of the epiphyseal cartilage to bone
(Figs 7.18,7.19) in this zone (i.e. on the diaphyseal Tissues grow by two methods. In some of these,
surface of the epiphyseal cartilage). At the same growth takes place by multiplication of cells (or
time, the thickness of the epiphyseal cartilage is by increase in intercellular material) throughout
maintained by active multiplication of cells in the the substance of the tissue. This is called interstitial
zone of ptoliferation. When the bone has attained growth. As a result, the tissue expands equally in
its full length, cells in the epiphyseal cartilage stop all directions and its shape is maintained. Cartilage
proliferating. The process of ossification, however, (and most other tissues) grow in this way.
Formation of Tissues of the Body

On the other hand, bone grows only by localised to a particular part of the skeleton, or
deposition of more bone on its surface, or at its may be generalised. Anomalies of individual parts
ends. This is called appositional growth. of the skeleton are considered in Chapter 10. Some
anomalies that affect the skeleton as a whole are
Remodelling as follows:
We have seen above that when a tissue grows by
interstitial growth it is easy for it to maintain its 1. Disorderly and excessive proliferation of
shape. However, this is not true of bone, which cartilage cells in the epiphyseal plate, or
can grow only by apposition. This will be clear the failure of normally formed cartilage to
from Fig. 7.20. In this figure the brown line be replaced by bone, leads to the formation
represents the shape of a bone end. The green line of irregular masses of cartilage within the
represents the same bone end after it has grown metaphysis. This is called dyschondroplasia
for some time. It will be clear that some areas of or enchondromatosis.
the original bone have to disappear if proper shape 2. Abnormal masses of bone may be formed
is to be maintained. This process of removal of in the region of the metaphysis and may
unwanted bone is called remodelling. protrude from the bone. Such a protrusion
The trabeculae of spongy bone and the is called an exostosis, and the condition is
Haversian systems of compact bone are so called multiple exostoses or diaphyseal
arranged that they are best fitted to bear stresses aclasis. This condition may be a result of
interference with the process of remodelling
imposed on them. This arrangement can change
of bone ends.
with change in stresses acting on the bone. This
process is often called internal remodelling. 3. Calcification of bone may be defective
[osteogenesis imperfecta) and may result in
multiple fractures.
4. Parts of bone may be replaced by fibrous
Bone and cartilage formation may sometimes be tissue {fibrous displasia).
abnormal as a result of various genetic and 5. Bones may show increased density or
environmental factors. The anomalies may be osteosclerosis. One disease characterised by
increased bone density is known as
osteopetrosis, or marble bone disease.
Early outline 6. In the condition called achondroplasia, there
of bone end
Area of bone that is insufficient, or disorderly, formation of
needs to be removed bone in the region of the epiphyseal
to maintain shape
of bone cartilage. This interferes with growth of long
bones. The individual does not grow in
height and becomes a dwarf. A similar
condition in which the limbs are of normal
length, but in which the vertebral column
remains short, is called chondro-osteo-
dystrophy.
7. Anomalous bone formation may be confined
to membrane bones. One such condition is
cleido-cranial dysostosis in which the
Fig. 7.20 Remodelling of bone ends during clavicle is absent and there are deformities
growth. of the skull. On the other hand, anomalies
Human Embryology

like achondroplasia and exostoses are FORMATION OF MUSCLE


confined to cartilage bones.
Fate of Somites
Generalized underdevelopment (dwarfism),
or overdevelopment, (gigantism) of bone We have seen that the paraxial mesoderm becomes
may be present. Sometimes all bones of one segmented to form a number of somites that lie on
half of the body are affected (asymmetric either side of the developing neural tube. A cross-
development). Overdevelopment or section through a somite shows that it is a
underdevelopment may be localised, e.g. to triangular structure and has a cavity (Fig. 7.21A).
a digit, or to a limb. The somite is divisible into three parts:

Surface ectoderm

Myotome

Cells of sclerotome migrate


medially and surround the
neural tube.

Spinal nerve
Myotome

Epidermis derived
Cells of the dermatome from surface ectoderm
migrate to the surface
and form the dermis
of the skin.
Epimere
• innervated by Myotome gives
dorsal ramus origin to muscles

Cells derived from the


sclerotome form the
mesenchymal basis of vertebrae.

Fig. 7.21 (A) Somites lying on either side of the neural tube. Note subdivisions of somite.
(B) The cells of the sclerotome have migrated medially and n o w surround the neural tube.
The myotome is innervated by nerves growing out of the neural tube. (C) The cells of the
dermatome have migrated to form the dermis of the skin.
Formation of Tissues of the Body

(a) T h e v e n t r o m e d i a l p a r t is c a l l e d t h e derived from the myotomes and subsequently


sclerotome. The cells of the sclerotome migrates to these regions. Such migration of
migrate medially. They surround the neural myotomes can be seen in embryos of some lower
tube and give rise to the vertebral column animals, but not in the human embryo. In m a n ,
and ribs (Figs 7.21B, C). the myotomes appear t o give origin only to the
(b) The lateral part is called the dermatome. musculature of the trunk, in whole or in part.
The cells of this part also migrate and come T h e occipital myotomes are believed to give
to line the deep surface of the ectoderm rise to the musculature of the tongue, while the
covering the entire body. These cells give extrinsic muscles of the eyeball are regarded
rise t o t h e d e r m i s of t h e skin a n d t o as derivatives of the pre-occipital myotomes.
subcutaneous tissue. Soon after its formation, each myotome, in
(c) The intermediate part is the myotome. It the neck and trunk, separates into a dorsal part
gives rise to striated muscle as described in (epimere) w h i c h gives rise t o the muscles
the following section. supplied by the dorsal primary ramus of the
spinal nerve and a ventral part (hypomere),
In the cervical, thoracic, lumbar and sacral which gives origin to the muscles supplied by
regions one spinal nerve innervates each myotome. the ventral ramus (Fig. 7.21C). The epimeres
The number of somites formed in these regions, give origin to the muscles of the back (extensors
therefore, corresponds to the number of spinal of the vertebral column), while the hypomeres
nerves. In the coccygeal region, the somites exceed give origin to the muscles of the body wall and
the number of spinal nerves but many of them limbs.
subsequently degenerate. In addition to the myotomes, striated muscle
The first cervical somite is not the most cranial may also arise in-situ from mesenchyme of the
somite to be formed. Cranial to it, there are: region. The limb muscles develop in this way,
in the mesenchyme of the limb b u d s . T h e
1. the occipital somites (four to five) which give muscles of the anterior parts of the abdominal
rise t o muscles of the t o n g u e a n d are and thoracic walls probably also arise in-situ.
supplied by the hypoglossal nerve;
2. the pre-occipital (or pre-otic) somites
Smooth Muscle
(somitomeres) supplied by the third, fourth
and sixth cranial nerves. A l m o s t all s m o o t h m u s c l e is d e r i v e d f r o m
mesenchyme. Smooth muscle in the walls of viscera
(e.g. the stomach) is formed from splanchnopleuric
Development of Striated Muscle
mesoderm in relation to them.
Striated muscle is derived from somites and also However, the muscles of the iris (sphincter and
from mesenchyme of the region. dilator pupillae) and myoepithelial cells of the
sweat glands are derived from ectoderm.
We have seen that each myotome establishes
c o n t a c t w i t h o n e segmental nerve. H e n c e ,
Cardiac Muscle
theoretically, the embryological derivation of
a muscle should be indicated by its nerve supply. This is derived from splanchnopleuric mesoderm
O n this basis it would be presumed that all the in relation to the developing heart tubes a n d
musculature of the body wall a n d limbs is pericardium.
Human Embryology

NERVOUS TISSUE The neural tube is at first lined by a single layer


of cells (Fig. 7.22A). These proliferate to form
Nervous tissue consists of cells, fibres and blood several layers (Fig. 7.22B). Nearest the lumen of
vessels. Two different categories of cells are found the tube is the matrix cell layer (also called
in nervous tissue, i.e. neurons and neuroglial cells. primitive ependymal or germinal layer). The
The neurons are cells which generate and conduct cells of this layer give rise to nerve cells, to
nerve impulses while the neuroglial cells are neuroglial cells, and also to more germinal cells.
supporting structures. Neurons have many Next comes the mantle layer in which are seen
processes, i.e. axons and dendrites. These processes the developing nerve cells, and neuroglial cells.
collect to form nerves. Blood vessels of nervous The outermost layer, termed the marginal
tissue are not derived from the neural tube but zone, contains no nerve cells. It consists of a
enter it from surrounding mesoderm. reticulum formed by protoplasmic processes of
developing neuroglial cells (spongioblasts). It
provides a framework into which the processes of
Formation of Neurons and nerve cells developing in the mantle layer can
Neuroglial cells grow.
The neurons and many neuroglial cells are formed The stages in the formation of a nerve cell are
in the neural tube. as follows:

Germinal layer

Mantle layer ^

Fig. 7.22 Layers of the neural tube. Although the germinal (neuroepithelial) layer is shown as a simple
epithelium it is actually pseudostratified.
Formation of Tissues of the Body

A
s> Apolar
neuroblast

Multipolar T*_

0 neuroblast

D
\ Unipolar
/ ~ neuroblast

C
<y-
Fig. 7.23 Stages in formation of a typical neuroblast.

One of the germinal cells passes from the The process of the cell which does not
germinal layer to the mantle layer and disappear now elongates, and on the side
becomes an apolar neuroblast {Fig. 7.23A). opposite to it numerous smaller processes
Two processes develop and convert the form. At this stage the cell is called a
apolar neuroblast to a bipolar neuroblast multipolar neuroblast (Fig. 7.23D).
(Fig. 7.23B). The main process of the multipolar
One of the processes of the neuroblast neuroblast now grows into the marginal
disappears, and it can now be called a layer, and becomes the axon of the nerve
unipolar neuroblast (Fig. 7.23C). cell (Fig. 7.22C). The axon can grow to a

Derivation of nerve cells (neurons) and neuroglial cells

From neurai tube From neural crest From mesenchymal cells

Neurons Schwann cells Microglia


Fibrous astrocytes Dorsal n. root ganglion cells Oligodendroglia (?)
Protoplasmic astrocytes Cells of other sensory ganglia
Oligodendroglia (?) Neurons in sympathetic ganglia
Ependymal cells

Fig. 7.24 Various types of cells derived from neuroepithelium.


Human Embryology

considerable length. It may either remain


within the central nervous system, or may Axon lying near
grow out of it as an efferent nerve fibre of a Schwann cell
p e r i p h e r a l - n e r v e . At its d e s t i n a t i o n it
establishes connections, either with the cell
Axon invaginates
bodies and dendrites of other neurons, or the Schwann cell.
with an effector organ (e.g. muscle).
6. The smaller processes of the neuroblast are
the dendrites. These ramify and establish
connections with other nerve cells.
7. At first the cytoplasm of the nerve cell is
homogeneous. Later Nissl's granules make
their a p p e a r a n c e . After their formation,
neurons lose the ability to divide.
^ Mesaxon formed by further
invagination by the axon
Neuroglial cells are also formed from germinal
cells of the ependymal layer (Fig. 7.24). These
cells (glioblasts) migrate to the mantle and
marginal zones as medulloblasts (also called
spongioblasts), which differentiate either into
astroblasts, and subsequently into astrocytes, or
Mesaxon elongates
i n t o oligodendroblasts and then into and begins to wind * • '
oligodendrocytes. T h e r e is a t h i r d t y p e of round the axon.
neuroglial cell called microglia. This type does
not develop from the cells of the neural tube, but
migrates into it along with blood vessels. These
cells are believed to be of mesodermal origin. • * - Mesaxon makes several
turns around the axon.
We h a v e seen t h a t t h e e p e n d y m a l (or Lipids are deposited
neuroepithelial) cells give rise both to neuroblasts between the layers
and to neuroglia. However, these two cell types of mesaxon. The myelin
sheath is thus formed.
are not formed simultaneously. The neuroblasts
are formed first. Neuroglial cells are formed after
the differentiation of neuroblasts is completed. A thin layer of Schwann
cell cytoplasm persists
as the neurilemma.
Formation of Myelin Sheath
Nerve fibres, which remain within the brain and Fig. 7.25 Invagination of Schwann cell by an axon
spinal c o r d , receive s u p p o r t f r o m , a n d a r e and formation of myelin sheath. (A) A x o n
ensheathed by, neuroglial cells. However, the lies near Schwann cell. (B) A x o n invagi-
nerve fibres, which leave the central nervous nating Schwann cell. (C) Mesaxon
formed (by t w o layers of cell membrane).
system to become constituents of peripheral nerves,
(D) Mesaxon elongates. (E) Several layers
acquire a special sheath called the neurolemma.
of mesaxon spiral around the axon to
This sheath is derived from some cells of the neural
form the basis of the myelin sheath.
crest that are called Schwann cells. At a later stage
Lipids are deposited between layers of
of development, a large number of nerve fibres, the mesaxon.
Formation of Tissues of the Body

both inside and outside the central nervous system,


develop another sheath between the neurolemma
and the axon. This is called the myelin sheath.
The myelin sheaths of peripheral nerves are derived
from the same Schwann cells which form the
neurolemma. In the central nervous system itself,
however, there are no Schwann cells and the myelin
sheath is formed by neuroglial cells called
oligodendrocytes.
The relationship of an axon to a Schwann cell
is illustrated in Fig. 7.25. Note the following
points:
Fig. 7.26 Relation of unmyelinated nerve fibres
1. Each axon invaginates the cytoplasm of a to a Schwann cell. Note that a number
Schwann cell and thus comes to be comp- of axons invaginate into a single
letely surrounded by it (Figs 7.25A, B). Schwann cell; and thai the mesaxons
Along the line of invagination, the cell remain short.
membrane of the Schwann cell becomes
drawn in to form a double layered of development. The process begins during the
mesentery-like membrane called the fourth month of intrauterine life, but is not
mesaxon (Fig. 7.25C). completed until the child is two to three years old.
2. In the case of myelinated nerve fibres, the Nerve fibres become fully functional only after
mesaxon elongates and becomes spirally they have acquired their myelin sheaths.
wound around the axon (Figs 7.25D, E). The blood vessels of the brain, and their
Some fatty substances are deposited between surrounding connective rissue, are not derived from
adjacent layers of the mesaxon and, together the neural tube. These arc mesodermal in origin
with it, form the myelin sheath. and invade the developing brain and spinal cord
3. In the case of unmyelinated fibres, there is from the surrounding mesoderm.
no elongation of the mesaxon. Several The development of the pia mater and the
unmyelinated fibres may invaginate the arachnoid mater {leptomeninges) is not definitely
same Schwann cell (Fig. 7.26). understood. According to some workers, these are
derived from the neural crest. The dura mater
Nerve fibres in different parts of the brain, and develops from the mesoderm surrounding the
spinal cord, become myelinated at different stages neural tube.
Chapter

8 The Skin and its Appendages

HIGHLIGHTS
• The epidermis is derived from surface ectoderm.
• The dermis is formed by mesenchyme derived
from dermatomes of somites.
• Naib develop from ectoderm at the tip of each
digit. Later this ectoderm migrates to the dorsal
aspect.
• Hair are derived from surface ectoderm that is
modified to form hair follicles.
• Sebaceous glands (ectoderm) arise as diver-
ticula from hair follicles.
• Sweat glands develop as downgrowths from
the epidermis, which are later canalized.
• Mammary glands arise from surface ectoderm.
They are formed along a milk line extending
from axilla to the inguinal region.
SKIN

The skin is derived from three diverse components


(Fig. 8.1).

(a) The epidermis is derived from the surface


ectoderm. This is, at first, single-layered
(Fig. 8.2A). The ectodermal cells proliferate
(multiply) to give rise to typical stratified
s q u a m o u s epithelium (Figs 8.2B, C, D).
Many of the superficial layers are shed off.
These get mixed up with secretions of
sebaceous glands to form a whitish sticky
substance {vernix caseosa) which covers the
skin of the newborn infant (Fig. 8.2D). The
vernix caseosa has a protective function.
The Skin and its Appendages

Surface ectoderm
forms epidermis -

Neural crest
gives rise to
dendritic cells.

Dermatome of somite
forms the dermis.—

Myotome

Fig. 8.1 Derivation of components of the skin.

(b) Epidermal ridges develop between


the third and fifth months of fetal
age. Soon, thereafter, characteristic TrTrrrTryryrTTrrrirT^rr^ Ectoderm
patterns (whorls, loop and arch)
•' •'"' ' — Mesenchyme
are formed on the tips of fingers
and toes. The p a t t e r n s are
genetically d e t e r m i n e d a n d are
different for each person.
(c) T h e melanoblasts (or dendritic
cells) of the epidermis are derived
from the neural crest. These cells Ectodermal cells
multiply to form
synthesize melanin pigments several layers
r e s p o n s i b l e for s k i n a n d h a i r
colour.
(d) Cells ofMerket and of Langerhans
appear in the epidermis between
8 a n d 12 weeks of intrauterine
life.
(e) T h e dermis is f o r m e d by
condensation and differentiation of
mesenchyme underlying the surface , Vemix caseosa
e c t o d e r m . T h i s m e s e n c h y m e is __ „ Stratum comeum
believed to be derived from the == * - Stratum lucidum
dermatome of somites. •*- Stratum granulosum
Stratum spinosum
The line of junction between dermis
a n d e p i d e r m i s is a t first s t r a i g h t . Germinal layer
Subsequently, the epidermis shows
regularly spaced thickenings that project Dermal papilla
into the dermis. The portions of dermis
intervening between these projections
form the dermal papillae. Still later, Fig. 8.2 Stages in the development of the epidermis.
Human Embryology

surface elevations {epidermal Stratum corneum


ridges) are formed by further Stratum lucidum
thickening of the epidermis in the
same situation.
Nail substance
—A
NAILS
Germinal layer
The nails develop from the surface
ectoderm. The ectoderm at the tip
of each digit becomes thickened to
form a primary nail field.
Subsequently this thickening
migrates from the tip of the digit
onto its dorsal aspect. Germinal matrix
The cells in the most proximal
part of the nail field proliferate to Fig. 8.3 Derivation of a nail.

Epidermis

Sebaceous
gland

. Developing
hair
- Papilla

Germinal
layer

-Bud for
sebaceous gland

- Papilla
-Papilla E

Fig. 8.4 Development of a hair follicle.


The Skin and its Appendages

form the root of the nail. Here the cells of the SEBACEOUS GLANDS
germinal layer multiply to form a thick layer of
cells called the germinal matrix. As the cells in A sebaceous gland is formed as a bud arising from
this matrix multiply, they are transformed into ectodermal cells forming the wall of a hair follicle.
the nail suhstance which corresponds to the stratum Some stages in the formation of a sebaceous gland
lucidum of the skin (Fig. 8.3). are shown in Figs 8.4C to E.
The migration of the primary nail fields from
the tips of the digits to their dorsal aspect explains SWEAT GLANDS
why the skin of the dorsal aspect of the terminal
part of the digits is supplied by nerves of the ventral A sweat gland develops as a downgrowth from
aspect. the epidermis (Fig. 8.5A). The downgrowth is at
first solid but is later canalized. The lower end of
HAIR the downgrowth becomes coiled (Fig. 8.5B} and
forms the secretory part of the gland.
The hair are also derived from surface ectoderm.
At the site where a hair follicle is to form, the
Anomalies of Skin and its Appendages
germinal layer of the epidermis proliferates to form
a cylindrical mass that grows down into the dermis 1. Albinism: Absence of pigment in skin, hair
(Figs 8.4A, B). The lower end of this downgrowth and eyes occurs because melanocytes are
becomes expanded and is invaginated by a unable to synthesize melanin. In this
condensation of mesoderm, which forms the autosomal recessive genetic condition, skin
papilla (Figs 8.4C, D). The hair itself is formed is depigmented all over the body. This
by proliferation of germinal cells overlying the should be distinguished from vitiligo
papilla. As the hair grows to the surface, the cells which is not congenital. In vitiligo the
forming the wall of the downgrowth surround it absence of pigment is patchy. In the
and form the epithelial root sheath. An additional affected areas there is degeneration of
dermal root sheath is formed from the surrounding already existing melanocytes. It is an
mesenchymal cells. A thin band of smooth muscle autoimmune disease.
{arrector pili ) is formed by mesodermal cells. It 2. Aplasia: The skin may fail to develop in
gets attached to the dermal root sheath. A typical certain regions.
hair follicle is thus formed (Fig. 8.4E).

_
MO
I
for sweat gland _ ,

secretory part
,pL B
er
Fig. 8.5 Development of a sweat gland.
Human Embryology

Dysplasia: The skin may be abnormal in


structure. Numerous varieties of dysplasia
are described. There may be congenital
growths of the skin. Dysplasia may be part
of maldevelopment of various ectodermal
derivatives including hair, teeth, sweat
glands and sebaceous glands.
Hair may be absent over the scalp
(congenital alopecia). The eyebrows and
eyelashes may also be absent. Absence of
hair in any part of the body is called
atrichia, while overgrowth of hair is called
hypertrichosis.
Anonychia: Nails may be absent. Occa-
sionally they may show overdevelopment.

MAMMARY GLANDS
In some animals (e.g. bitches} a series of
mammary glands are present on either side of the
midline, on the ventral surface of the trunk. These
are situated along a line that extends from the
Fig. 8.6 The mammary line passing from the
axilla to the inguinal region. In the human embryo,
axilla t o the inguinal region.
the ectoderm becomes thickened along this line to
form mammary ridges or lines
(Fig. 8.6). Most of this line soon
disappears. Each mammary gland
develops from a part of this line
that overlies the pectoral region.
In the region where the
mammary gland is to form, a
thickened mass of epidermal cells
is seen projecting into the dermis
(Fig. 8.7A). From this thickened
mass, sixteen to twenty solid
outgrowths arise and grow into the
B
W
surrounding dermis (Figs 8.7B, C).
The thickened mass of epidermis,
as well as these outgrowths, are
now canalized (Fig. 8.7D). The
secretory elements of the gland are
formed by proliferation of the
terminal parts of the outgrowths.
The proximal end of each
outgrowth forms one lactiferous Fig. 8.7 Stages in the development of the mammary gland.
The Skin and its Appendages

duct. The ducts at first open into a pit formed by opening into a pit. This causes difficulty
cavitation of the original epithelial thickening. in suckling.
However, the growth of underlying mesoderm 6. The gland may be abnormally small
progressively pushes the wall of this pit outwards, {micromastia) or abnormally large
until it becomes elevated above the surface and (macromastia).
forms the nipple (Fig. 8.7E). The mammary gland 7. Gynaecomastia: The male breast may
remains rudimentary in the male. In females, the enlarge as in the normal female and may
ducts and secretory elements undergo extensive even secrete milk.
development during puberty and pregnancy.
TIMETABLE OF SOME EVENTS
Developmental Anomalies of tin MENTIONED IN THIS CHAPTER
Mammary Glands
Age Developmental Events
1. Amostia: The gland may be absent on one
or both sides. 7th week Mammary line is
2. Athelia: The nipple may be absent. established.
3. Polythelia and polymastia: Supernume- 8th week Melanoblasts start
rary nipples may be present anywhere appearing.
along the milk line. They may remain 1st to 3rd month Cells of neural crest
rudimentary (polythelia) or may form migrate to skin.
accessory mammary glands (polymastia). 2nd month Surface ectoderm is single
4. Accessory breasts may be found away from layered.
the milk line. They have been observed in 2nd to 4th month Surface ectoderm becomes
the neck, cheeks, femoral triangle and
multiple layered.
vulva.
3rd to 4th month Dermal papillae are
5. Inverted or crater nipple: The nipple may
fail to form resulting in lactiferous ducts formed.
The Pharyngeal Arches

HIGHLIGHTS • The cartilages of the fourth and sixth arches


give rise to the cartilages of the larynx.
• Pharyngeal arches are rod-like thickenings of
• The nerves of the pharyngeal arches are as
mesoderm present in the wall of the foregut.
follows: First arch = mandibular; second arch
• At first there are six arches. The fifth arch = facial; third arch = glossopharyngeal-, fourth
disappears and only five remain. a r c h = superior laryngeal; sixth a r c h =
• The ventral ends of the arches of the right and recurrent laryngeal. The muscles supplied by
left sides meet in the middle line in the floor of these nerves are derived from the mesoderm of
the pharynx. the arch concerned.
• In the interval between any two arches, the • The external acoustic meatus develops from
e n d o d e r m (lining the p h a r y n x ) is pushed the first ectodermal cleft.
outwards to form a series of pouches. These • T h e first e n d o d e r m a l p o u c h (and p a r t of
are called endodermal, or pharyngeal pouches. second) gives off a diverticulum called the
• Opposite each pouch the surface ectoderm dips tubotympanic recess. The middle ear and the
inwards as an ectodermal cleft. auditory tube develop from the tubotympanic
recess.
• Each p h a r y n g e a l a r c h c o n t a i n s a skeletal
element (cartilage that may later form bone), • The palatine tonsil arises from the second
striated muscle supplied by the nerve of the pouch.
arch and an arterial arch. • The inferior parathyroid gland and the thymus
• T h e c a r t i l a g e of the first a r c h {Meckel's are derived from the third pouch.
cartilage) gives origin to the incus and malleus • The superior parathyroid gland is derived from
(of middle ear). the fourth pouch.
• The cartilage of the second arch forms the stapes, • The thyroid gland develops mainly from the
the styloid process and part of the hyoid bone. thyroglossal duct. This duct is formed as a
• The cartilage of the third arch forms the greater median diverticulum arising from the floor of
part of the hyoid bone. the pharynx (at the foramen caecum).
The Pharyngeal Arches

d e s t i n e d t o form t h e p h a r y n x ) , b e f o r e t h e
INTRODUCTION
appearance of the pharyngeal arches, is shown in
The formation of the foregut has been considered Fig. 9.2A. At this stage, the endodermal wall of
in Chapter 5. Reference to Fig. 5.14 will show the foregut is separated from the surface ectoderm
that after the establishment of the head fold, the by a layer of mesoderm. Soon, thereafter, the
foregut is bounded ventrally by the pericardium, mesoderm comes to be arranged in the form of six
and dorsally by the developing brain. Cranially, bars that run dorsoventrally in the side wall of the
it is at first separated from the stomatodaeum by foregut. Each of these 'bars' grows ventrally in
the b u c c o p h a r y n g e a l m e m b r a n e . W h e n this the floor of the developing pharynx and fuses with
membrane breaks down, the foregut opens to the the corresponding 'bar' of the opposite side to form
exterior through the stomatodaeum. a pharyngeal or branchial arch. In the interval
At this stage, the head is represented by the between any two adjoining arches, the endoderm
bulging caused by the developing brain (Fig. 5.14), e x t e n d s o u t w a r d s in t h e form of a p o u c h
while the p e r i c a r d i u m may be considered as {endodermal or pharyngeal pouch) to meet the
occupying the region of the future thorax. The ectoderm which dips into this interval as an
two are separated by the stomatodaeum which is ectodermal cleft (Fig. 9.2B).
the future mouth. It is, thus, apparent that a neck The first arch is also called the mandibular arch;
is not yet present. and the second, the byoid arch. The third, fourth
The neck is formed by the elongation of the and sixth arches do not have special names. The
region between the stomatodaeum and the pericar- fifth arch disappears soon after its formation, so
dium. This is achieved, partly, by a 'descent' of that only five arches remain.
the developing heart. 1 lowcver, this elongation is T h e following structures are formed in the
due mainly t o the a p p e a r a n c e of a series of mesoderm of each arch (Fig. 9.3):
mesodermal thickenings in the wall of the cranial-
most part of the foregut. These are called the I. A skeletal element: This is cartilaginous to
pharyngeal or branchial arches (see Fig. 9.1). begin with. It may remain cartilaginous,
A coronal section through the foregut (the part may develop into bone, or may disappear.

Bulging produced by
developing brain Pharyngeal arches
Pharyngeal arches

Stomal od aeum'J

Hindlimb
bud
Foreiimb
Pericardial bulge
bud

Fig. 9.1 Lateral views of embryos showing the formation of pharyngeal arches between stomatodaeum
and the pericardial bulge.
Human Embryology

Buccopharyngeal
membrane

Ectoderm (blue)

Primitive pharynx
derived from Endoderm (red)
foregut

Mesoderm (green)

Buccopharyngeal membrane
disappears.

(5th arch disappears


soon after formation.)

Fig. 9.2 Coronal sections through cranial part of foregut. (A) Before, and (B) After
formation of pharyngeal arches.

2. Striated muscle: This is supplied by the nerve with them, and their embryological
nerve of the arch. In later development, this origin can thus be determined from their
musculature may, or may not, retain its nerve supply.
attachment to the skeletal elements derived An arterial arch: Ventral to the foregut, an
from rhe arch. It may subdivide to form a artery called the ventral aorta develops.
number of distinct muscles, which may Dorsal to the foregut, another artery called
migrate away from the pharyngeal region. the dorsal aorta, is formed. A series of
When they do so, however, they carry their arterial arches [aortic arches) connect the
The Pharyngeal Arches

ventral and dorsal aortae. One such arterial In some lower animals, each arch is supplied
a r c h lies in e a c h p h a r y n g e a l a r c h . In by t w o nerves (Fig. 9.4). The nerve of the arch
subsequent development, the arrangement itself runs along the cranial border of the arch.
of these arteries become greatly modified. This is called the post-trematic nerve of the arch
The fate of the arterial arches is considered (trema • trench). Each arch also receives a
in Chapter 15. branch from the nerve of the succeeding arch.
This runs along the caudal border of the arch,
Each pharyngeal arch is supplied by a nerve. and is called the pre-trematic nerve of the arch.
In addition to supplying the skeletal muscle of the In the h u m a n e m b r y o , however, a d o u b l e
arch, it supplies sensory branches to the overlying i n n e r v a t i o n is to be seen only in the first
ectoderm and endoderm (Fig. 9.3). pharyngeal arch.

DERIVATIVES OF THE
SKELETAL ELEMENTS
1. T h e cartilage of the first arch is called
Meckel's cartilage (Fig. 9.5). The incus wad
malleus (of the middle ear) are derived from
its dorsal e n d . T h e ventral p a r t of the
cartilage is surrounded by the developing
mandible and is absorbed. The part of the
cartilage extending from the region of the
middle ear t o the mandible disappears, but
its sheath (perichondrium) forms the anterior
ligament of the malleus and the
sphenomandibular ligament.
M e s e n c h y m e of the first arch is also
responsible for formation of bones of the
face including the maxilla, the mandible,
the zygomatic bone, the palatine bone and
Fig. 9.3 Structures to be seen in a pharyngeal part of the temporal bone. (Also see first
arch. arch syndrome.)

Nerve I
Pre-trematic branch
" Nerve II
Post-trematic branch
Nerve III

Fig. 9.4 Arrangement of nerves supplying the pharyngeal arches in some lower animals.
Human Embryology

Spine of sphenoid

Stapes

Malleus

Styloid process

Sphenomandibular
ligament

Remnants of
Meckel's cartilage
within mandible

Greater cornu

From cartilage Lesser cornu


of third arch

Body of hyoid bone

From cartilages of Thyroid cartilage


fourth and sixth arches

Cricoid cartilage

Fig. 9.5 Fate of the cartilages of the pharyngeal arches. The left half of the figure
shows an earlier stage of development. The derivatives of the first arch
are shown in red, second arch in green, third arch in orange, and
fourth, fifth and sixth arches in blue.

T h e cartilage of the second arch forms the (e) Superior parr of body of hyoid bone.
following: (Note that all structures listed start with 'S').
(a) Stapes. The following structures are formed from
(b) Styloid process. the cartilage of the third arch:
(c) Stylohyoid ligament (from sheath). (a) Greater Cornu of hyoid bone.
(d) Smaller (lesser) cornu of hyoid bone. (b) Lower part of the body of hyoid bone.
The Pharyngeal Arches

4. The cartilages of the larynx are derived from while the chorda tympani (branch of facial nerve)
the fourth and sixth arches with a possible is the pre-trematic nerve. This double innervation
contribution from the fifth arch, but their is reflected in the nerve supply of the anterior two-
exact derivation is controversial. third of the tongue which is derived from the ventral
part of the first arch.
NERVES AND MUSCLES
OF THE ARCHES Some recent investigations suggest that
mesenchyme giving rise to muscles of the
All the muscles derived from a pharyngeal arch pharyngeal arches is derived from paraxial
are supplied by the nerve of the arch and can, mesoderm that is cranial to the occipital somites
therefore, be identified by their nerve supply. The (i.e. from the region of the pre-occipital
nerves of the arches and the muscles supplied by somites); and that its organization is influenced
them are given in Table 9.1. by neural crest cells. Although paraxial
mesoderm here does not form typical somites,
Table 9.1 Nerves of pharyngeal arches and it shows partial segmentation into seven masses
muscles supplied by them of mesenchyme called somitotneres. The
structures derived from the seven somitomeres
Arch Nerve of Arch Muscles of Arch and from five occipital somites that follow them,
Mandibular Medial and lateral have been described as given in Table 9.2.
pterygoids, Masseter,
Temporalis,
Mylohyoid, Anterior Table 9.2 Muscles derived from somitomeres and
belly of digastric, somites
Tensor tympani,
Tensorpalati. Somitomere/Somites Muscles Derived

Second Facial Muscles of face, Somitomere 1 and 2 Muscles supplied by


Occipitofrontalis, o c u l o m o t o r nerve
Platysma, Stylohyoid, Somitomere 3 Superior o b l i q u e muscle
Posterior belly of supplied by trochlear nerve
digastric, Stapedius, Muscles of first pharyngeal
Somitomere 4
Auricular muscles.
arch supplied by mandibular
Third Closso- Stylopharyngeus nerve
pharyngeal Somitomere 5 Lateral rectus muscle
Fourth Superior supplied by abducent nerve
laryngeal Muscles of larynx and Somitomere 6 Muscles of the second
Sixth Recurrent pharynx pharyngeal arch supplied by
laryngeal the facial nerve
Somitomere 7 Stylopharyngeus (from 3rd
We have already seen that these nerves not only arch) supplied by
supply muscles, but also innervate the parts of glossopharyngeal nerve
skin and mucous membrane derived from the
O c c i p i t a l somites Laryngeal muscles (from 4th
arches. Some of the nerves (e.g. glossopharyngeal) 1 and 2 to 6th arches) supplied by
have only a small motor component and are the vagus nerve
predominantly sensory. As stated above, the first
arch has a double nerve supply. The mandibular O c c i p i t a l somites Muscles of tongue supplied
3 to 5 by hypoglossal nerve
nerve is the post-trematic nerve of the first arch,
Human Embryology

If we accept this view of the origin of branchial swellings that lie in the neck, along the anterior
m u s c u l a t u r e , there would be n o significant border of the sternocleidomastoid. These are called
reason to distinguish between it and muscle branchial cysts, and are most commonly located
derived from somites. just below the angle of the mandible. If such a
cyst opens onto the surface, it becomes a branchial
sinus. Rarely, a cervical sinus may open into the
FATE OF ECTODERMAL CLEFTS lumen of the pharynx in the region of the tonsil.
After the formation of the pharyngeal arches, the
region of the neck is marked on the outside by a FATE OF ENDODERMAL POUCHES
series of grooves or ectodermal clefts. T h e dorsal
part of the first cleft (between the first and second T h e e n d o d e r m a l p o u c h e s t a k e p a r t in t h e
arches) develops into the epithelial lining of the formation of several important organs (Fig. 9.7).
external acoustic meatus. The pinna (or auricle) These are listed below.
is formed from a series of swellings or hillocks,
that arise on the first and second arches, where First Pouch
they adjoin the first cleft (for the development of (a) Its ventral part is obliterated by formation
the pinna refer to Chapter 19). T h e ventral parr of of the tongue.
this cleft is obliterated. (b) Its dorsal part receives a contribution from
The second arch grows much faster than the the dorsal part of the second pouch, and
succeeding arches and comes to overhang them these two together form a diverticulum that
(Fig. 9.6). T h e space between the overhanging grows t o w a r d s the region of the developing
second arch and the third, fourth and sixth arches ear. This diverticulum is called the tubo-
is called the cervical sinus. Subsequently, the lower tympanic recess. The proximal part of this
overhanging border of the second arch fuses with r e c e s s g i v e s rise t o t h e auditory
tissues caudal to the arches. The side of the neck (pharyngotympanic) tube and the distal part
(which was so far marked by the ectodermal clefts) to the middle ear cavity, including the
now becomes smooth. The cavity of the cervical tympanic antrum.
sinus (which is lined by ectoderm) normally gets
obliterated. Part of it may persist and give rise to Second Pouch
(a) T h e e p i t h e l i u m of t h e
ventral part of this pouch
contributes to the forma-
tion of the tonsil.
(b) The dorsal part takes part
Cervical sinus in the formation of the
lying deep to tubotympanic recess.
downward projection
Projection growing
downwards from Third Pouch
second arch
This gives rise to the inferior
parathyroid glands, a n d the
thymus.

Fourth Pouch
Fig. 9.6 Cervical sinus. The left half of the figure shows an earlier
stage than the right half. This gives origin to the superior
The Pharyngeal Arches

Tubotympanic recess

Tonsil

Parathyroid II

Parathyroid IV

Lateral thyroid (?)

Thymus

Fig. 9.7 Scheme to show the fate of the pharyngeal pouches (numbered 1 to 4).

parathyroid glands, and may contribute to the Early in development, this pouch is cut off, both
thyroid gland. from the pharyngeal wall and from the surface
ectoderm.
Fifth or Ultimobranchial Pouch After separation from the inferior parathyroid
A fifth pouch is seen for a brief period during rudiment, each thymic rudiment has a thinner
development. In some species it gives rise to the cranial part and a broader caudal part. The thinner
ultimobranchial body. Its fate in m a n is portion forms the cervical part of the thymus. The
c o n t r o v e r s i a l . It is generally believed t o be broader parts, of the two sides, enter the thorax and
i n c o r p o r a t e d into the fourth p o u c h , the t w o become united to each other by connective tissue.
together forming the caudal pharyngeal complex. Theendodermal cells of the thymus are invaded
The superior parathyroid glands arise from this by vascular mesoderm which contains numerous
complex. The complex probably also gives origin lymphoblasts. This invading mesenchyme partially
to the parafollicular cells of the thyroid gland. breaks up the thymic tissue into isolated masses,
and thus gives the organ its lobulated appearance.
DEVELOPMENT OF THE THYMUS Fragmentation of the cervical part of the thymus
may give rise to accessory thymic tissue. Such
The thymus develops from the endoderm of the t i s s u e , p r e s e n t in r e l a t i o n t o t h e s u p e r i o r
third pharyngeal pouch (which also gives rise to parathyroid glands, is believed to arise from the
the inferior parathyroid glands). fourth pouch.
Human Embryology

T h e t h y m u s is relatively large at b i r t h . It In keeping with their developmental history,


c o n t i n u e s t o increase in w e i g h t till p u b e r t y . the superior p a r a t h y r o i d glands are relatively
Thereafter, it gradually undergoes atrophy. constant in position, but the inferior parathyroid
glands may descend into the lower part of the neck
or even i n t o t h e a n t e r i o r m e d i a s t i n u m .
DEVELOPMENT OF PARATHYROID
Alternatively, they may remain at their site of
GLANDS
origin and are then seen near the bifurcation of
Parathyroid glands are derived as follows: the common carotid artery.

1. The inferior parathyroid glands develop from DEVELOPMENT OF THE


endoderm of the third pharyngeal pouch THYROID GLAND
(parathyroid III).
2. The superior parathyroid glands develop The thyroid gland develops mainly from the
from endoderm of the fourth pharyngeal thyroglossal duct. Parafollicular cells are derived
pouch (parathyroid IV). from the caudal pharyngeal complex {derived from
the fourth and fifth pharyngeal pouches).
As the third pouch also gives origin to the After the formation of the pharyngeal arches,
thymus, this organ is closely related to parathyroid the floor of the pharynx has the appearance as
111. When the thymus descends towards the thorax, shown in Fig. 9.9. The medial ends of the two
parathyroid 111 is carried caudally along with it m a n d i b u l a r arches are separated by a midline
for some distance. Meanwhile, parathyroid IV is swelling called the tuherculutn impar. Immediately
prevented from descending caudally, because of behind the tuberculum, the epithelium of the floor
the close relationship of the fourth pouch to the of the pharynx shows a thickening in the middle
developing thyroid gland. As a result, parathyroid line (Fig. 9.10A). This region is soon depressed
III becomes caudal to parathyroid IV. Hence, the below the surface to form a diverticulum called
parathyroid glands derived from the fourth pouch the thyroglossal duct (Fig. 9.10B).
become the superior parathyroid glands and those T h e site of origin of the diverticulum is n o w
derived from the third pouch become the inferior seen as a depression called the foramen caecum.
parathyroid glands (Fig. 9.8). The diverticulum grows down in the midline into

Thyroid
gland

. Parathyroid III

Fourth pouch

Parathyroid IV

Fig. 9.8 Derivation of superior and inferior parathyroid glands. Note that the
relative position of parathyroid III and IV is reversed during development.
The Pharyngeal Arches

represent the ultimobranchial


Tuberculum impar
body of lower animals.
Lingual swelling
Anomalies of the
X i
First arch *-t. :J D!O Thyroid Gland
A. Anomalies of Shape
1. The pyramidal lobe is
Second arch »• t ' " ) ^ * ^ / ~ , Foramen caecum present so often that it
"'/ (blue dot)
is regarded as a normal
Third arch *• y T"""" Hypobranchial structure. It may arise
/ eminence from the isthmus
(Fig. 9.11A) o r from

YwJ
o n e of t h e l o b e s
Fourth arch •^^^
(Figs9.11B,C).Itmay
Fifth arch has
disappeared.

Sixth arch
\ \ II o ^ Tracheal groove
have no connection
w i t h t h e rest of t h e
thyroid, and may
A / be divided into two or
Fig. 9.9 Floor of the pharynx showing the foramen caecum from \ more parts (Fig. 9.1 ID).
the thyroglossal duct arises. In extent, it may vary
from a s h o r t s t u m p
Site of foramen caecum (Fig. 9.11A) to a pro-
cess reaching the hyoid
Floor of bone (Fig. 9.11C).
pharynx Thyroglossal duct
2. T h e i s t h m u s m a y be
absent (Fig. 9.12A).

h- Thyroglossal
duct
Lateral thyroid (?)
3. O n e of the lobes of the
g l a n d m a y be v e r y
small (Fig. 9.12B), or
from 4th pouch
absent (Fig. 9.12C).
Thyroid developing
from thyroglossal duct B. Anomalies of Position
(Fig. 9.13)
Bifid lower end

^r 1. Lingual thyroid: T h e
thyroid may lie under
Fig. 9.10 Stages in the development of the thyroid gland.
t h e m u c o s a of t h e
the neck. Its tip soon bifurcates (Fig. 9.IOC). d o r s u m of the t o n g u e a n d m a y form a
Proliferation of the cells of this bifid end gives s w e l l i n g t h a t m a y c a u s e difficulty in
rise to the t w o lobes of the thyroid gland. swallowing.
The developing thyroid comes into intimate Intra-lingual thyroid: The thyroid may be
relationship with the caudal pharyngeal complex embedded in the muscular substance of the
and fuses with it (Fig. 9.10D). Cells arising from tongue.
this complex are believed to give origin to the Suprahyoid thyroid: The gland may lie in
parafollicular cells of the thyroid which may the midline of the neck, above the hyoid bone.
Human Embryology

Fig. 9.11 Variations in the pyramidal process Fig. 9.12 Anomalies of the thyroid gland. The
of the thyroid gland. parts of the gland shown in dotted
outline are missing.
4. Infrahyoid thyroid: The gland may lie below
the hyoid bone, but above its normal Tongue
position. Lingual thyroid

5. Intrathoracic thyroid: The entire gland, or


part of it, may lie in the thorax.
Intralinguai thyroid
Note that when thyroid tissue is present in the
anomalous positions described above, an
J i _— Suprahyoid thyroid
additional thyroid may or may not be present at
the normal sire.
Infrahyoid thyroid

C. Ectopic Thyroid Tissue Hyoid bone

Small masses of thyroid tissue may be present at Thyroid cartilage


abnormal sites. Thyroid tissue has been observed
in the larynx, trachea, oesophagus, pons, pleura, Path of thyroglossal duct
pericardium and ovaries. Masses of ectopic thyroid Normal position of thyroid
tissue have been described in relation to the deep
cervical lymph nodes {lateral aberrant thyroids)
but these are now believed to represent metastases Fig. 9.13 Path of thyroglossal duct. Note that
in the lymph nodes from a carcinoma of the thyroid thyroid tissue may come to lie anywhere
gland. along the course of this duct.
The Pharyngeal Arches

D. Remnants of the Thyroglossal Duct to be remembered that the duct is intimately related
to the hyoid bone (Fig. 9.13).
These r e m n a n t s m a y persist a n d lead t o t h e
formation of the following:
TIMETABLE O F S O M E EVENTS
1. Thyroglossal cysts, that may occur M E N T I O N E D IN THIS C H A P T E R
anywhere along the course of the duct. They
Age Developmental Events
may acquire secondary openings o n the
surface of the neck t o form fistulae. 4th week Appearance of 1st and 2nd
2. Thyroglossal fistula opening at the foramen (22nd day) arches.
caecum. 5th week Four arches are seen.
3. Carcinoma of the thyroglossal duct. (29th day) Thyroid, parathyroid and
thymus start forming.
In the surgical removal of thyroglossal cysts
7th week Thyroid gland reaches its
and fistulae, it is important to remove all remnants
of the thyroglossal duct. In this connection, it has definitive position.
The Skeleton

HIGHLIGHTS
• T h e vertebral column is derived from the
sclerotomes of somites. Each sclerotome divides
into three parts: cranial, middle and caudal.
• A vertebra is formed by fusion of the caudal
part of one sclerotome and the cranial part of
the n e x t s c l e r o t o m e . It is, t h e r e f o r e ,
intersegmental in position.
• The middle part of the sclerotome forms an
intervertebral disc, w h i c h is t h e r e f o r e
segmental in position.
• The sternum is formed by fusion of right and
left sternal bars.
• The skull develops from mesenchyme around
the developing brain. Some skull bones are
formed in m e m b r a n e (e.g. p a r i e t a l ) ; some
partly in membrane and partly in cartilage
(e.g. sphenoid); and a few entirely in cartilage The process of bone formation has been considered
(e.g. ethmoid). in Chapter 7. We have seen that all bone is of
m e s o d e r m a l o r i g i n , a n d t h a t b o n e s c a n be
• The mandible is formed in membrane from the classified as cartilage bones or membrane bones,
mesenchyme of the mandibular process.
on the basis of their mode of ossification. We shall
• Limbs are first seen as outgrowths {limb buds) now consider the development of some individual
from the side wall of the embryo. Each bud parts of the skeleton.
grows and gets subdivided to form parts of the
I i m b.
THE VERTEBRAL COLUMN
• Limb bones develop from mesenchyme of the
limb b u d s . Joints are formed in intervals T h e v e r t e b r a l c o l u m n is f o r m e d f r o m the
between bone ends. sclerotomes of the somites.

The cells of each sclerotome get converted into


loose mesenchyme. This mesenchyme migrates
medially and surrounds the notochord
The Skeleton

Neural tube Mesenchymal basis


of neural arch ..-: :

I 1 /"^Y Somite — Neural tube

fe
Mesenchyme M ; . • ; v7 'Sclerotome
from
sclerotome Notochord '7
Mesenchymal basis - Notochord
of costal element
Fig. 10.1 Formation of mesenchymal basis of the
body of a vertebra from a sclerotome.
Mesenchymal basis
of vertebra! body
(Fig. 10.1). The mesenchyme then extends
backwards on either side of the neural tube and Fig. 10.2 Formation of mesenchymal basis of the
surrounds it (Fig. 10.2). Extensions of this neural arch and of the costal element.
mesenchyme also take place laterally in the
position to be subsequently occupied by the (Fig. 10.3A). The mesenchymal cells of each
transverse processes, and ventrally in the body segment are at first uniformly distributed.
wall, in the position to be occupied by the ribs. However, the cells soon become condensed in a
region that runs transversely across the middle of
For some time the mesenchyme derived from the segment. This condensed region is called the
each somite can be seen as a distinct segment perichordal disc. Above and below it there are

£ ^Vertebra

Intervertebral
disc

. Nucleus
pulposus

Caudal less t
condensed part

Fig. 10.3 (A) Mesenchyme derived f r o m somites is seen in the form of segments. (B) Each segment has a
central condensed part, and cranial and caudal less condensed parts. (C) A vertebra is formed by
fusion of adjoining uncondensed parts of t w o somites. Hence it is an intersegmental structure.
Each intervertebral disc is derived from the condensed part of one somite. Hence it is segmental
in position.
Human Embryology

less condensed parts (Fig. 10.3B). The mesenchy- neural arch and one for the greater part of the
mal basis of the body (or centrum) of each vertebra body (centrum). At birth the centrum and the
is f o r m e d by fusion of t h e a d j o i n i n g , less two halves of the neural arch are joined by
condensed parts of two segments (Fig. 10.3B). The c a r t i l a g e (Fig. 1 0 . 4 A ) . T h e s e a r e t e r m e d
perichordal disc becomes the intervertebral disc. neurocentral joints. Note that the posterolateral
parts of the vertebral body are formed from the
The neural arch, the transverse processes and neural arch (Fig. 10.4B). After the centrum and
the costal elements are formed in the same way neural arch have fused, the junction between
as the body. T h e interspinous and intertrans- the t w o is indicated by the neurocentral line.
verse ligaments are formed in the same manner
as the intervertebral disc.
Congenital Anomalies of
T h e notochord disappears in the region of
Vertebral Column
the vertebral bodies. In the region of the inter-
vertebral discs, the notochord becomes expand- O n e or more vertebrae may be absent, the
ed and forms the nucleuspulposus (Fig. 10.3C). caudal vertebrae being more commonly
affected. Absence of the coccyx alone, or
From the above account we may note that: of the sacrum and coccyx, may be seen.
Additional vertebrae may be present. The
1. T h e vertebra is an intersegmental structure sacrum may show six segments.
made up from portions of two somites. The Part of a vertebra may be missing. This
position of the centre of the somite is m a y r e s u l t in v a r i o u s a n o m a l i e s ,
represented by the intervertebral disc. depending on the part t h a t is absent.
2. The transverse processes and ribs are also
intersegmental. They separate the muscles (a) The two halves of the neural arch may
derived from two adjoining myotomes. fail t o fuse in t h e m i d l i n e . T h i s
3. Spinal nerves are segmental s t r u c t u r e s . c o n d i t i o n is c a l l e d spina bifida
They, therefore, emerge from between two (Fig. 10.5). T h e g a p b e t w e e n the
adjacent v e r t e b r a e and lie
between t w o adjacent ribs.
Part of vertebral body
4. The blood vessels supplying formed by neural arch
structures derived from the
m y o t o m e (e.g. i n t e r c o s t a l
vessels) are intersegmental
like the vertebrae. Therefore,
the intercostal and l u m b a r
a r t e r i e s He o p p o s i t e t h e
vertebral bodies.

T h e m e s e n c h y m a l basis of t h e
vertebra is converted into cartilage Neurocentral line
by t h e a p p e a r a n c e of s e v e r a l
centres of chondrification. Three Fig. 10.4 (A) A vertebra at birth consisting of three separate pieces
p r i m a r y centres of ossification of bone: a centrum and t w o neural arches. (B) Diagram
appear in the cartilaginous model to show the neurocentral line w h i c h is the line along
for each vertebra: one for each w h i c h body and neural arch have fused.
The Skeleton

fusion may occur in the cervical region


{Klippel-Feil syndrome). The atlas
vertebra may be fused to the occipital bone
(occipitilization of atlas). The fifth lumbar
vertebra may be partially or completely
fused to the sacrum {sacralization of 5th
lumbar vertebra).
5. Parts of the vertebral column t h a t are
n o r m a l l y fused t o each o t h e r may be
separate. The first sacral vertebra may be
s e p a r a t e from the rest of the s a c r u m
(lumbarization of the first sacral vertebra).
Fig. 10.5 Spina bifida produced by non-fusion of
The odontoid process may be separate
the t w o halves of the neural arch.
from the rest of the axis vertebra.
6. The articular facets may be abnormal in
neural arches may not be o b v i o u s orientation, or may be deficient. When
(spina bifida occulta), or may be large there is deficiency of both the inferior
e n o u g h for m e n i n g e s a n d n e u r a l a r t i c u l a r p r o c e s s e s of t h e fifth l u b a r
e l e m e n t s t o b u l g e o u t of it (see vertebra, the body of the vertebra may slip
meningocoele and meningo- forwards over the sacrum. This is called
myelocoele). spondylolisthesis.
Spina bifida in a fetus can be recog- 7. The vertebral canal may be divided into
nized by u l t r a s o u n d e x a m i n a t i o n . two lateral halves by a projecting shelf of
E x a m i n a t i o n of a m n i o t i c fluid b o n e , which splits the spinal cord
shows increased levels of alpha feto longitudinally into two halves
proteins (AFP) in a case with spina (diastematomylia).
bifida. 8. Ossification of the vertebral bodies may
(b) The vertebral body may ossify from be defective thus reducing the total length
two primary centres which soon fuse. of the spine. This can lead to the formation
One of these parts may fail to develop, of dwarfs w h o have a short trunk but have
resulting in only half of the body limbs of normal length (chondro-osteo-
b e i n g p r e s e n t . T h i s is c a l l e d dystrophy),
hemivertebra. It is usually associated 9. A peculiar t u m o u r arising from cells of
with absence of the corresponding rib.
the primitive knot may be seen attached
(c) The t w o halves of the vertebral body to the lower end of the spine. Various
may be formed normally but may fail tissues may be seen in it. Such a growth is
to fuse. T h e v e r t e b r a l b o d y t h e n called a sacrococcygeal teratoma.
c o n s i s t s of t w o h e m i v e r t e b r a e . A n o m a l i e s of t h e v e r t e b r a e are of
Sometimes the gap between the t w o
practical importance in that:
halves is large enough for meninges
and nerves to bulge forward between (a) They may cause deformities of the
them (anteriorspina bifida). spine. The spine may be bent on itself
(congenital scoliosis). Deformities of
4. Two or more vertebrae that are normally cervical vertebrae may lead to tilting
separate may be fused to each other. Such of the head to one side and its rotation
Human Embryology

to the opposite side (congenital either side of the midline. Mesenchymal


torticolis). This deformity may be condensations forming at these sites become
secondary to a contracture of the cartilaginous in the seventh week of intrauterine
sternocleidomastoid muscle. life. Laterally, the sternal bars are continuous with
(b) The spinal nerves, or even the spinal ribs. The fusion of the two sternal bars first occurs
cord, may be implicated. They may at their cranial end (manubrium) and then proceeds
be subjected to abnormal pressure caudally (Figs 10.7B, C).
leading to paralysis. The manubrium and the body of the sternum
(c) They are frequently the cause of are ossified, separately. The xiphoid process ossifies
backache. only late in life.

THE RIBS Anomalies of the Sternum and Ribs


1. Some ribs that are normally present may
The ribs are derived from ventral extensions of be missing. Unilateral absence of a rib is
the sclerotomic mesenchyme that forms the often associated with hemivertebra.
vertebral arches. These extensions are present not
2. Accessory ribs may be present. Such a rib
only in the thoracic region but also in the cervical,
may be attached to the seventh cervical
lumbar and sacral regions. They lie ventral to the
vertebra (cervical rib), or to the first
mesenchymal basis of the transverse processes with
lumbar vertebra (lumbar rib).
which they are continuous.
In the thoracic region, the entire
extension (called the primitive costal
arch) undergoes chondrification, and
subsequent ossification, to form the
ribs. However, some mesenchyme
between it and the developing
transverse process does not undergo
chondrification: it becomes loose and
forms the costotransverse joint. In the
cervical, lumbar and sacral regions,
chondrification and ossification of the
costal arch is confined to the region
in immediate relationship to the
transverse process. The bone formed
Rib
from the arch is fused to the transverse
Neurocentra) line
process and is referred to as the costal
element of the process. The contri-
butions made by the costal element to From neural arch
the cervical, lumbar and sacral
vertebrae are shown in Fig. 10.6.
From costal element

THE STERNUM Fig. 10.6 Relative contribution to vertebrae by the centrum, the
neural arch and the costal element in different
The sternum is formed by fusion of two regions. Note that a small part of the body of the
sternal bars, or plates, that develop on vertebra is derived from the neural arch.
The Skeleton

(b) The developing internal ear (otic vesicle),


and the region of the developing nose, are
surrounded by mesenchymal condensations
c a l l e d t h e otic a n d nasal capsules
respectively. These capsules also take part
in forming the mesenchymal basis of the
skull.
(c) The first branchial arch is closely related
to the developing skull. It soon shows two
subdivisions, called the mandibular and
Fig.10.7 Development of the sternum. (A) Sternal maxillary processes. Some bones of the skull
bars formed on each side of the middle a r e f o r m e d in t h e m e s o d e r m of these
line. (B) The sternal bars begin to fuse processes.
with each other at the cranial end.
(C) Fusion progresses caudally. S o m e b o n e s of t h e s k u l l a r e f o r m e d in
membrane, some in cartilage, and some partly in
3. When the fusion of the two sternal bars is membrane and partly in cartilage, as listed below.
faulty, the body of the sternum shows a
. partial or even a complete midline cleft. Bones that are Completely Formed in
Minor degrees of non-fusion may result in Membrane
a bifid x i p h o i d process or in midline
f o r a m i n a . T r a n s v e r s e clefts m a y a l s o (a) The frontal and parietal bones are formed
occur. in relation to mesenchyme covering the
4. In the condition called funnel chest, the developing brain.
lower part of the sternum and the attached (b) The maxilla (excluding the premaxilla),
ribs are d r a w n inwards into the thorax. zygomatic and palatine bones, and part of
The primary defect is that the central ten- t h e temporal b o n e s a r e f o r m e d by
don of the diaphragm is abnormally short. i n t r a m e m b r a n o u s o s s i f i c a t i o n of t h e
5. The upper part of the sternum (and related mesenchyme of the maxillary process.
costal cartilages) may project forwards (c) T h e nasal, lacrimal and vomer hones are
{pigeon breast). ossified in the membrane covering the nasal
capsule.

THE SKULL Bones that are Completely Formed in


Cartilage
T h e skull is developed from the mesenchyme
s u r r o u n d i n g the developing brain. This The ethmoid bone and the inferior nasal concha
mesenchyme comes into close relationship with are derived from the cartilage of the nasal capsule.
the following structures that also contribute to the The septal and alar cartilages of the nose represent
development of the skull. p a r t s of t h e c a p s u l e t h a t d o n o t u n d e r g o
ossification.
(a) Cranial t o the first cervical somite, there
are four occipital somites- The mesenchyme Bones that are Partly Formed in Cartilage
arising from the sclerotomes of these somites and Partly in Membrane
helps to form part of the base of the skull in (a) Occipital: The interparietal p a r t (lying
the region of the occipital bone. above the superior nuchal lines) is formed
Human Embryology

in membrane; the rest of the bone is formed 5. The occipital bone may be fused to the
by endochondral ossification. atlas vertebra.
(b) Sphenoid: The lateral part of the greater 6. See mandibulo-facial dysostosis.
wing and the pterygoid laminae are formed
in membrane; the rest is cartilage bone. FORMATION OF THE LIMBS
(c) Temporal: The squamous and tympanic
parts are formed in membrane. The petrous The bones of the limbs, including the bones of the
and mastoid parts are formed by ossification shoulder and pelvic girdles, are formed from
of the cartilage of the otic capsule. The mesenchyme of the limb buds. With the exception
styloid process is derived from the cartilage of the clavicle (which is a membrane bone), they
of the second branchial arch. are all formed by endochondral ossification.
(d) Mandible: Most of the bone is formed in The limb buds are paddle-shaped outgrowths
membrane in the mesenchyme of the that arise from the side-wall of the embryo at the
mandibular process. The ventral part of beginning of the second month of intrauterine life
Meckel's cartilage gets embedded in the (Fig. 10.8). Each bud is a mass of mesenchyme
bone. The condylar and coronoid processes covered by ectoderm.
are ossified from secondary cartilages that
appear in these situations. At the tip of each limb bud, the ectoderm is
thickened to form the apical ectodermal ridge.
The development of the hyoid bone has been This ridge has an inducing effect on underlying
described in Chapter 9. mesenchyme causing it to remain undifferen-
tiated and to proliferate. Areas away from the
Anomalies o f the Skull ridge undergo differentiation into cartilage,
muscle, etc.
1. The greater part of the vault of the skull is
missing in cases of anencephaly.
The forelimb buds appear a little earlier than
2. The skull may show various types of defor-
the hindlimb buds. As each forelimb bud grows, it
mity. In one syndrome, deformities of the
skull are associated with absence of the
clavicle [cleidocranial dysostosis). Prema- Pharyngeal arches
ture union of the sagittal suture gives rise
to a boat-shaped skull {scaphocephaly).
Early union of the coronal suture results
in a pointed skull (acrocephaty).
Asymmetrical union of sutures results in a
twisted skull (plagiocephaly). When the
brain fails to grow the skull remains small
{microcephaly).
3. The bones of the vault of the skull may be
widely separated by expansion of the cra-
nial cavity in congenital hydrocephalus.
4. In a rare congenital condition called Hand-
Schuller-Christian disease, large defects
are seen in the skull bones. The primary
defect is in the reticuloendothelial system;
the changes in the bones are secondary. Fig. 10.8 Embryo showing limb buds.
The Skeleton

Fig. 10.9 Stages in differentiation of the forelimb bud.

becomes subdivided by constrictions into arm, Each bud has a preaxial (or cranial) border and a
forearm and hand. The hand itself soon shows postaxial border (Fig. 10.11). The thumb and great
outlines of the digits. The interdigital areas show toe are formed on the preaxial border.
cell death because of which the digits separate The radius is the preaxial bone of the forearm.
from each other (Fig. 10.9). Similar changes occur In later development, the forelimb is adducted to
in the hindlimb. the side of the body (Fig. 10.11). The original
While the limb buds are growing, the ventral surface forms the anterior surface of the
mesenchymal cells in the buds form cartilaginous arm, forearm and hand. In the case of the lower
models, which subsequently ossify to form the limb, the tibia is the preaxial bone of the leg.
bones of the limb. Adduction of this limb is accompanied by medial
The limb buds are at first directed forward and rotation with the result that the great toe and tibia
laterally from the body of the embryo (Fig. 10.10). come to lie on the medial side. The original ventral

Preaxial border

Forelimb
" bud

Hindlimb
bud

Fig. 10.10 Scheme to show that the longitudinal Fig. 10.11 Scheme showing that with 'adduction' of
axis of the l i m b buds is transverse to the the embryonic limb, the preaxial border
long axis of the embryonic body. becomes the lateral border.
Human Embryology

surface of the limb is represented by the inguinal


joints
region, the medial side of the lower part of the
thigh, the popliteal surface of the knee, the back The tissues of joints are derived from mesenchyme
of the leg and the sole of the foot. intervening between developing bone ends. This
The forelimb bud is derived from the part of mesenchyme may differentiate into fibrous tissue,
the body wall belonging to segments C4, C5, C6, forming a fibrous joint (syndesmosis), or into
C7, C8, Tl and T2. It is, therefore, innervated by cartilage forming a cartilaginous joint. In the case
the corresponding spinal nerves. The hindlimb bud of some cartilaginous joints (synchondrosis or
is formed opposite the segments L2, L3, L4, L5, primary cartilaginous joints) the cartilage
SI and S2. connecting the bones is later ossified, with the result

Cartilage - :•«*;

Dense
Connective tissue connective tissue
of presumptive joint
Loose
connective tissue

Cartilage -

Surroundin_ _
mesenchyme

Capsule

Capsule (^>'^ai;'^JV

» •• •

Fig. 10.12 Development of a synovial joint.


The Skeleton

that the two bones become continuous. This is seen, Supernumerary digits may be present
typically, at the joints between the diaphyses and (Polydactyly). A digit (most commonly the
epiphyses of long bones. thumb) may have an extra phalanx.
The palm or sole may show a deep
At the site where a synovial joint is to be formed, longitudinal cleft {lobster claw).
the mesenchyme is usually seen in three layers. The limbs may remain short in
The two outer layers are continuous with the achondroplasia.
perichondrium covering the cartilaginous ends Sometimes the bone ends forming a joint
of the articulating bones. The middle layer are imperfectly formed (congenital
becomes loose and a cavity is formed in it. The dysplasia). This can lead to congenital
cavity comes to be lined by a mesothelium that dislocation. The hip joint is most
forms the synovial membrane (Fig 10.12). The commonly affected.
capsule, and other ligaments, are derived from
the surrounding mesenchyme.
TIMETABLE OF SOME EVENTS
Anomalies of Limbs MENTIONED IN THIS CHAPTER
1. One or more limbs of the body may be Age Developmental Events
partially, or completely, absent (phoco-
melia, atnelia). These conditions may be 4th week Forelimb bud appears.
produced by ingestion of harmful drugs. (26th day)
2. Parr of a limb may be deformed. Deformi- 4th week Hindlimb bud appears.
ties are most frequently seen in the region (28th day)
of the ankle and foot, and are of various 5th week Limbs become paddle shaped.
types. In the most common variety of 6th week Formation of future digits can be
deformity, the foot shows marked plantar (36th day) seen.
flexion (equinus: like the horse) and Cartilaginous models of bone
inversion (varus). Hence this condition is start forming.
called talipes equinovarus or club foot.
7th week Rotation of limbs occurs.
3. Congenital strictures, congenital ampu-
8th week The elbow and knee are
tations or congenital contractures may be
present. (50th day) established, and fingers and toes
4. There may be abnormal fusion (bony or are free.
fibrous) between different bones of the Primary cenrres of ossification are
limb. Adjoining digits may be fused seen in many bones.
[syndactyly). The phalanges of a digit may 12th week Primary centres of ossification are
be fused to one another (synphalangia). seen in all the long bones.
5. A digit may be abnormally large
{macrodactyly) or abnormally short The extremities are most susceptible to
(brachydactyly). In arachnodactyly, the teratogens during the 4th to 7th weeks; and
fingers are long and thin (spider fingers). slightly less susceptible in the 8th week.
Chapter

11 Face, Nose and Palate

HIGHLIGHTS • T h e palate is formed by fusion of t h r e e


components. These are the right and left palatal
• T h e stomatodaeum (future m o u t h ) is a processes (arising from the maxillary process);
depression b o u n d e d cranially by a bulging and the primitive palate (derived from the
p r o d u c e d by the brain a n d caudally by a frontonasal process) (Fig. 11.19). Deficiency
bulging produced by the pericardial cavity. in fusion leads to various forms of cleft palate
• T h r e e p r o m i n e n c e s a p p e a r a r o u n d the (Fig. 11.20).
s t o m a t o d a e u m . These are the frontonasal
process (above) a n d t h e r i g h t a n d left
mandibular arches (first pharyngeal arches)
(Fig. 11.3A).
• The mandibular arch divides into a maxillary
process and a mandibular process (Fig. 11.3B).
• The right and left mandibular processes meet
in the midline and fuse (Fig. 11.4A). They
form the lower Up and lower jaw.
• T h e upper lip is formed by fusion of the
frontonasal process with the right and left INTRODUCTION
maxillary processes. Failure to fuse completely
leads to various forms of harelip. We have seen (Chapter 5) that, after the formation
• T h e cheeks are formed by fusion of (the of the head fold, the developing brain and the
p o s t e r i o r p a r t s of) the m a x i l l a r y a n d pericardium form two prominent bulgings on the
mandibular processes. ventral aspect of the embryo (Fig. 5.14). These
bulgings are separated by the s t o m a t o d a e u m
• T h e nose is derived from the f r o n t o n a s a l
(Fig. 11.1). T h e floor of the s t o m a t o d a e u m is
process.
formed by the buccopharyngeal membrane, which
• The nasal cavity is formed as follows. An separates it from the foregut. Soon, mesoderm
e c t o d e r m a l t h i c k e n i n g , the nasal p l a c o d e , covering the developing forebrain proliferates, and
a p p e a r s over the f r o n t o n a s a l process forms a d o w n w a r d projection that overlaps the
(Fig. 11.4A). The placode gets depressed below upper part of the stomatodaeum. This d o w n w a r d
the surface to form the nasal pit (Fig. 11.4B). p r o j e c t i o n is called the frontonasal process
The nasal pits enlarge to form the nasal cavity. (Fig. 11.2). We have also seen (Chapter 9) that the
• Paranasal sinuses appear as outgrowths from pharyngeal arches are laid d o w n in the lateral
the nasal cavity. and ventral walls of the most cranial part of the
Face, Nose and Palate

Mesenchyme
covering forebrain

Stomatodaeum

Buccopharyngeal
membrane
Foregut

Fig. 11.1 Head end of an embryo just before Fig. 11.2 Formation of frontonasal process.
formation of the frontonasal process. Compare w i t h Fig. 11.1.

foregut (Fig. 9.2B). These are also, therefore, in This arch gives off a bud from its dorsal end. This
very close relationship to the stomatodaeum. It bud is called the maxillary process (Fig. 11.3B). It
will now be readily appreciated that the face is grows ventro-medially cranial to the main part of
derived from the following structures that lie the arch which is now called the mandibular
around the stomatodaeum: process.
The ectoderm overlying the frontonasal process
(a) the frontonasal process; and soon shows bilateral localized thickenings, that
(b) the first pharyngeal (or mandibular) arch are situated a little above the stomatodaeum (Fig.
of each side (Fig. I 1.3A). 11.4A). These are called the nasal placodes. The
formation of these placodes is induced by the
At this stage each mandibular arch forms the underlying forebrain. The placodes soon sink
lateral wall of the stomatodaeum (Fig. 11.3A). below the surface to form nasal pits (Fig. 11.4B).

Frontonasal
process

Stomatodaeum

Mandibular
arch

Fig. 11.3 Development of face: Formation of mandibular and maxillary processes.


Human Embryology

Frontonasal
process Medial nasal
process
Lens placode Lateral nasal
process

Nasal pit
Eye
\-f~^ Maxillary
j / process

Fig. 11.4 Development of face (continued). (A) The right and left mandibular processes fuse and form the
lower boundary of the future mouth. The nasal placodes appear over the frontonasal process.
The lens placode appears. (B) The nasal placode is converted into the nasal pit. Elevations of the
pit form the medial and lateral nasal processes.

The pits arc continuous with the stomatodaeum nasal process (Fig. 11.5) and then with the
below. The edges of each pit are raised above the medial nasal process (Fig. 11.6). The medial
surface: the medial raised edge is; called the medial and lateral nasal processes also fuse with
nasal process and the lateral edge is called the each other. In this way the nasal pits (now
lateral nasal process. called external nares) arc cut off from the
stomatodaeum.
DEVELOPMENT O F THE FACE The maxillary processes undergo
considerable growth (Fig. 11.6). At the same
We are n o w in a position to study the formation time the frontonasal process becomes much
of the various parts of the face. narrower from side to side, with the result
that the t w o external nares come closer
Lower Lip together.
The mandibular processes of the two sides grow The stomatodaeum is now bounded above
towards each other (Fig. 11.3B) and fuse in the by the upper lip which is derived as follows
midline (Fig. 11.4A). They now form the lower (Figs 11.7, 11.8}.
margin of the stomatodaeum. If it is remembered
(a) The mesodermal basis of the lateral part
that the mouth develops from the stomatodaeum,
of the lip is formed from the maxillary
it will be readily u n d e r s t o o d t h a t the fused
process. The overlying skin is derived
mandibular processes give rise to the lower lip,
from ectoderm covering this process.
and to the lower jaw (Fig. 11.7).
(b) The mesodermal basis of the median
p a r t of the lip (called philtrum) is
Upper Lip
formed from the frontonasal process.
1. E a c h m a x i l l a r y p r o c e s s n o w g r o w s The ectoderm of the maxillary process,
medially and fuses, first with the lateral however, overgrows this mesoderm to
Face, Nose and Palate

Maxillary process
Naso-optic fuses with
furrow Fronto- lateral nasal
nasal process

Primitive mouth
Fused
mandibular proceses

Fig. 11.5 Development of the face (continued). (A) The right and left nasal pits come close to each other.
The lateral nasal process is separated from the maxillary process by the naso-optic furrow.
(B) The maxillary process fuses with the lateral nasal process obliterating the naso-optic furrow.

m e e t t h a t of t h e o p p o s i t e
Frontonasal process m a x i l l a r y p r o c e s s in t h e
midline (Fig. 11.8). As a result,
the skin of the entire upper lip
is innervated by the maxillary
nerves.
Maxillary process 4. The muscles of the face (including
(fused with medial
nasal process) those of the lips) arc derived from
mesoderm of the second branchial
Mandibular process arch and are, therefore, supplied by
the facial nerve.
Frontonasal process
(has become narrow)
Nose

Maxillary and
The nose receives contributions from the
mandibular processes frontonasal process, and from the medial
fuse to form cheek and lateral nasal processes of the right and
Mouth is narrower.
left sides.
We have seen that the external nares
are formed when the nasal pits are cut off
from the stomatodaeum by the fusion of
the maxillary process with the medial
Fig. 11.6 Development of the face (continued). (A) The
nasal process. We have also noted that the
maxillary process extends b e l o w the nasal pit and
external nares gradually approach each
fuses with the medial nasal process. In this way
other. This is a result of the fact that the
the nasal pit is separated from the stomatodaeum.
(B) The maxillary and mandibular processes
frontonasal process becomes progressively
partly fuse to form the cheek. W i t h growth of the narrower and its deeper part ultimately
maxillary processes the nasal pits c o m e closer t o forms the nasal septum. Mesoderm
each other. becomes heaped up in the median plane
Human Embryology

to form the prominence of

From frontonasal
process"
<W> r\<m> Upper part of cheek
the nose. Simultaneously, a
groove appears between the
from maxillary process regions of the nose and the
From lateral - bulging forebrain (which
nasal process
m a y n o w be c a l l e d t h e
Lower part of cheek
from mandibular process forehead) (Fig. 11.10}. As
the nose becomes prominent,
Lateral part of upper lip the external nares come to
from maxillary process
Median part of open downwards instead of
upper lip from
frontonasal process
forwards (Fig. 11.10). The
Lower lip from
mandibular process external form of the nose is
thus established. The
Fig. 11.7 Derivation of parts of the face. development of the nasal
cavity is considered later.

From frontonasal Cheeks


process
After the formation of the upper and lower lips,
the sromatodaeum (which can now be called the
m o u t h ) is very b r o a d . In its lateral part, it is
bounded above by the maxillary process and below
Maxillary process by the m a n d i b u l a r p r o c e s s . T h e s e processes
overgrows
frontonasal process. undergo progressive fusion with each other to form
the cheeks (compare Figs 11.6A and B; also sec-
Figs 11.9 and 11.10).
We have already seen (while studying the
Fig. 11.8 Formation of upper l i p : Scheme t o show formation of the upper lip) that the maxillary
h o w the maxillary process 'overgrows' process fuses with the lateral nasal process. This
the frontonasal process. fusion not only occurs in the region of the lip but

Lens placode

Maxillary
Nasal /
^ process Eye y0*'
placode /
, Mandibular
o ^ ^ r process
v° J - 1st cleft
7— 2nd arch

c Nasal pit /-^


-
/
X Pericardium

A
fK
1st cleft developing
into external ear
B
Fig. 11.9 Early stages in development of the face seen from lateral side.
Face, Nose and Palate

Maxillary and s> - .


mandibular
y processes fuse
to form cheek. f \
)°* Auricle
" of ear
From maxillary

From frontonasal^
\
process-^/
*.*_. j)
/
/
/
process \ \

\ ^ From mandibular -^^"^~y——"


process /
J

From 2nd arch


/
A B
Fig 11.10 Later stages in the development of the face as seen from the lateral aspect.

also extends from the stomatodaeum to the medial External Ear


angle of the developing eye (Figs 11.6, I1.9B).
The external ear is formed around the dorsal part
For some time this line of fusion is marked by a
of the first ectodermal cleft (Fig. 11.9B). A series
g r o o v e c a l l e d t h e naso-optic furrow or
of mesodermal thickenings (often called tubercles
nasolacrimal sulcus (Fig. 11.5A). A strip of
or hillocks) appear on the mandibular and hyoid
ectoderm becomes buried along this furrow and arches where they adjoin this cleft. The pinna (or
gives rise to the nasolacrimal duct (Chapter 19). auricle) is formed by fusion of these thickenings
Eye (see Chapter 19, Fig. 19.25).

The development of the eye itself will be dealt From a study of Figs 11.9 and 11.10, it will be
with later (Chapter 19}, bur a brief reference to it seen that when first formed, the pinna lies caudal
is necessary to form a c o m p l e t e idea of the to the developing jaw. It is pushed upwards and
development of the face. backwards to its definitive position due to the
T h e region of the eye is first seen as an great enlargement of the mandibular process.
ectodermal thickening, the lens placode, which If the mandibular process fails to enlarge, the
appears on the ventro-lateral side of the developing ears remain low d o w n . See mandibulofacial
forebrain, lateral and cranial to the nasal placode dysostosis in the following section.
(Fig. 11.4A). The lens placode sinks below the
surface and is eventually cut off from the surface
Developmental Anomalies of the Face
e c t o d e r m . T h e developing eyeball p r o d u c e s a
bulging in this situation (Fig. 11.5). The bulg- It has been seen that the formation of various
ings of the eyes are at first directed laterally parts of the face involves fusion of diverse
(Figs 11.5, 1 1.6} and lie in the angles between the c o m p o n e n t s . T h i s fusion is o c c a s i o n a l l y
maxillary processes and the lateral nasal processes. i n c o m p l e t e a n d gives rise t o v a r i o u s a n o -
With the narrowing of the frontonasal process they malies.
come to face forwards (Figs 11.6, 11.7).
The eyelids are derived from folds of ectoderm 1. Harelip: The upper lip of the hare normally
that are formed above and below the eyes, and by has a cleft. Hence the term harelip is used
mesoderm enclosed within the folds. for defects of the lips.
Human Embryology

(a) When one or both maxillary processes


do not fuse with the medial nasal
process, this gives rise to defects in
the upper lip. These may vary in
degree and may be unilateral (Figs
11.11A-C) or bilateral (Fig. 11.11D).
(b) Defective development of the lower-
most part of the frontonasal process
may give rise to a midline defect of
the upper lip (Fig. 11.1 IE).
(c) When the two mandibular processes
do not fuse with each other the lower
lip shows a defect in the midline. The
defect usually extends into the jaw
(Fig. 11.11F).
Oblique facial cleft: Non-fusion of the
maxillary and lateral nasal processes gives Fig. 11.11 Varieties of harelip. For explanation see
rise to a cleft running from the medial text.
angle of the eye to the mouth (Fig. 11.12).
The nasolacrimal duct is not formed.
Inadequate fusion of the mandibular and
maxillary processes with each other may
lead to an abnormally wide mouth
(macrostomia). Lack of fusion may be
unilateral: this leads to formation of a
lateral facial cleft. Too much fusion may
result in a small mouth {microstomia).
The nose may be bifid. This may be
Fig. 11.12 O b l i q u e facial cleft.
associated with median cleft lip. Both these
occur due to bifurcation of the frontonasal
process. Occasionally one-half of it may
be absent. Very rarely the nose forms a
cylindrical projection, or proboscis
(Fig. 11.13) jutting out form just below
the forehead. This anomaly may sometimes
affect only one-half of the nose and is
Note that the
usually associated with fusion of the two nose is missing
eyes (cyclops).
The entire first arch may remain
underdeveloped on one or both sides,
affecting the lower eyelid (coloboma type Fig. 11.13 Abnormal face showing single median
defect), the maxilla, the mandible and the eye (cyclops). A rod-like projection is
external ear. The prominence of the cheek seen above the eye (proboscis). Also see
is absent and the ear may be displaced Fig. 19.12.
Face, Nose and Palate

ventrally and caudally. There may be The eyes may be. widely separated
presence of cleft palate and of faulty {hypertelorism). The nasal bridge may be
dentition. This condition is called broad. This condition results from the
mandibulofacial dysostosis, Treacher presence of excessive tissue in the
Collins syndrome or first arch syndrome. frontonasal process.
This is a genetic condition inherited as 10. The lips may show congenital pits or
autosomal dominant. fistulae. The lip may be double.
6. One-half of the face may be underdeve-
loped or overdeveloped.
Development of Nasal Cavities
7. The mandible may be small compared to
the rest of the face resulting in a receding The nasal cavities are formed by extension of the
chin {retrognathia). In extreme cases it nasal pits. We have seen that these pits are at first
may even fail to develop (agnathia). in open communication with the stomatodaeum
8. Congenital tumours may be present in (Fig. 11.14A). Soon the medial and lateral nasal
relation to the face. These may represent processes fuse and form a partition between the
attempts at duplication of some parts. pit and the stomatodaeum. This is called the

Pharynx

Stomatodaeum

Tongue Posterior nares


formed by breaking
down of bucconasal
membrane

Fig. 11.14 (A) Parasagittal sections through developing nasal cavity. (A) Nasal pit formed. (B) Nasal pit
deepens. It is separated from the stomatodaeum by the primitive palate. (C) The nasal pit enlarges
to form the nasal sac. Posterior to the primitive palate the sac is separated from the oral cavity by
the bucconasal membrane. (D) Bucconasal membrane breaks d o w n .
Human Embryology

primitive palate (Fig. 11.14B) and is derived from frontonasal process becomes progressively
the frontonasal process. n a r r o w e r . This n a r r o w i n g of the frontonasal
The nasal pits now deepen to form the nasal process, and the enlargement of the nasal cavities
sacs which expand both dorsally and caudally t h e m s e l v e s , b r i n g t h e m closer together. T h e
(Fig. 11.14C). The dorsal part of this sac is, a t intervening tissue becomes much thinned to form
first, separated from the s t o m a t o d a e u m by a the nasal septum (Figs 11.15C, D). The ventral
thin m e m b r a n e called t h e bucconasal mem- part of the nasal septum is attached below to the
brane (or nasal fin). This soon breaks d o w n primitive palate (Fig. 11.15C). M o r e posteriorly,
(Figs 11.14D, 11.15B). The nasal sac now has a ventral the septum is at first attached t o the bucconasal
orifice that opens on the face {anterior or external membrane (Fig. 11.15D), but on disappearance
nares) and a dorsal orifice that opens into the of this membrane it has a free lower edge. The
stomatodaeum {primitive posterior nasal aperture). nasal cavities are separated from the mouth by
The two nasal sacs are at first widely separated the development of the palate.
from one a n o t h e r by the frontonasal process The lateral wall of the nose is derived, on each
(Figs 11.15A, B). We have seen, however, t h a t t h e side, from the lateral nasal process. The nasal
conchae appear as elevations on the
- Frontonasal process - lateral wall of each nasal cavity.
^^^ ^ ^ Nasal sac ^ The original olfactory placodes form
the olfactory epithelium that lies in
\yJ-y- ^ - « Primitive 1 J
the roof, and adjoining parrs of the
walls, of the nasal cavity.

Anomalies of the Nasal Cavity

1. There may be atresia of the


cavity at the external nares,
at t h e p o s t e r i o r n a s a l
a p e r t u r e o r in the cavity
proper. This m a y be
unilateral o r bilateral. Very
rarely, there may be total
Bucconasal membrane a b s e n c e of t h e n a s a l
has disappeared. passages.

Fig. 11.15 Formation of the nasal septum. (A) and (C) are coronal 2. C o n g e n i t a l defects in t h e
sections through the anterior part of the nasal sac. (B) c r i b r i f o r m p l a t e of t h e
and (D) are sections through the posterior part. (A) Right ethmoid bone may lead to a
and left nasal sacs are widely separated by the frontona- communication between the
sal process. Anterior part of nasal sac is separated from cranial cavity and the nose.
the stomatodaeum by the primitive palate. (B) Posterior 3. The nasal septum may not
part of nasal sac is separated from the stomatodaeum be in the middle line, i.e. it
by the bucconasal membrane. (C) Nasal sacs enlarge may be deflected to one side.
and come close together. The frontonasal process is The septum may be absent.
narrow and forms the nasal septum. The lower edge of
4. T h e n a s a l c a v i t y m a y
the septum reaches the primitive palate. (D) Bucconasal
membrane breaks d o w n . As a result the posterior part
communicate with the
of the nasal sac opens into the stomatodaeum. mouth.
Face, Nose and Palate

Paranasal Sinuses From Figs 11.6 and 11.10, it will be seen that
these processes not only form the upper lip but
The paranasal sinuses appear as diverticula from also e x t e n d b a c k w a r d s on either side of the
the nasal cavity. The diverticula gradually invade s t o m a t o d a e u m . T h e y c a n , t h e r e f o r e , be
the b o n e s after w h i c h they are n a m e d . T h e diagrammatically illustrated as in Fig. 11.16A. If
maxillary and sphenoidal sinuses begin to develop we cut a coronal section through the region (along
before birrh. The other sinuses develop after birth. the line XY in Fig. 11.16A) the maxillary processes
Enlargement of paranasal sinuses is associated will be seen as in Fig. 11.16B. Finally, if we n o w
with overall enlargement of the facial skeleton, c o r r e l a t e Fig. 1 1 . 1 6 B with Fig. 1 1 . 1 5 D t h e
including the jaws. This provides space in the jaws relationship of the maxillary processes to the
for growth and eruption of teeth. Growth of the d e v e l o p i n g nasal cavity a n d m o u t h is easily
facial skeleton is responsible for the gradual change understood (Fig. 11.16C).
in looks of a baby.
From each maxillary process, a plate-like shelf
grows medially (Fig. 11.16D). This is called the
DEVELOPMENT OF THE PALATE palatal process. We now have three components
To understand the development of the palate, let from which the palate will be formed. These are
us have another look at the maxillary process. (Fig. 11.19):

Frontonasal
process
Riqht Maxillary Left
/ process \

0
\!^J Stomatodaeum \ ^ J
0
Vertical section across right and left
Right Maxillary |_eft maxillary processes along
process axis XY shown in Fig. A.
Imaginary horizontal section through
right and left maxillary processes
and the frontonasal process

Maxillary process Palatal process

View similar to (8) after establishment View similar to (C) at a later stage.
of nasal cavity. The nasal septum is Palatal processes grow medially
beginning to form. from each maxillary process.

Fig. 11.16 Development of the palate.


Human Embryology

Median part of the upper lip 1. the two palatal processes; and
2. the p r i m i t i v e p a l a t e f o r m e d from the
frontonasal process.

Premaxilla - Palatal
process The definitive palate is formed by the fusion of
these three parts as follows:

\ Maxillary (a) Each palatal process fuses with the posterior


\ process
• K \ margin of the primitive palate (Fig. 11.19).
(b) The two palatal processes fuse with each
I other in the midline (Fig. 11.18A). Their
Position of nasal septum fusion begins a n t e r i o r l y a n d p r o c e e d s
Fig. 11.17 Constituents of the developing palate as backwards.
seen in a schematic horizontal section (c) The medial edges of the palatal processes
through the maxillary processes. fuse with the free lower edge of the nasal
s e p t u m (Fig. 11.1 SB), t h u s
separating the two nasal cavities
from each other a n d from the
mouth.

At a later stage, the mesoderm


in the palate u n d e r g o e s intra-
membranous ossification to form
t h e hard palate. However,
ossification does not extend into
the most posterior portion, which
Fig. 11.18 Separation of nasal cavities from each other, and from the
mouth. Compare w i t h Fig. 11.16D.
remains as the soft palate. The
part of the palate derived from
the frontonasal process forms the premaxilla, which
carries the incisor teeth.
From primitive palate
— (Frontonasal process)
This part forms premaxilla.
Cleft Palate
From palatal process Defective fusion of the various components of the
(Maxillary process)
palate gives rise to clefts in the palate. These vary
considerably in degree, as illustrated in Fig. 11.20.
Clefts of the palate that extend to its anterior
Soft palate end are associated with h a r e l i p , as both the
upper lip and the palate are formed by fusion of
the maxillary processes w i t h the f r o n t o n a s a l
process.
Fig. 11.19 Embryological subdivisions of the palate Clefts of t h e p a l a t e r e s u l t in a n o m a l o u s
and the lines of fusion of these communications between the mouth and the nose.
subdivisions. These mav be unilateral or bilateral.
Face, Nose and Palate

Median part of lip


-Premaxilla

Fig. 11.20 Varieties of cleft palate. (A) Complete non-fusion, giving rise to a Y-shaped cleft, accompanied by
bilateral harelip. (B) The left maxillary process has fused with the premaxilla, but not with the right
maxillary process. The cleft is accompanied by unilateral harelip. (C) Midline cleft extending into
the hard palate. (D) Cleft of soft palate. (E) Bifid uvula.

TIMETABLE OF SOME EVENTS DESCRIBED IN THIS CHAPTER


Age Developmental Events
4th week The frontonasal, maxillary
(28th day) and mandibular processes can be identified.
Lens and nasal placodes are also present.
5th week (31 to 35 days) The nasal pits are established.
6th week Tubercles for the development of pinna begins to be formed.
On each side, palatal process appears from the maxillary process.
7th week Eyelids are established.
Maxillary processes fuse with medial nasal process.
8th week Eyes shift from lateral to frontal position.
Bucconasal membrane ruptures.
10th week The palatal processes and nasal septum fuse with each other.
Teratogens are likely to cause lip defects if the embryo is exposed to them during the 5th and 6th
weeks. The palate is most susceptible between the 7th and 8th weeks.
Chapter

1 12 Alimentary System I: Mouth,


Pharynx and Related Structures

HIGHLIGHTS
• The oral cavity is derived partly from the
stomatodaeum (ectoderm) and partly from the
foregut (endoderm). These two are separated
by t h e b u c c o p h a r y n g e a l m e m b r a n e t h a t
disappears later (Fig. 12.1).
• Teeth are formed in relation to the dental
lamina (Fig. 12.2). An enlargement of the
lamina is formed for each tooth. It is called
the enamel organ (Fig. 12.6).
• Ameloblasts (derived from ectoderm) form the
enamel. Odontoblasts (derived from mesoderm)
form dentine. T h e pulp is formed by
mesenchyme that invaginates into the enamel
organ (Fig. 12.6E).
• Three swellings a p p e a r in the floor of the
pharynx, in relation to the first pharyngeal
a r c h . These are the right and left lingual
swellings, a n d a m e d i a n swelling the
tuberculum impar (Fig. 12.11). Another median T h e m o u t h is d e r i v e d p a r t l y f r o m t h e
swelling is formed in relation to the third and stomatodaeum and partly from the foregut. Hence
f o u r t h a r c h e s . This is the hypobranchial its epithelial lining is partly ectodermal and partly
eminence. c n d o d e r m a l . A f t e r d i s a p p e a r a n c e of t h e
• The anterior two-third of the tongue is formed buccopharyngeal membrane, the line of junction
from the lingual swellings and the tuberculum between the ectoderm and endoderm is difficult to
impar. define. The epithelium lining the inside of the lips
• The posterior one-third of the tongue is formed by a n d cheeks, a n d the palate, is m o s t probably
the cranial part of the hypobranchial eminence. e c t o d e r m a l . T h e teeth a n d g u m s are also of
• Salivary glands develop as o u t g r o w t h s of ectodermal origin. The epithelium of the tongue
buccal epithelium. is, however, derived from endoderm (Fig. 12.1).
• The palatine tonsil develops in relation to the In the region of the floor of the mouth, the
second pharyngeal pouch. mandibular processes take part in the formation
of three structures. These are:
• The pharynx is derived from the foregut.
Alimentary System I: Mouth, Pharynx and Related Structures

Stomatodaeum-

({ Alveolar process
Buccopharyngeal (teeth and gums)
membrane

- Endoderm—
-Toungue
— Foregut— 1 B
Fig. 12.1 Derivation of the ectodermal part, and endodermal pari of the floor of the mouth.
(A) Stomatodaeum separated from foregut by buccopharyngeal membrane. (B) Buccopharyngeal
membrane disappears. (C) Lips, cheeks and gums lined by ectoderm, tongue by endoderm.

(a) the lower lip (and the lower part of cheeks);


TEETH
(b) the lower jaw; and
(c) the tongue. The teeth are formed in relation to the alveolar
process. T h e epithelium overlying the convex
At first these regions are not demarcated from border of this process becomes thickened and
each other (Fig. 12.2A). Soon the tongue forms a projects into the underlying m e s o d e r m . This
recognizable swelling, which is separated laterally epithelial thickening is called the dental lamina
from the rest of the mandibular process by the (Figs 12.2C, D). The dental lamina is, in fact,
linguo-gittgival sulcus {Fig. 1 2 . 2 B ) . S o o n , apparent even before the alveolar process itself is
thereafter, another more laterally placed sulcus defined (Fig. 12.2C). As the alveolar process is
makes its appearance. This is called the labio- semicircular in outline (Fig. 12.3) the dental lamina
gingival sulcus (Fig. 12.2C). This sulcus deepens is similarly curved (Fig. 12.5A).
rapidly and the tissues of the mandibular arch The dental lamina now shows a series of local
lateral to it form the lower lip (or cheek). With thickenings, each of which is destined to form one
the deepening of these two sulci, the area lying milk tooth. These thickenings are called enamel
between them becomes a raised alveolar process organs. There are ten such enamel organs (five on
(Fig. 12.2D). This process forms the jaw, and the each side) in each alveolar process (Fig. 12.5B).
teeth develop in relation to it. The tongue, the The stages in the formation of an enamel organ
alveolar process (or jaw) and the lips (or cheeks) and the development of a tooth are as follows:
are thus separated from one another (Fig. 12.3).
The roof of the mouth is formed by the palate, 1. As already stated each enamel organ is
the d e v e l o p m e n t of w h i c h has a l r e a d y been formed by localized proliferation of the cells
considered. The alveolar process of the upper jaw of the dental lamina (Figs 12.6A, B).
is separated from the upper lip and cheek by 2. As the enamel organ grows downwards into
appearance of a labio-gingival furrow, just as in the mesenchyme (of the alveolar process)
the lower jaw. The medial margin of the alveolus its l o w e r e n d a s s u m e s a c u p - s h a p e d
becomes defined when the palate becomes highly appearance (Fig. 12.6C). The cup comes to
arched (Fig. 12.4). be occupied by a mass of mesenchyme
Some anomalies in the region of the mouth are called the dental papilla. (According to
described in Chapter 11. some authorities, this mesenchyme is of
Human Embryology

Developing tongue
Linguo-gingival sulcus
Floor of developing mouth
L
Mandibular
process

Labio-gingival sulcus

Up

Alveolar process

Fig. 12.2 (A) The floor of mouth formed by fused mandibular processes. (B) Linguo-gingival sulcus
separates the developing tongue from the rest of mandibular processes. (C) Labio-gingival sulcus
separates alveolar process from lip (or cheek). The dental lamina, seen in the alveolar process,
gives origin to teeth. Also see Fig. 12.3.

Alveolar process neural crest origin.) The enamel organ and


the denral papilla together constitute the
tooth germ. At this stage the developing
tooth looks like a cap: it is, therefore,
described as the cap stage of tooth
Tongue - development.
Cheek - _ Linguo-gingival 3. The cells of the enamel organ that line the
sulcus
papilla become columnar. These are called
ameloblasts (Fig. 12.6D).
4. Mesodermal cells of the papilla that are
adjacent to the ameloblasts arrange
Fig. 12.3 Floor of mouth s h o w i n g labio-gingival themselves as a continuous epithelium-like
and linguo-gingival sulci. layer. The cells of this layer are called
Alimentary System I: Mouth, Pharynx and Related Structures

A m e l o b l a s t s lay d o w n
enamel on the superficial
_ Maxillary process ( o u t e r ) s u r f a c e of t h e
(Upper lip and cheek) basement m e m b r a n e . T h e
o d o n t o b l a s t s lay d o w n
dentine on its deeper surface.
The process of laying down
of enamel and of dentine is
Labio-gingival furrow similar to that of formation
of bone by osteoblasts. As
layer after layer of enamel
and dentine are laid down,
the layer of ameloblasts and
Arched palate Xj*/f \
the layer of o d o n t o b l a s t s
Labio-gingival furrow move away from each other
Alveolar process
(Fig. 12.7).
After t h e e n a m e l is fully
Fig. 12.4 Development of some structures seen in relation to the formed the ameloblasts
roof of the m o u t h . (A) Maxillary processes and palate. d i s a p p e a r leaving a t h i n
(B) Labio-gingival furrow separates upper lip (or upper m e m b r a n e , the dental
part of cheek) from alveolar process (of upper jaw). cuticle^ over the enamel. The
(C) Medial margin of alveolar process becomes distinct
odontoblasts, however,
because of upward arching of the palate.
c o n t i n u e t o s e p a r a t e the
d e n t i n e from the p u l p
t h r o u g h o u t the life of the
Dental lamina
Enamel organ tooth.
T h e alveolar p a r t s of the
maxillae and mandible are
formed by ossification in the
corresponding alveolar
process. As ossification pro-
gresses, the roots of the teeth
become surrounded by bone.

Fig. 12.5 Formation of enamel organs from dental lamina. T h e root of t h e t o o t h is


(A) Dental lamina f o l l o w i n g the curve of the alveolar established by continued growth
process (Compare w i t h Fig. 12.3). (B) Enamel organs into underlying mesenchyme.
formed in relation to the dental lamina.
O d o n t o b l a s t s in this region lay
down dentine. As layers of dentine
odontoblasts (Fig. 12.6E). The ameloblasts are deposited, the pulp space becomes progressively
a n d o d o n t o b l a s t s a r e s e p a r a t e d by a narrower and is gradually converted into a canal
basement membrane. The remaining cells through which nerves and blood vessels pass into
of the papilla form the pulp of the tooth. the tooth.
The developing tooth now looks like a bell In t h e r e g i o n of t h e r o o t t h e r e a r e n o
[bell stage). a m e l o b l a s t s . T h e d e n t i n e is c o v e r e d by
Human Embiyology

Dental papilla

Mesenchymal cells Odontoblasts


Cells of pulp

Fig. 12.6 Stages in (he formation of a tooth germ. (A) Dental lamina formed by proliferation of ectoderm
lining the alveolar process. (B) Deeper part of dental lamina enlarges to form enamel organ.
(C) Mesodermal cells invaginate the enamel organ to form the papilla. (D) Layer of ameloblasts
(ectoderm) formed from deepest cells of enamel organ. (E) Odontoblasts, derived from
mesodermal cells, form a layer next to the ameloblasts.

m e s e n c h y m a l cells t h a t d i f f e r e n t i a t e i n t o the temporary teeth and of the permanent incisors,


cementohlasts. These cells lay d o w n a layer of c a n i n e s a n d first m o l a r s s h o w c o n s i d e r a b l e
dense bone called the cementum. Still further to d e v e l o p m e n t . T h e g e r m s of t h e p e r m a n e n t
t h e o u t s i d e , m e s e n c h y m a l cells f o r m t h e p r e m o l a r s a n d of t h e s e c o n d m o l a r s a r e
periodontal ligament which connects the root to rudimentary. The germ of the third molar is formed
the socket in the jaw bone. after birth. The developing tooth germs undergo
The permanent teeth are formed as follows: calcification. All the t e m p o r a r y teeth and the
permanent lower first molar begin to calcify before
(a) The dental lamina gives off a series of buds, birth; the other permanent teeth begin to calcify
one of which lies on the medial side of each at varying ages after birth.
developing milk tooth (Fig. 12.9). These The eruption of a tooth is preceded by a major
b u d s form e n a m e l o r g a n s e x a c t l y as development of its root. The ages at which teeth
described a b o v e . T h e y give rise t o the e r u p t vary c o n s i d e r a b l y . T h e a v e r a g e age of
permanent incisors, canines and premolars. eruption is as follows:
(b) The permanent molars are formed from buds
that arise from the dental lamina posterior (a) Temporary or Milk Teeth
to the region of the last milk tooth. Lower central incisor 6 - 9 months
Upper incisors 8 - 1 0 months
T h e dental lamina is established in the sixth Lower lateral incisors 1 2 - 2 0 months
week of intrauterine life. At birth, the germs of all First molar 12-20 months
Alimentary System I: Mouth, Pharynx and Related Structures

Anomalies of Teeth
Dental lamina
1. One or more teeth may be
Outer wall of enamel
organ (disappearing) absent. Complete absence is
called anodentia.
2. Supernumerary teeth may be
present.
3. Individual teeth may be
abnormal. They may be too
large or too small. They may
have supernumerary cusps or
Pulp
roots. Alternatively, cusps or
roots may be less than normal.
4. Two (or more) teeth may be
fused to each other [gemination).
Odontoblasts
5. The alignment of the upper and
lower teeth may be incorrect
(malocclusion). This may be
Part of
caused by one or more of the
developing jaw above anomalies or by defects
of the jaws.
Fig. 12.7 Parts of a developing tooth. Ameloblasts lay d o w n 6. Eruption of teeth may be
enamel. Odontoblasts lay d o w n dentine.
precocious (i.e. too early).
Ossification in relation to mesenchymal cells
surrounding the developing tooth forms the jaw.
Lower incisors may be present
at birth.
Canines 16-20 months 7. Eruption of teeth may be
Second molars 20-39 months delayed. The third molar frequently fails
(b) Permanent Teeth to erupt.
First molar 6-7 years Teeth may form in abnormal situations,
Central incisors 6-8 years e.g. in the ovary or in the hypophysis
Lateral incisors 7-9 years cerebri.
Premolars 10-12 years There may be an improper formation of
Canines 10-12 years the enamel or dentine of the tooth.
Second molars 11-13 years
Third molars 17-21 years TONGUE
Summary of derivation of parts of tooth The tongue develops in relation to the pharyngeal
arches in the floor of the developing mouth. We
Ectoderm Ameloblasts — > - ENAMEL
have seen that each pharyngeal arch arises as a
mesodermal thickening in the lateral wall of the
foregut and that it grows ventrally to become
Mesoderm continuous with the corresponding arch of the
(of neural crest origin ?) Odontoblasts — • DENTINE opposite side (Fig. 12.10). The medial-most parts
of the mandibular arches proliferate to form two
lingual swellings (Fig. 12.11). The lingual
Mesenchyme around tooth CEMENTUM swellings are partially separated from each other
PERIODONTAL LIGAMENT by another swelling that appears in the midline.
Human Embryology

Tooth erupting through gum

Fifth arch

Tracheal groove

Sixth arch

Fig. 12.10 Floor of primitive pharynx: Stage 1. Note


that the right and left pharyngeal arches
meet in the m i d l i n e to form the floor of
the pharynx.
Fig. 12.8 Diagram of an erupting temporary tooth.
Note its relationship to the jaw. Al<
observe germ of permanent tooth. subsequently marked by a depression called the
foramen caecum.
Another, midline swelling is seen in relation to
the medial ends of the second, third and fourth
arches. This swelling is called the hypo-branchial
Milk teeth eminence (Fig. 12.1 I). The eminence soon shows
--^ In •v C a subdivision into a cranial part related to the
In -^
C second and third arches (called the copula) and a
\ PM
M *'• caudal part related to the fourth arch (Fig. 12.12A).
bkM The caudal part forms the epiglottis.
PMQ- The anterior two-third of the tongue is formed
Germs of
permanent by fusion of:
teeth
MO
(a) the tuberculum impar, and
M0 (b) the two lingual swellings.
\ MIJ
The anterior two-third of the tongue is thus
derived from the mandibular arch (Figs 12.12B, C).
Fig. 12.9 O r i g i n of germs of permanent teeth. According to some, the tuberculum impar does
Germs of permanent incisors, canines,
not make a significant contribution to the tongue.
and premolars are formed in relation to
temporary teeth (as seen in Fig. 12.8).
The posterior one-third of the tongue is derived
Permanent molars arise from the dental from the c r a n i a l p a r t of the h y p o b r a n c h i a l
lamina behind the part that gives rise to eminence (copula) (Fig. 12.12). In this situation,
temporary teeth. the second arch mesoderm gets buried below the
surface. The third arch mesoderm grows over it
This median swelling is called the tuberculum to fuse with the m e s o d e r m of the first a r c h
impar. Immediately behind the tuberculum impar, (Fig. 12.13}. The posterior one-third of the tongue
the epithelium proliferates to form a downgrowth is thus formed by third arch mesoderm.
{thyroglossal duct) from which the thyroid gland The posterior-most part of the tongue is derived
d e v e l o p s . T h e site of t h i s d o w n g r o w t h is from the fourth arch (Fig. 12.13).
Alimentary System I: Mouth, Pharynx and Related Structures

Tuberculum impar Lingual swelling

Lingual swelling

Tuberculum impar

Foramen caecum

. of hypobranchial
eminence

Fourth
arch
The fifth arch
has disappeared

Sixth arch

Fig. 12.11 Floor of primitive pharynx: Stage 2. The


fifth pharyngeal arch has disappeared.
Note the right and left lingual swellings,
and the tuberculum impar formed in
relation to the first arch; and the
hypnhranchial eminence formed in
relation to the medial ends of the third
and fourth arches.

In keeping with its embryological origin, the


anterior two-third of the tongue is supplied by the Anterior two-third
lingual branch of the mandibular nerve, which is of tongue
the post-trematic nerve of the first arch, and by
the chorda tympani which is the pre-trematic nerve
of this arch. The posterior one-third of the tongue
is supplied by the glossopharyngeal nerve, which
is the nerve of the third arch. The most posterior
part of the tongue is supplied by the superior laryn- Posterior one-third
ot tongue
geal nerve, which is the nerve of the fourth arch.
The musculature of the tongue is derived from Sulcus terminalis
the occipital myotomes. This explains its nerve
supply by the hypoglossal nerve, which is the nerve Foramen caecum
of these myotomes.
Epiglottis
The epithelium of the tongue is at first made
up of a single layer of cells. Later it becomes
stratified and papillae become evident. Taste buds
are formed in relation to the terminal branches of Fig. 12.12 Scheme to show the origin of different
the innervating nerve fibres. parts of the tongue.
Human Embryology

2nd arch 4th arch Anomalies of the Tongue


1st arch 3rd arch 1. The tongue may be too large {macro-
glossia) or too small {microglossia). Very
rarely the tongue may be absent {aglossia).
2. The tongue may be bifid because of non-
fusion of the two lingual swellings.
3. The apical part of the tongue may be
anchored to the floor of the mouth by an
overdeveloped frenulum. This condition is
called ankyloglossia or tongue-tie. It
interferes with speech. Occasionally, the
tongue may be adherent to the palate
{ankyloglossia superior).
4. A red, rhomboid-shaped smooth zone may
be present on the tongue in front of the
foramen caecum. It is considered to be the
Fig. 12.13 Scheme to show h o w the second arch is
buried by overgrowth of the third arch,
result of persistence of the tuberculum impar.
during development of the tongue. 5. Thyroid tissue may be present in the
tongue either under the mucosa or within
the muscles.
The derivation of various components is 6. Remnants of the thyroglossal duct may
summarized in Table 12.1. form cysts at the base of the tongue.
Development of the tongue starts in the fourth 7. The surface of the tongue may show
week of intrauterine life. fissures.

Table 12.1 Summary of derivation of components of the tongue

Part of tongue Embryonic part Nerve supply


from which derived General Taste Motor
sensation

Mandibular Facial
Epithelium over First arch (Chorda
anterior two-thirds {lingual br.)
tympani)

Epithelium over Third arch Glosso- Glosso-


posterior one-third pharyngeal pharyngeal

Epithelium over Fourth arch Superior Superior


posterior-most part laryngeal br. laryngeal br.
of vagus of vagus

Occipital
MUSCLE myotomes Hypoglosal
Alimentary System I; Mouth, Pharynx and Related Structures

Salivary Glands endodermal pouches are formed in relation to the


lateral wall of this part of the foregut. In addition,
The salivary glands develop as outgrowths of the
the floor of the foregut gives rise to a midline
buccal epithelium. The outgrowths are at first solid
diverticulum (Fig. 14.20} from which the entire
and are later canalized. They branch repeatedly
respiratory system is developed (Chapter 14).
to form the duct system. The terminal parts of the
Most of the endodermal pouches lose contact
duct system develop into secretory acini.
with the pharyngeal wall. T h e opening of the
As t h e s a l i v a r y g l a n d s d e v e l o p n e a r t h e
p h a r y n g o - t y m p a n i c tube represents t h e site of
junctional area between t h e e c t o d e r m of t h e
origin of the tubotympanic recess. The site of the
stomatodaeum and the endoderm of the foregut,
midline respiratory diverticulum is represented by
it is difficult t o d e t e r m i n e w h e t h e r they a r e
the inlet of the larynx.
ectodermal or endodermal.
With the establishment of the palate and mouth,
The outgrowth for the parotid gland arises in
the pharynx shows a subdivision into nasopharynx,
relation to the line along which the maxillary and
oropharynx and laryngopharynx.
mandibular processes fuse t o form the cheek. It is
The muscles forming the wall of the pharynx
generally considered to be ectodermal.
are derived from the third a n d subsequent
The outgrowths for the submandibular and
pharyngeal arches.
sublingual glands arise in relation to the linguo-
gingivai sulcus. They are usually considered t o
TIMETABLE OF SOME EVENTS
be of endodermal origin.
DESCRIBED IN THIS CHAPTER
O n e o r m o r e of t h e salivary g l a n d s m a y
sometimes be absent. Age Developmental events
4 weeks Tongue starts forming, i.e.
Tonsils two lateral lingual swelling
The palatine tonsil develops (on each side) in and tuberculum impar
relation to the lateral part of the second pharyngeal become visible.
pouch. The endoderm lining the pouch undergoes 5 weeks Hypobranchial eminence
c o n s i d e r a b l e p r o l i f e r a t i o n . It i n v a d e s t h e becomes visible.
underlying mesoderm of second arch, which forms 6 weeks Dental laminae of upper and
the tonsillar stroma. As a result, most of the pouch lower jaws are established.
is obliterated. Lymphocytes collect in relation to 7 weeks Salivary glands starts
the tonsillar stroma beneath the epithelium. It is developing.
not certain whether these lymphocytes differentiate 8 weeks Enamel organs are formed.
in situ or are derived from blood (possibly, they 10 weeks Enamel organ becomes cup-
come to the tonsil from the liver as lymphoblasts). shaped.
The intratonsillar cleft or tonsillar fossa is believed 3 months Formation of tonsil begins.
t o r e p r e s e n t a persisting p a r t of t h e second 5 months The tonsil is infiltrated by
pharyngeal pouch. lymphatic tissue.
Similar epithelial proliferations and 6 months Enamel and dentine have
aggregations of lymphoid tissue give rise t o the formed considerably.
tubal tonsils, the lingual tonsil and the pharyngeal Formation of tongue is
tonsils. almost complete.
Just before birth Cementum is formed.
Pharynx After birth Periodontal ligaments are
The pharynx is derived from the cranial-most part formed before eruption of
teeth.
of the foregut. We have already seen that the
Chapter

13 Alimentary System II:


Gastrointestinal Tract

HIGHLIGHTS come to lie on the right side; and the jejunum


and ileum lie mainly in the left-half of the
• Endoderm, which is at first in the form of a abdominal cavity.
flat sheet, is converted into a tube by formation
of head, tail and lateral folds of the embryonic
disc. This tube is the gut.
• T h e gut c o n s i s t s of foregut, midgut and
hindgut. T h e m i d g u t is at first in w i d e
communication with the yolk sac (Fig. 13.1).
Later it becomes tubular. Part of it forms a
loop t h a t is divisible into prearterial and
postarterial segments (Fig. 13.2),
• The most caudal part of the hindgut is the
cloaca. It is partitioned to form the primitive
rectum (dorsal) and the primitive urogenital
sinus (Fig. 13.4).
• The oesophagus is derived from the foregut.
• T h e stomach is derived from the foregut
INTRODUCTION
(Fig. 13.12).
• Duodenum: The superior part and the upper The epithelial lining of the various parts of the
half of the descending part is derived from the gastrointestinal tract is of endodermal origin. In
foregut. The rest of the duodenum develops the region of the m o u t h and the anal c a n a l ,
from the midgut. however, some of the epithelium is derived from
• The jejunum and ileum are derived from the the ectoderm of the s t o m a t o d a e u m and of the
prearterial segment of the midgut loop. proctodaeum respectively.
We h a v e seen t h a t as a r e s u l t of t h e
• The postarterial segment of the midgut loop
establishment of the head and tail folds, part of
gives off a c a e c a l b u d . T h e caecum and
the cavity of the yolk sac is enclosed within the
appendix are formed by enlargement of this
bud. embryo to form the primitive gut (Fig. 5.14). The
primitive gut is in free communication with the
• T h e ascending colon d e v e l o p s from the rest of the yolk sac. The part of the gut crania! to
postarterial segment of the midgut loop. this communication is the foregut, the part caudal
• After its formation the gut undergoes rotation. to the communication is the hindgut, while the
As a result the caecum and ascending colon i n t e r v e n i n g p a r t is the midgut (Fig. 1 3 . 1 ) .
Alimentary System II: Gastrointestinal Tract

Foregut Midgut Hindgut


Buccopharyngeal
membrane

/ Cloacal
Stomatodaeum " membrane

Proctodaeum
Allantoic diverticulum

Fig. 13.1 Parts of the primitive gut. Compare w i t h Fig. 5.11.

Cranially, the foregut is s e p a r a t e d from the to lie outside the abdominal cavity of the embryo.
stomatodaeum by the buccopharyngeal memb- It passes through the umbilical opening into a part
rane. Caudally, the hindgut is separated from the of the extra-embryonic coelom that persists in
proctodaeum by the cloacal membrane. At a later relation to the most proximal part of the umbilical
stage of development, these membranes disappear cord. The loop is subsequently withdrawn into the
and the gut opens to the exterior at its two ends. abdominal cavity.
While the gut is being formed, the circulatory While considering the formation of the allantoic
system of the e m b r y o undergoes considerable diverticulum it was seen that the diverticulum
development. A midline artery, the dorsal aorta, opens into the ventral aspect of the hindgut
is established and comes to lie just dorsal to the (Figs 5.14, 13.1). The part of the hindgut caudal
gut (big. 13.2). It gives off a series of branches ro to the attachmenc of the allantoic diverticulum i*.
the gut. Those in the region of the midgut, initially, called t h e cloaca. T h e cloaca soon s h o w s a
run right up to the yolk sac and are, therefore, subdivision into a broad ventral part and a narrow
called vitelline arteries. Subsequently, most of these dorsal part (Fig. 13.3). These two pans are separated
ventral branches of the dorsal aorta disappear and from each other by the formation of the urorectal
only three of them remain; one for the foregut, septum, which is first formed in the angle between
one for the midgut and one for the hindgut. The the allantois and the cloaca (Figs 13.4A, B). The
artery of the abdominal part of the foregut is the ventral subdivision of the cloaca is now called the
coeliac, that of the midgut is the superior mesenteric primitive urogenital sinus and gives origin to some
and that of the hindgut is the inferior mesenteric. parts of the urogenital system. The dorsal part is
The wide communication between the yolk sac called the primitive rectum. It forms the rectum
and the midgut is gradually n a r r o w e d d o w n and part of the anal canal. The urorectal septum
(Fig. 13.2B> with the result t h a t the m i d g u t g r o w s t o w a r d s the cloacal m e m b r a n e and
becomes tubular. Thereafter, the midgut assumes eventually fuses with it (Fig. 13.4C). The cloacal
the form of a loop (Fig.13.2C). T h e superior membrane is now divided into a ventral urogenital
mesenteric artery now runs in the mesentery of membrane, related to the urogenital sinus, and a
this loop to its apex. The loop can, therefore, be dorsal anal membrane related t o the rectum.
said to have a proximal, or prearterial, segment Mesoderm around the anal membrane becomes
and a distal, orpostarterial, segment. A bud (called heaped up with the result that the anal membrane
caecal bud) soon arises from the postarterial comes to lie at the bottom of a pit called the anal
segment very near the apex of the loop (Fig. 13.2D}. pit, or proctodaeum. The anal pit contributes to
the formation of the anal canal.
For a number of weeks, the midgut loop comes
Human Embryology

Midgut
Prearterial
segment
Superior
mesenteric
artery

Yolk sac_—....-
remnant t—^ f
Vitello-intestinai
duct disappears
Vitelline artery

Abdominal wail

Midgut - Midgut loop


Vitello-
intestinai duct

B Yolk sac

Fig. 13.2 Establishment of the midgut loop. (A) Midgut in w i d e c o m m u n i c a t i o n w i t h the yolk sac. Note
vitelline artery passing from dorsal aorta to the yolk sac. (B) Yolk sac m u c h smaller and attached to
midgut through a narrow vitello-intestinai duct. The original vitelline artery gives branches to the
midgut. (Cl The midgut increases in length and forms a l o o p . The l o o p has a prearterial segment
and a postarterial segment. (D) M i d g u t loop passes out of abdominal cavity. The caecal bud arises
from the postarterial segment.

Ventral part of cloaca


Urogenital sinus

Primitive
rectum
Dorsal part of cloaca Urorectal
septum
A B

Fig. 13.3 Formation of urorectal septum as seen in transverse sections. This septum divides the cloaca into
the primitive urogenital sinus and the primitive rectum.
Alimentary System II: Gastrointestinal Tract

Hindgut

Primitive urogenital sinus

Urogenital membrane

Anal membrane

Fig. 13.4 Formation of urorectal septum as seen in longitudinal sections through the cloaca.

Derivatives of Foregut 2. Jejunum


3. Ileum
1. Part of the floor of the mouth, including the 4. Caecum and appendix
tongue 5. Ascending colon
2. Pharynx 6. Right two-thirds of transverse colon
3. Various derivatives of the pharyngeal
pouches, and the thyroid
4. Oesophagus Derivatives of Hindgut
5. Stomach 1. Left one-third of transverse colon
6. Duodenum: Whole of the superior (first) part 2. Descending and pelvic colon
and upper half of the descending (second) 3. Rectum
part (up to the major duodenal papilla) 4. Upper part of anal canal
7. Liver and extra-hepatic biliary system 5. Parts of the urogenital system derived from
8. Pancreas the primitive urogenital sinus
9. Respiratory system
At this stage, it may be noted that endoderm of
Derivatives of Midgut the foregut, midgut and hindgut gives rise only to
the epithelial lining of the intestinal tract. The
1. Duodenum: Descending (second) part distal smooth muscle, connective tissue and peritoneum
to the major papilla; horizontal (third) and are derived from splanchnopleuric mesoderm
ascending (fourth) parts (Fig. 13.6).
Human Embryology

^Ectoderm

~"]0f Intra-embryonic
coelom
|.'.,.|.|.|.|.1.|.'.|.'.|.|..N-l-k.H-.|..|.|.I.M-T
Endoderm
Submucosa,
Notochord Muscle wall, and
Serous layer from
splanchnopleuric
mesoderm

Fig. 13.6 Derivation of the coats of the gut.

the neck, with the descent of the diaphragm, and


Dorsal with the enlargement of the pleural cavities.
\ mesogastrium The musculature of the oesophagus is derived
from mesenchyme surrounding the foregut. Around
the u p p e r t w o - t h i r d s of the o e s o p h a g u s , the
mesenchyme forms striated muscle. Around rhe
lower one-third, the muscle formed is smooth (as
Ventral over the rest of the gut).
mesogastrium

Stomach
Fig. 13.5 Scheme to show h o w the gut is formed
by lateral folding of the embryonic The stomach is first seen as a fusiform dilatation
disc. (A) Embryonic disc before lateral of the foregut, just distal to the oesophagus. Its
folding. (B) The lateral edges of the disc d o r s a l b o r d e r is a t t a c h e d t o t h e p o s t e r i o r
g r o w in a ventral direction. (C) The abdominal wall by a fold of peritoneum called
edges pass medially to meet in the rhe dorsal mesogastrium. Its ventral border is
middle line. In this way, the layer of attached to the septum transversum by another
endoderm is converted into a tube fold of peritoneum called the ventral mesogastrium
w h i c h is the future gut. The ectoderm
(Figs 13.7A, B).
also meets in the m i d l i n e and cuts off
the coelom from the exterior. Subsequently, the liver and the diaphragm are
formed in the substance of the septum transversum.
The ventral mesogastrium now passes from the
stomach to the liver and from the liver to the
DERIVATION OF INDIVIDUAL PARTS d i a p h r a g m and anterior a b d o m i n a l wall (Fig.
OF ALIMENTARY TRACT 13.7C). The part of the ventral mesogastrium
between the liver and the stomach becomes the
Oesophagus lesser omentum, while the part between the liver
T h e oesophagus is developed from rhe part of the and the diaphragm (and anterior abdominal wall)
foregut between the pharynx and the stomach. It gives rise to the coronary and falciform ligaments.
is at first short but elongates with the formation of Similarly, the dorsal mesogastrium is divided
Alimentary System II: Gastrointestinal Tract

Lienorenal ligament Kidney

Spleen

Liver

Fig. 13.7 (A) Side view of stomach showing the dorsal and ventral mesogastriums.
(B) Transverse section through (A) showing that the ventral mesogastrium
connects the stomach to the septum transversum. (C) The most important
remnant of the ventral mesogastrium is the lesser omentum. It passes from the
stomach to the liver (which develops in the septum transversum). The spleen is
formed in relation to the dorsal mesogastrium. Its formation divides this part of
the dorsal mesogastrium into (he gastrosplenic ligament and the lienorenal
ligament.

by the development of the spleen into a part anterior surface and the original right surface
b e t w e e n s t o m a c h a n d spleen {gastrosplenic becomes the posterior surface.
ligament) and a part between spleen and posterior
abdominal wall called the lienorenal ligament
Duodenum
<Fig. 13.7C).
The stomach undergoes differential g r o w t h The superior (or first) part and the upper half of
resulting in considerable alteration in its shape the descending (or second) part of the duodenum
and orientation. The original ventral border comes are derived from the foregut. The rest of the
to face upward and to the right and becomes the duodenum develops from the most proximal part
lesser curvature. The dorsal border n o w points of the midgut (Fig. 13.8A). T h e part of the gut
downwards and to the left and becomes the greater that gives rise to the d u o d e n u m forms a l o o p
curvature. T h e original left surface becomes its attached t o the posterior abdominal wall by a
Human Embryology

From |
foregut \ ^ / - — - Bile duct _ Dorsal
mesogastrium

B
Mesoduodenum
From
midgut

Fig. 13.8 Development of the duodenum. (A) Part of the duodenum above the entry of
the bile duct is derived from the foregut; and the part below this level is derived
from the midgut. (B) & {Q At first the duodenum has a mesentery called the
mesoduodenum. As seen in (B) this is continuous, cranially, with the dorsal
mesogastrium. The mesoduodenum later disappears (Fig. 13.9).

mesentery (mesoduodenum) (Figs 13.8B, C). Later, In keeping with its development, the proximal
this loop falls to the right. The mesoduodenum part of the duodenum is supplied by branches of
then fuses with the peritoneum of the posterior the coeliac artery and the distal part by branches
abdominal wall, with the result that most of the of the superior mesenteric.
duodenum becomes retroperitoneal (Fig. 13.9). The
mesoduodenum persists in relation t o a small part
of the duodenum adjacent to the pylorus. This is Jejunum and Ileum
the part seen in radiographs as the duodenal cap. The jejunum and most of the ileum are derived

-*- Mesoduodenum

Fig. 13.9 Scheme to show how the mesoduodenum disappears. The duodenum then becomes
retroperitoneal.
Alimentary System II: Gastrointestinal Tract

Artery
midgut loop (Fig. 13.10). The
caecum and appendix are formed
by enlargement of this bud. The
Postarterial proximal part of the bud grows
segment of gut rapidly to form the caecum. Its
distal part remains narrow and
forms the appendix (Fig. 13.11).
During the greater part of fetal
life, the appendix arises from the
apex of the caecum (Fig. 13.11).
Caecal bud -*- Subsequently, the lateral (or right)
wall of the caecum grows much
more rapidly than the medial (or
left) wail with the result that the
Fig. 13.10 Midgut loop. In this figure the loop has been drawn to point of attachment of the
correspond with the orientation of the ileocaecal region in appendix comes to lie on the
postnatal life. (Actually, the prearterial segment is cranial medial side (Fig. 13.11).
to the postarterial segment.)

from the prearterial segment of the midgut loop.


Ascending Colon
The terminal portion of the ileum is derived from
the postarterial segmenr proximal ro the caecal It develops from the postarterial segment of the
bud (Fig. 13.12). midgut loop (Fig. 13.12) distal to the caecal bud.

Caecum and Appendix Transverse Colon


We have seen that the caecal bud is a diverticulum The right two-thirds of the transverse colon develop
that arises from the postarterial segment of the from the postarterial segmenr of the midgut loop.
The left one-third arises from the
hindgut. This mode of origin is
reflected in its arterial supply; the
Large _
intestine right two-thirds are supplied by the
superior mesenteric artery and the
left one-third by the inferior
mesenteric.

Descending Colon
The descending colon develops
from the hindgut.

Rectum
The rectum is derived from the
Fig. 13.11 Development of caecum and appendix. The orientation i primitive rectum, i.e. the dorsal
as in Fig. 13.10. subdivision of the cloaca.
Human Embryology

Stomach and
part of duodenum
from foregut

Part of duodenum
(below entry of bile duct)
s*T\ \T n * — V / ^ jejunum and most of
r t \ 1) \ ^ / \ ileum from prearterial
segment of midgut

Midgut Prearterial and

Distal parts of colon,


- rectum, and part of
anal canal from hindgut
Terminal ileum, caecum,
ascending colon, and right
2/3 of transverse colon from
postarterial segment of midgut

Fig. 13.12 Derivation of various abdominal parts of the gut.

According to some authorities, the upper part of (Fig. 13.13). The line of junction of the endodermal
the rectum is derived from the hindgut proximal and ectodermal parts is represented by the anal
to the cloaca. valves (pectinate line).

Anal Canal ROTATION OF THE GUT


The anal canal is formed partly from the endodcrm We have seen that after its formation, the midgut
of the primitive rectum and partly from the loop lies outside the abdominal cavity of the
ectoderm of the anal pit or proctodaeum embryo, in a part of the extra-embryonic coelom

Hindgut Anal membrane


- lined by disappears. The
endoderm anal canal is thus

n
- established. It is
lined partly by
Anal endoderm and partly
rtnai
membrane by ectoderm.
membrane
Anal pit lined
by ectoderm M I
Anal valves

Fig. 13.13 (A) Anal membrane separates hindgut from anal pit. (B) Anal membrane disappears. ( Q Scheme
to show the parts of the anal canal in w h i c h the lining epithelium is derived from ectoderm or
endoderm.
Alimentary System If: Gastrointestinal Tract

that persists near the umbilicus. The loop has a important part in establishing the definitive
prearterial (or proximal), segment and a relationships of the various parts of the intestine.
postarterial (or distal), segment (Fig. 13.2C). The steps of the rotation must, therefore, be clearly
Initially, the loop lies in the sagittal plane, its understood.
proximal segment being cranial and ventral to
the distal segment (Fig. 13.14A). The midgut loop Viewed from the ventral side, the loop
now undergoes rotation. This rotation plays a very undergoes an anticlockwise rotation by 90°,

Foregut (stomach)
Midgut loop rotates.
The prearterial segment
comes to lie to the right
and the postarterial
segment to the left.

Right
Postarterial
segment of
midgut (caudal)
Coils of jejunum
and ileum formed
by elongation of
prearterial segment

Coils of jejunum
and ileum pass behind Superior
the artery to the left side. mesenteric artery

As a result the
hindgut (des. colon)
is pushed to the left.

Fig. 13.14 Stages in rotation of the gut. Study these figures carefully along w i t h the corresponding
description in the text. In (E) note that the caecum moves to the right and the transverse c o l o n
now lies in front of the superior mesenteric artery.
Human Embryology

Finally, the p o s t a r t e r i a l
Pre arterial
^^^^^^ segment segment of the midgut loop
/^^\. r e t u r n s t o the a b d o m i n a l
A [> cavity. As it does so, it also

segment
f^i
^r ~ V J J
i Arrows indicate
direction of
(o»Q)° rotates in an anticlockwise
direction (Fig. 13.14E) with
the result the transverse
^^^^^^r rotation.
Artery colon lies a n t e r i o r t o the
superior mesenteric artery
and the caecum conies to lie
Outline of umbilical on the right side. N o t e that
^ ^. opening
all rotation has taken place

•t ^ V f^TY in an anticlockwise direction

iy
{Fig. 13.15).

\
S>
Fig. 13.15 Scheme to show the orientation of the proximal and distal
At this stage the caecum lies
just below the liver, and an
ascending colon cannot be
demarcated. Gradually, the
caecum descends to the iliac
ends of the midgut loop at different phases of the rotation fossa, a n d the ascending,
of the gut. Arrows indicate the direction of rotation. transverse and descending
Compare with Fig. 13.14. parts of the colon become
distinct.

with the result t h a t it n o w lies in the FIXATION OF THE GUT


horizontal plane. The prearterial segment
c o m e s to lie on the right side a n d the At first all parts of the small and large intestines
postarterial segment on the left (Compare have a mesentery by which they are suspended
Figs 13.14A and B.) from the posterior a b d o m i n a l wall. After the
T h e prearterial segment n o w undergoes completion of rotation of the gut, the d u o d e n u m ,
great increase in length to form the coils of the ascending colon, the descending colon and the
the jejunum and ileum. These loops still lie rectum become retroperitoneal by fusion of their
outside the abdominal cavity, to the right mesenteries with the posterior abdominal wall (as
side of the distal limb (Fig. 13.14C). indicated in Fig. 13.9). The original mesentery
The coils of jejunum and ileum (prearterial persists as the mesentery of the small intestine,
segment) n o w return t o the a b d o m i n a l the transverse mesocolon and the pelvic mesocolon.
cavity. As they do so, the m i d g u t l o o p
undergoes a further anticlockwise rotation. Developmental Anomalies of the Gut
A s a result, the coils of jejunum and ileum
pass behind the superior mesenteric artery Congenital Obstruction
into the left-half of the abdominal cavity This may be due to a variety of causes.
(Fig. 13.14D). The d u o d e n u m , therefore,
comes to lie behind the artery, and the coils 1. Atresia: The continuity of the lumen of
of jejunum and ileum occupy the posterior the gut is interfered w i t h d u e t o t h e
and left part of the abdominal cavity. following:
Alimentary System II: Gastrointestinal Tract

' Constricted segment


(site of obstruction)

Fig. 13.17 Megacolon.

condition is called megacolon or


Hirschsprung's disease (Fig. 13.17).
Fig. 13.16 Varieties of atresia and stenosis of the
4. Abnormal thickening of muscular wall:
gut.
This is seen typically at the pyloric end of
the stomach (Congenital pyloric stenosis)
(a) A segment of the gut may be missing (Fig. 13.8). The thickened muscle bulges
(Fig. 13.16A). into the lumen and narrows it. According
(b) A segment of the gut may be replaced to some authorities, the primary cause of
by fibrous tissue (Fig. 13.16B). this defect is the same as in megacolon.
(c) A septum may block the lumen
5. External pressure by abnormal peritoneal
(Fig. 13.16C).
bands or abnormal blood vessels: Such
2. Stenosis: The lumen may be abnormally bands are often seen in relation to the
narrow (Fig. 13.16D). duodenum (Fig. 13.19). The duodenum
(Comment: As a normal developmental may also be compressed by an annular
process, there is epithelial occlusion of the pancreas (Fig. 13.20).
lumen of gut in early stages of develop-
ment. The gut later gets recanalized. Some
cases of atresia, duplication and stenosis Hypertrophied pyloric
sphincter
of gut may be due to abnormal recanali-
zation)
3. Non-development of nerve plexuses in the
wall of a part of the intestinal tract: This
may result in difficulty in the passage of
intestinal contents through the part. Such
a defect in the lower part of the colon gives
rise to a condition in which the colon
proximal to the defective segment becomes
greatly distended with its contents. This Fig.13.18 Congenital pyloric stenosis.
Human Embryology

Gall bladder

Fig. 13.19 Obstruction of d u o d e n u m by a


cystocolic band passing from the gall
bladder to the transverse c o l o n .

Fig. 13.21 Various types of imperforate anus.


(A) Stenosis of anal canal. (B) Persistent
anal membrane. (C) The proctodaeum is
represented by a solid mass of
ectodermal cells and there is a big gap
between it and the hindgut (rectum).
(D) Upper and lower parts of rectum
separated by a gap.

1A Larynx-^

((I (/I
Fig. 13.20 Annular pancreas surrounding the
duodenum. Trachea- ', I —
m
6. Imperforate anus: This is caused by A B C D
stenosis or atresia or the lower part of the Oesophagus
rectum or the anal canal. Some varieties
of this condition are shown in Figs 13.21 Fig. 13.22 (A) Normal arrangement of trachea and
and 13.23. oesophagus. (B), (C), (D) Various forms of
li,u hro-oesoph.ige.il ristul.ie.
Abnormal Communications or Fistulae
(a) Tracheo-oesophageal fistula: Atresia of the
Parts of the gut may have abnormal oesophagus is often accompanied by
communications with other cavities or with the abnormal communications between the
surface of the body. These are most frequently oesophagus and trachea, as illustrated in
seen in relation to the oesophagus and the Fig. 13.22.
rectum, and are usually associated with atresia (b) Incomplete septation of the cloaca: The
of the normal passage. rectum may communicate with the urinary
Alimentary System II: Gastrointestinal Tract

Sacrum
Pubic
Urinary bladder symphysis \ ^ W

Anal pit

D
J
- \ Anal pit
Abnormal opening

Vagina
Vagina Anal pit

Urethra * /
A n a l pit
Vagina

Fig. 13.23 Various types of rectal fistulae in the male (A to D) and female (E to H). The fistula may be between
rectum and urinary bladder (i.e. recto-vesical) as in (A) and (F), between rectum and urethra
(recto-urethra!) as in (B) and (C), and between rectum and vagina (recto-vaginal) as in (G), (H) and
(F). More than one type may be present at the same time (F). The rectum may open on to the
perineum at an abnormal site (D), (E). In these cases the anal pit is formed at the normal site.
Human Embryology

Fig. 13.24 Degrees of duplication of the gut represented by a cyst on the terminal ileum as in (A), and by
duplication of the entire c o l o n and terminal ileum as in (B).

bladder, urethra, or vagina (Figs 13.23A-


C, E, F), or may open onto the perineum
at an abnormal site (Figs 13.23D, G).
These conditions are associated with
imperforate anus.

Duplication
Varying lengths of the intestinal tract may be
duplicated. The duplicate part may form only
a small cyst, or may be of considerable length.
It may or may not communicate with the rest
of the intestine (Fig. 13.24).

Diverticula
These may arise from any part of the gut. They Fig. 13.25 Sites at w h i c h congenital diverticula may
are most common near the duodenum arise from stomach and d u o d e n u m .
(Fig. 13.25).
Persistence of a part of the vitello-intestinal may be present in its wall. (In such cases
duct may give rise to the presence of a ulceration and perforation can occur in the
diverticulum attached to the terminal part of diverticulum.) Occasionally the whole of the
the ileum. This is called Meckel's diverticulum vitello-intestinal duct, or its distal part alone,
or diverticulum ilei. It is of surgical importance may be patent. The former condition leads to a
as it may undergo inflammation giving rise to fecal fistula at the umbilicus. The latter
symptoms similar to those of appendicitis. condition leads to formation of an umbilical
Meckel's diverticulum is also of interest, as sinus. The vitello-intestinal duct may be
pancreatic tissue or a gastric type of mucosa represented by cysts (enterocystoma or vitelline
Alimentary System II: Gastrointestinal Tract

Anterior
abdomial wall

Cyst—1;

Fig. 13.26 Anomalies in relation to the vitello-intestinal duct (See arrows). (A) Meckel's diverticulum.
(B) Patent vitello-intestinal duct. (C) Umbilical sinus. (D) Cyst attached to the abdominal wall. A
cyst may also be seen attached to the gut, or embedded in the abdominal wall as shown in (E).
(E) Stenosis of gut at the site of attachment of duct. (F) Vitello-intestinal duct represented by a
fibrous cord. An umbilical growth arising from remnants of the duct is also shown.

cyst) or by fibrous cords (Fig. 13.26). Fibrous (c) Non-return of umbilical hernia:
cords constitute a danger in later life, as coils Sometimes, the coils of intestine that
of intestine may get twisted round these leading develop from the midgut loop remain
to strangulation. Remnants of the vitello- outside the abdominal cavity. The child is
intestinal duct may also give rise to growths. born with loops of intestine hanging out
of the umbilicus. This condition is called
Errors of Rotation exomphalos or omphalocoele (Fig. 13.28).
(a) Non-rotation ofthe midgut loop: In this Loops of intesrine and other abdominal
condition the small intesrine lies towards contents may also be seen outside the
the right side of the abdominal cavity, and abdominal cavity for an entirely different
the large intestine towards the left reason. In congenital umbilical hernia rhe
(Fig. 13.27A). muscle layer and skin are absent in the
(b) Reversed rotation: The transverse colon region of the umbilicus, creating a defect
crosses behind the superior mesenteric in the abdominal wall through which
artery and the duodenum crosses in front abdominal contents can pass. Such contents
of it (Fig. 13.27B). are covered with peritoneum, but in
Human Embryology

Fig. 13.27 Errors of rotation. (A) Non-rotation. Coils of small intestine lie in the right-half of the abdomen and
colon in the left half. (B) Reversed rotation. The duodenum lies anterior to the superior mesenteric
artery and the colon crosses behind it.

exomphalos they are covered only by


amnion.

Errors of Fixation
(a) Parts of the intestine that are normally
retroperitoneal may have a mesentery.
Abnormal mobility of this part of the
Umbilical cord
intestine may result in its twisting. This
condition is called volvulus. Twisting of
blood vessels to the loop can lead to
obstruction of its blood supply.
(b) Parts of the intestines, which normally,
have a mesentery, may be fixed by
abnormal adhesions of peritoneum.
Coils of intestine (c) The caecum may remain sub-hepatic or
• Body wall may descend only to the lumbar region.
Alternatively, it may descend into the
pelvis (Fig. 13.29).
Fig. 13.28 Exomphalos. Coils of intestine derived
from the midgut loop fail to return into
Situs Inversus
the abdominal cavity. In this condition all abdominal and thoracic
Alimentary System II: Gastrointestinal Tract

Fig. 13.29 Errors in descent of the caecum. (A) Subhepatic. (B) Lumbar. <C) Pelvic. The normal position is
shown in dotted line in (A) and (B).

viscera are laterally transposed, i.e. all parts and duodenum He on the left side and the
normally on the right side are seen on the left stomach on the right side.
side, and vice versa. For example, the appendix

TIMETABLE OF SOME EVENTS DESCRIBED IN THIS CHAPTER


Age Developmental Events
16 days Allontoic diverticulum starts appearing.
3 weeks Gut begins to acquire tubular form because of head and tail foldings.
At the end of third week the buccopharyngeal membrane ruptures.
4 weeks The fusiform shape of the stomach becomes visible.
5 weeks Stomach rotates and dilates. Intestinal loop begins to form.
Caeca! bud can be identified.
6 weeks Intestinal loop is well formed.
Urorectal septum starts dividing the cloaca.
Allantois and appendix become clearly visible.
Stomach completes its rotation.
7 weeks Septation of cloaca into rectum and urogenital sinus is completed.
Intestinal loop herniates out of the abdominal cavity.
8 week Intestinal loop rotates counterclockwise.
9 weeks Anal membrane breaks down.
3 months Head and tail foldings are completed.
Herniated coils of intestine return to the abdominal cavity.
Chapter

14 Respiratory System; Body Cavities

HIGHLIGHTS • The peritoneal cavity is derived from the right


and left limbs of the intra-embryonic coelom.
• The liver and biliary passages (endoderm) are The two limbs unite to form a single cavity
derived from the hepatic bud. This bud arises after formation of lateral folds of the embryonic
from the gut at the junction of foregut and disc.
midgut.
• The pleural cavities are formed from right and
• The pancreas (endoderm) develops from two left pericardio-peritoneal canals that connect
buds, dorsal and ventral, that arise from the the pericardial and peritoneal cavities. Each
gut near the junction of foregut and midgut. canal is invaginated by the corresponding lung
Most of the pancreas is formed from the dorsal bud. Enlargement of the bud leads to great
bud. The ventral bud forms part of the head of enlargement of the canal and formation of the
the pancreas. pleural cavity.
• The spleen (mesoderm) develops in the dorsal • The diaphragm develops in relation to the
mesogastrium. septum transversum. It receives contributions
• The respiratory system develops from a median from the pleuro-peritoneal membranes, the
diverticulum of the foregut (endoderm). At its b o d y wall a n d the m e s e n t e r i e s of the
caudal end the diverticulum divides into right oesophagus.
and left lung buds.
• The larynx and trachea develop from the part LIVER, PANCREAS AND SPLEEN
of the respiratory diverticulum cranial to its
division. LIVER
• The lung buds undergo repeated division to The liver develops from an endodermal bud that
establish the bronchial tree and alveoli of the arises from the ventral aspect of the gut, at the
lungs. point of junction between foregut and midgut
• The peritoneal, pericardial and pleural cavities (Fig. 14.1A). This bud grows into the ventral
develop from the intra-embryonic coelom. This mesogastrium and passes through it into the septum
coelom at first consists of right and left halves transversum (Fig. 14.IB). It enlarges and soon
t h a t are connected, across the middle line, shows a division into a larger cranial part called
cranial to the prochordal plate. the pars hepatica, and a smaller caudal portion
• The pericardial cavity is derived from the called the pars cystica (Fig. 1 4 . I C ) . T h e pars
median midline part of the intra-embryonic h e p a t i c a d i v i d e s i n t o r i g h t a n d left p a r t s ,
coelom. After formation of the head fold this e a c h of w h i c h f o r m s o n e l o b e of t h e liver
cavity comes to lie ventral to the foregut. (Figs 14.1D, E).
Liver, Pancreas, Spleen; Respiratory System; Body Cavities

Foregut — —

Hepatic bud

Ventral
mesogastrium
Midgut

Pars ^^0
_ hepatica ^^m

Gall bladder —
rs cystica I

Fig. 14.1 Development of the biliary apparatus. (A) Hepatic bud arises from the gut at the junction of
foregut and midgut. (B) It grows towards the septum transversum through the ventral
mesogastrium. (C) The bud divides into the pars hepatica (that forms the liver) and the pars cystica
(that forms the gall bladder). The part of the hepatic bud proximal to its division forms the bile
duct.

As the right and left divisions of the pars formation (haemopoiesis). Large aggregations of
hepatica enlarge and extend into the septum blood-forming cells are present between hepatic
transversum, the cells arising from them are cells and blood vessels (Fig. 14.2).
broken up into interlacing columns called hepatic Bile formation begins when the fetus is about
trabeculae. In this process, the umbilical and three months old. The bile is responsible for the
vitelline veins which lie in the septum transversum, black colour of the first stools {meconium) passed
are broken up to form the sinusoids of the liver. by the newborn.
Sinusoids are also formed from the mesenchyme
of the septum transversum.
GALL BLADDER AND BILIARY PASSAGES
The endodermal cells of the hepatic bud give
rise to the parenchyma of the liver and to bile The pars cystica of the hepatic bud gives origin to
capillaries. The mesoderm of the septum the gall bladder and to the cystic duct (Fig. 14.1).
transversum forms the capsule and fibrous tissue The part of the hepatic bud proximal to the pars
basis of the liver. cystica forms the bile duct. The bile duct at first
The fetal liver is an important centre of blood opens on the ventral aspect of the developing
Human Embryology

Hepatic
artery

Portal
triad

Kupffer
vein
eel! Haematopoietic
cells (in fetal life)

Fig. 14.2 Some histological components in developing liver. Derivatives of endoderm


(hepatic bud) are shown in shades of pink, those of mesenchyme of septum
transversum in green, and those from vitelline and umbilical veins in yellow.

duodenum. As a result of differential growth of the uncinate process of the pancreas, while the
the duodenal wall, and as a result of the rotation upper part of the head, the body and the tail are
of the duodenal loop, it comes to open on the formed from the dorsal bud (Fig. 14.6).
dorsomediat aspect of the duodenum along with The duct system of the pancreas is established
the ventral pancreatic bud (Fig. 14.5). as follows. The ducts of the dorsal and ventral
buds anastomose with each other (Fig. 14.7). The
duct of the dorsal bud, between this anastomosis
PANCREAS and the duodenum, remains narrow and forms the
The pancreas develops from two endodermal buds, accessory pancreatic duct {Fig. 14.7C). The main
dorsal and ventral, which arise from the part of pancreatic duct is formed, in its distal part, by the
the gut that later forms the second part of the
duodenum. The ventral bud arises in close relation
to the hepatic bud, in the inferior angle between it
and the duodenum (Fig. 14.3). The dorsal bud
arises from the dorsal aspect of the gut (Figs 14.3,
14.5A) and grows into the mesoduodenum and Hepatic bud
the dorsal mesogastrium. When the duodenal loop
falls to the right, the ventral bud comes to point to
the right and the dorsal bud to the left (Fig. 14.5B). Ventral
pancreatic bud
Thereafter, as a result of differential growth of the
wall of the gut, the attachment of the ventral bud
(along with the bile duct) also shifts to the left
side (Fig. 14.5C). Pancreatic tissue formed from Midgut
these two buds now fuses to form one mass. The
ventral bud forms the lower part of the head and Fig. 14.3 Dorsal and ventral pancreatic buds.
Liver, Pancreas, Spleen; Respiratory System; Body Cavities

duct of the dorsal bud and,


Pharynx in its proximal part, by the
duct of the ventral bud. The
Respiratory main pancreatic duct,
diverticulum
therefore, opens into the
Septum transversum duodenum at the major
duodenal papilla, along
with the bile duct.
The secretory elements
of the pancreas are formed
by proliferation of the
primitive ducts. The islets
of Langerhans are also
derived from the primitive
duct system.
Developing liver
Cystic bud (gall bladder)
SPLEEN
Ventral pancreatic bud The spleen develops as a
Dorsal collection of mesenchymal
mesentery cells in the dorsal mesogas-
Ventral mesentery trium (Fig. 14.8A). Some of
these cells are contributed
Fig. 14.4 Diagram showing developing liver. Note the pancreatic buds and by the coclomic epithelium
the respiratory diverticulum. lining the mesogastrium.
The mesenchymal cells
differentiate into lymphoblasts and other blood
forming cells.
As the mesenchymal cells proliferate, they form
s a mass which projects to the left and is covered by

,00
- Ventral bud
peritoneum (Fig. 14.8B). The dorsal mesogastrium,

From dorsal bud

B
1
Fig. 14.5 Changes in relative position of ^
pancreatic buds. (A) Initial position in
w h i c h the ventral and dorsal buds lie in
. J
the direction indicated by their names.
(B) Position after duodenal loop falls to )uct
the right. The ventral b u d n o w points to creatic tissue
the right, and the dorsal bud to the left. From ventral bu J
(C) The attachment of the ventral bud
moves to the left and the t w o buds now Fig. 14.6 Parts of pancreas derived from ventral
lie close together. and dorsal buds.
Human Embryology

in this region, can now be divided into


Duct of dorsal bud
a part extending from the stomach to
D A the spleen (gastrosplenic ligament) and
\77L another part extending from the spleen
to the posterior abdominal wall. The
• Duct of ventral bud
l a t t e r p a r t fuses w i t h the p o s t e r i o r
abdominal wall (Fig. 14.9A) with the
Bile duct
result that a fold of peritoneum n o w
passes from the spleen to the left kidney
D B (lienorenal ligament). As a consequence
C7Z Two ducts anastomose
of this fusion, and as a result of a change
in t h e o r i e n t a t i o n of t h e s t o m a c h
Minor (Fig. 14.9B}, the spleen comes to lie on
duodenal Accessory pancreatic duct the left side and takes parr in forming
papilla ,
the left boundary of the lesser sac of
11 DC peritoneum.
\7ZL
Main pancreatic duct
Major 'r Anomalies of the Liver
duodenal
papilla Anomalies of the liver are rare. Some
of them are illustrated in Fig. 14.10.
Fig. 14.7 Formation of main and accessory pancreatic ducts.
Note that the distal part of the main pancreatic duct Anomalies of the Gall Bladder
is derived from the duct o\ the dorsal b u d ; and its
Anomalies of Shape
proximal part from the duct of the ventral bud.
(a) The fundus may be folded on

Dorsal
mesogastrium - Spleen

Fig. 14.8 Development of the spleen. The spleen appears in the dorsal mesogastrium as in (A) and soon
bulges to the left as in (B).
Liver, Pancreas, Spleen; Respiratory System; Body Cavities

pouch), which may be


Dorsal mesogastrium fuses with adherent to the cystic duct
peritoneum lining posterior or even to the bile duct
abdominal wall. \
Kidney (Fig. 14.11E).

Anomalies of Position
Spleen
(a) The organ may lie trans-
versely on the under-
surface of the right lobe of
the liver, or may lie under
the left lobe (Fig. 14.11G).
(b) The gall bladder may be
lined by peritoneum on all
sides. It may be attached
to the liver by a fold of peri-
toneum or may be comple-
Lienorenal ligament tely free {floating gall
bladder) (Fig. 14.11-1).
(c) It may be embedded in the
substance of the liver
(Fig. 14.11H).

Duplication
(a) The lumen may be,
partially, or completely
subdivided by a septum,
which may, or may not,
extend into the cystic duct
Lesser omentum (Figsl4.HA, B).
(b) The gall bladder may be
Fig. 14.9 Development of the spleen. First see Fig. 14.8. Note the
partially, or completely, dup-
changing relationship of the spleen to the dorsal licated (Figs 14.11B, C).
mesogastrium and to the lesser sac of peritoneum. In (A)
note how part of the dorsal mesogastrium fuses w i t h the Other Anomalies
posterior abdominal w a l l . As a result of this change the
dorsal mesogastrium is divided (at this level) into the
(a) The gall bladder may
gastrosplenic and lienorenal ligaments.
open directly into the bile
duct {sessile bladder)
(Fig. 14.1 IF).
itself, to form a cap-like structure called (b) The gal 1 bladder may be absent
the Phrygian cap (Fig. 14.11D). (Fig. 14.1 OD).
(b) The wall of the infundibulum may project (c) Diverticu la may arise from any part of
downwards as a pouch {Hartmann's the organ
Human Embryology

Ligamentum teres

^ ^ ~ P Porta hepatis _
Inferior vena cava

Fig. 14.10 Anomalies of the liver. (A) Rudimentary left lobe. (B) Anomalous lobation. (C) Reidel's lobe.
(D) Absence of quadrate lobe associated with absence of gall bladder. (E) Accessory liver in
falciform ligament.

Anomalies of the Extrahepatic Duct System right lobe may terminate in the right hepatic
duct, the cystic duct, the bile duct, or even
Abnormal Length
directly into the gall bladder.
There is considerable variation in the level at
which various ducts join each other, with the Anomalies of the Pancreas
result that occasionally some of them
may become abnormally long, or short (a) Annular pancreas: Pancreatic tissue
(Figsl4.12A-D). surrounds the duodenum completely and
may obstruct it (Fig. 13.20).
Abnormal Mode of Termination
(b) Divided pancreas: The parts of the
pancreas derived from the dorsal and
(a) The cystic duct may join the left side of ventral buds fail to fuse with each other
the common hepatic duct, passing either (Fig. 14.15).
in front of it or behind it, to reach its left (c) Accessory pancreatic tissue may be found
side (Figs 14.12E-G). in the stomach, duodenum, jejunum,
(b) The cystic duct may end in the right hepatic Meckel's diverticulum, gal! bladder and
duct (Fig. 14.12H). spleen.
(c) The cystic duct may pass downwards (d) Inversion of pancreatic ducts: The
anterior to the duodenum, before joining embryonic arrangement of the ducts
the common hepatic duct. persists and the greater part of the pancreas
(d) The bile duct may open into the pyloric, is drained through the minor duodenal
or even the cardiac, end of the stomach. papilla (Fig. 14.16).

Atresia Anomalies of the Spleen


Parts of the duct system, and sometimes the 1. The spleen may be lobulated.
whole of it, may be absent (Fig. 14.13). 2. The spleen may, very rarely, be absent.
3. Accessory spleens may be seen:
Duplication
(a) at the hilum of the spleen,
Parts of the duct system may be duplicated (b) in the gastrosplenic ligament,
(Fig. 14.14). Accessory ducts arising from the (c) in the lienorenal ligament,
Liver, Pancreas, Spleen; Respiratory System; Body Cavities

Ligamentum

S.
- ^ teres

Gall/ £"

Caudate lobe
Right lobe Inferior vena cava

Fig. 14.11 Anomalies of the gall bladder. (A) to (C) Various degrees of subdivision. In (C) there is complete
d u p l i c a t i o n . (D) Phrygian cap. (E) Hartmann's p o u c h . (F) Sessile gall bladder. (Gl Left-sided gall
bladder (1) and transverse gall bladder (2). (H) Gall bladder embedded in liver tissue. (I) Floating
gall bladder, in w h i c h the organ is covered all round by peritoneum.
Human Embryology

Ft. and L. hepatic ducts Long hepatic Very long


join within liver substance. ducts cystic duct

Cystic duct passes behind


common hepatic duct and
joints its left side.
Cystic duct passes
in front of common
hepatic duct and joins
its left side.
m Cystic duct joining
right hepatic duct

Fig. 14.12 Some anomalies of the extrahepatic duct system.

Gall bladder and


Complete agenesis Cystic duct missing
cystic duct missing

D ^

t)
Hepatic ducts missing Bile duct missing Terminal part of
bile duct missing

Fig. 14.13 Agenesis of parts of the extrahepatic biliary tract. Missing parts are indicated in light colour.
Liver, Pancreas, Spleen; Respiratory System; Body Cavities

Fig. 14.14 Duplication of parts of the extrahepatic biliary trac

(d) within the pancreas, and


From ventral bud
(e) along the splenic artery.

From dorsal bud 4. The spleen lies on the right side of the
abdomen in situ inversus. The liver and
pancreas are also reversed from side to
side.

THE RESPIRATORY SYSTEM


Fig. 14.15 Divided pancreas. The parts of the
pancreas arising from dorsal and ventral The respiratory system develops from a median
buds remain separate. diverticulum of the foregut. Its lining epithelium
is, therefore, of endodermal
origin. The connective
Bile duct
tissue, cartilage and muscle,
X Pancreatic duct from
\ / d u c t of dorsal bud in relation to the organs of
respiration, are derived from
splanchnopleuric mesoderm.
The diverticulum that is
Main drainage o f _ J —f^*7
pancreas through I V h destined to form the
minor papilla \ \J% r e s p i r a t o r y system is first
,. •• Y ^ ^ Duct of ventral bud seen as a m i d l i n e g r o o v e
\ / —'—T\ remains small.
{tracheobronchialgroove) in
Major duodenal' ^ * ^ , the floor of the developing
papilla pharynx just caudal to the
hypobranchial eminence
Fig. 14.16 Inversion of pancreatic ducts. The main pancreatic duct is (Fig. 14.17). This groove is
formed entirely by the duct of the dorsal bud and opens at the flanked by t h e sixth
minor duodenal papilla. The duct of the ventral bud remains pharyngeal arches. Soon after
small.
Human Embryology

its appearance, the distal part of the groove is


separated from the oesophagus (Fig. 14.18) but
the cranial part continues to communicate with
the pharynx.
The free caudal end of the diverticulum becomes
bifid, each subdivision being called a lung bud
Fourth - » (Fig. 14.19). The part of the diverticulum cranial
arch to the bifurcation forms the larynx and trachea,
Fifth arch has while the lung buds form the bronchi and lung
disappearead. * Tracheal
groove parenchyma.
Sixth arch

LARYNX
Fig. 14.17 Formation of tracheo-bronchial groove. T h e l a r y n x develops f r o m the cranial-most p a r t

o O
Foregut Tracheo-bronchial
groove

Fig. 14.18 Scheme to show how the respiratory diverticulum separates from the foregut. The upper figures
are transverse sections (along the axis XY) of the figures below them. In (C) and (D) note
progressive separation of respiratory diverticulum from the foregut, except at the cranial end.

1 B C
Fig. 14.19 Development of lung buds. (A) Right and left lung buds appear. (B) They divide into lobar bronchi
(3 right, 2 left). (C) Segmental bronchi established.
Liver, Pancreas, Spleen; Respiratory System; Body Cavities

of the respiratory diverticulum. The communi- Within the respiratory passages, some cells
cation between the diverticulum and the pharynx become specialized for production of surfactant.
persists as the inlet of the larynx. The caudal part This substance forms a thin layer over alveoli
of t h e h y p o b r a n c h i a l e m i n e n c e f o r m s t h e and reduces surface tension.
epiglottis. T h e thyroid, cricoid and arytenoid Before birth the respiratory passages are full
cartilages are derivatives of the fourth, fifth and of fluid which also contains surfactant. When
sixth pharyngeal arches. The laryngeal muscles the newborn begins to breathe, the fluid is rapid-
are also derived from branchial mesoderm as ly absorbed and partly expelled. The surfactant
indicated by their nerve supply. remains as a thin layer lining the alveoli. This
prevents collapse of alveoli during expiration.
In premature babies, a deficiency of surfactant
TRACHEA AND BRONCHI may cause difficulty in expansion of the lung
T h e trachea d e v e l o p s from t h e p a r t of t h e and can be a cause of death of the baby.
respiratory diverticulum, that lies between the
point of its bifurcation and the larynx. There is considerable increase in the number
The two primary divisions of the respiratory of alveoli in the postnatal period.
diverticulum form the right and left principal The pulmonary circulation is established early
bronchi. T h e left division comes to lie m o r e in fetal life. However, most of the blood is at first
transversely than the right (Fig. 14.19). It soon short-circuited through the foramen ovale and the
shows two subdivisions that represent the two ductus arteriosus. The amount of blood circulating
lobar bronchi of the left lung. The right division through the lungs progressively increases, and by
divides into three lobar bronchi. the seventh month of intrauterine life the circulation
is rich enough to provide adequate oxygen for
sustaining life. Hence an infant born, thereafter,
LUNGS is viable (i.e. it can live).
T h e substance of the lungs is formed by further T h e d e v e l o p m e n t of the pleural cavities is
subdivisions of the lobar bronchi (Fig. 14.19). (The considered later in this chapter.
total number of divisions of each main bronchus
are about 17 before birth, and six more after birth.) Anomalies of the Larynx
After the establishment of the bronchial tree,
(a) Laryngocoele: In this c o n d i t i o n , t h e
alveoli are formed by expansion of the terminal
laryngeal saccule is abnormally large. It
parts of the tree.
may extend beyond the larynx proper, and
The parts of the lung parenchyma, developing may even form a swelling in the neck.
from the lobar bronchi, are separated from one (b) Congenital stenosis or atresia: There may
another by mesoderm. This mesoderm forms the be stenosis or atresia of the larynx.
connective tissue basis of the lung and also gives (c) The entire larynx, or part of it (e.g. vocal
rise to the pleura. As the pleura lines the surface cords), may be duplicated.
of each lobe separately, the lobes come to be
(d) Laryngoptosis: The larynx lies low down in
separated by fissures.
the neck. Part of it may be behind the sternum.
During the fetal life, all subdivisions of the (e) One or more of the laryngeal cartilages
bronchial tree are lined by a cubical epithelium. may be absent.
This is the canalicular phase of lung development.
With the onset of respiration, after birth, the alveoli Anomalies of the Trachea
b e c o m e d i l a t e d a n d their lining e p i t h e l i u m 1. Tracheo-oesophageal fistulae have already
becomes thin. been described (see Fig. 13.22).
Human Embryology

Tracheal bronchus

Fig. 14.20 Some varieties of tracheal bronchi. (A) Blind bronchus. (B) Supplying accessory lobe.
(C) Replacing apical bronchus.

2. A diverticulum may arise from the trachea. Larynx


3. Accessory bronchi may arise from the
trachea. Such a bronchus may
(a) be blind (Fig. 14.20A);
(b) supply a mass of lung tissue (accessory
lobe) which is not a normal part of
the lungs (Fig. 14.20B);
(c) replace a normal bronchus (e.g.
apical) in one of the lungs
(Fig. 14.20C).
4. Very rarely the trachea may be absent. The
bronchi to the lungs may arise from a blind Oesophagus
'bifurcation1 (Fig. 14.21 B); or from the Fig. 14.21 Some anomalies of trachea. (A) Normal.
oesophagus (Fig. 14.21C). (B) Agenesis. (C) Agenesis along with
tracheo-oesophageal fistula.
Anomalies of the Lungs and Bronchi
Agenesis and Hypoplasia 2. Presence of Abnormal Fissures

A complete lung, or one of its lobes (and (a) A transverse fissure may be present
associated bronchi), may fail to develop, or may on the left side with the result that the
left lung has three lobes (Fig. 14.22B).
remain underdeveloped.
(b) The medial basal segment (cardiac
Abnormalities of Lobes lobe) of the left lung may be separated
by a fissure from the rest of the lower
1. Absence of fissures that are normally lobe (Fig. 14.22D).
present: It leads to a reduction in the (c) The superior segment of the lower
number of lobes, e.g. absence of the lobe may be similarly separated
transverse fissure of the right lung results (Fig. 14.22C).
in a right lung with only two lobes (d) A part of the upper lobe of the right
(Fig. 14.22A). lung may come to lie medial to the
Liver, Pancreas, Spleen; Respiratory System; Body Cavities

Fig. 14.22 Abnormal lobes of the lungs: (A) Right lung with only t w o lobes, (B) Left lung w i t h three lobes,
(C) Apical segment of lower lobe is separate. (D) Medial basal segment is separate.

Lung Normal position


of azygos vein

Parietal
" pleura (pink)
Azygos vein
_ Visceral
pleura (black)

Fig. 14.23 Azygos lobe of lung (B). The normal relationship of the azygos vein to the lung is shown in (A).

azygos vein. This part is called the


azygos lobe (Fig. 14.23). In this
condition, the azygos vein is suspended ^
from the wall of the thorax by a fold
of parietal pleura {mesoazygos).
/ y
Accessory lobes are usually connected to
bronchi that are not part of the normal
bronchial tree. Such bronchi may arise
from the:
(a) trachea above its bifurcation (upper
accessory lobe) (Fig. 14.20B).
(b) oesophagus (lower accessory lobe, Fig. 14.24 Accessory lobe of lung (indicated by
Fig. 14.24). Occasionally, the lobe arrow) supplied by bronchus arising
may not have any bronchus. from oesophagus.

Sequestration of lung tissue: An area of may have an independent pleural covering.


embryonic lung tissue may separate from In other cases the sequestrated tissue may
the tracheobronchial tree (sequestration = lie within a lobe (intralobarsequestration).
separation). Such tissue may form a The sequestrated lung tissue derives its
complete lobe (lobarsequestration), which blood supply from an abnormal branch of
Human Embryology

the aorta. The condition is most frequently of pericardium, pleura and peritoneum are formed
seen in the lower lobe of the left lung. from these layers of mesoderm. The mesodermal
cells lining the cavities differentiate into a flattened
Lung Hernia e p i t h e l i a l l i n i n g c a l l e d rttesothelium. The
mesothelium gives the peritoneum, pleura and
Part of a lung may herniate: (a) through the pericardium their smooth surfaces.
inlet of the thorax, (b) through a defect in the The intra-embryonic coelom is a horseshoe-
thoracic wall, (c) into the mediastinum, or shaped cavity having a narrow midline portion
(d) into the opposite pleural cavity. and two lateral parts. The midline part lies near
the cranial end of the embryonic disc (Fig. 14.25A),
Displaced Bronchi and it is from this part of the coelom that the
These may arise from the trachea above its pericardial cavity is formed. The two lateral limbs
bifurcation or even from the oesophagus. of the coclom form the peritoneal cavity. For some
They may supply: (a) a normal segment of time, the pericardial and peritoneal cavities are
one of the lungs (Fig. 14.20C), (b) an accessory connected t o each other by a pair of n a r r o w
lobe (Figs 14.20B, 14.24), or (c) they may be pericardio-peritoneal canals (Fig. 14.25B). These
blind (Fig. 14.20A). canals undergo great enlargement to form the
pleural cavities.
Details of the development of the pleural and
BODY CAVITIES p e r i t o n e a l cavities a r e c o n s i d e r e d h e r e . T h e
pericardial cavity is considered in Chapter 15.
Introduction
The pericardial, pleural and peritoneal cavities
PLEURAL CAVITY
are derivatives of the intra-embryonic coclom. We
have seen that by the formation of this cavity the After t h e f o r m a t i o n of t h e h e a d f o l d , t h e
lateral plate mesoderm is split into a parietal pericardium comes to lie on the ventral aspect of
(somatopleuric) and a visceral (splanehnopleuric) the embryo, and the pericardio-peritoneal canals
layer (Fig. 5.6). The parietal and visceral layers wind b a c k w a r d s on either side of the foregut

Median part of intra-embryonic coelom Pericardio-


forms pericardial cavity. peritoneal canal
s -*. /
Prochordal y

(©)
plate ~~y^y<

A U B
Embryonic^\

o lateral parts of intra-e mbryonic coelom


fus e to form the peritonea cavity

Fig. 14.25 (A) Intra-embryonic coelom, and (B) its subdivisions.


Liver, Pancreas, Spleen; Respiratory System; Body Cavities

Pericardio-
peritoneal canal
Peritoneal cavity

Hindgut

Foregut
Vitello-intestinal duct
Pericardial cavity Septum transversum

Fig. 14.26 Parts of intra-embryonic coelom and their relationship to the gut.

(Fig. 14.26). The lung buds, that arise from the pericardiopleural membranes and the formation
foregut, now invaginate these canals (Fig. 14.27). of the pleuro-peritoneal membranes, respectively
As the buds enlarge to form the kings, the canals (Figs L4.28B,C).
balloon out to form the pleural cavities.
Each pleural cavity now communicates with From Figs 14.28C and 14.29A, it will be seen
the pericardial cavity through the pericardio- that the pleural cavities are at first dorso-lateral
pleural opening, and with the peritoneal cavity to the pericardium. As the lungs increase in
through the pleuro-peritoneal opening size, the pleural cavities extend into the
(Fig. 14.28A). In subsequent development, these mesoderm of the body wall (which is expanding
openings are closed by the formation of the at the same time) and gradually come to He

Foregut Foregut
/~~\ ^ ^ /"V-Pericardio-peritoneal A
B
A Q f T cana, ^Q2
\o
Pericardial Lung bud
cavity

^ ^ ^ Oesophagus
Foregut

*• i
Lung bud
invaginating .

C D
Fig. 14.27 Invagination of pericardio-peritoneal canals by the lung buds.
Human Embryology

Pericardial cavity
Pericardio-peritoneal
canal enlarging to Pericardio-pleural
form pleural cavity membrane

Pleuro-peritoneal
membrane

Peritoneal cavity
A Peritoneal cavity peritoneal
opening

Position of pericardio-
pleural membrane
Pleural cavity

Position of
- pleuro-peritoneal
membrane

Fig. 14.28 Formation of pleural cavity and its separation from pericardial and peritoneal cavities.
(A) Pericardio-peritoneal canal enlarges to form the pleural cavity. The pleural cavity
communicates freely w i t h the pericardial and peritoneal cavities. (B) Pleural cavity is gradually
separated from the pericardial and peritoneal cavities by formation of pericardio-pleural and
pleuro-peritoneal membranes.

lateral, and to some extent ventral, to the membrane. Later this membrane forms the
pericardium (Fig. 14.28B). The pleural cavities fibrous pericardium. This explains the course
also extend downwards into the mesoderm of the phrenic nerve over the pericardium.
forming the posterior abdominal wall, and
upwards towards the neck (Fig. 14.28C). In Fig.
14.29B note that with the expansion of the PERITONEAL CAVITY
pleural cavity the mesoderm of the body wall We have seen that the peritoneal cavity is formed
is split into an outer part that forms the wall of from the two limbs of the horseshoe-shaped intra-
the thorax and an inner part over the pericardial embryonic coelom. The two parts are at first
cavity. The latter is called the pleuro-pericardial separate, but fuse to form one cavity, as a result
membrane. The phrenic nerve runs through this of lateral folding of the embryonic disc
Liver Pancreas, Spleen; Respiratory System; Body Cavities

Pleural cavity Pleural cavity extending


upwards into body wall

- Mesoderm of -
body wall

Septum
transversum

Peritoneal _
cavity r

Pleural cavity extending


downwards into body wall
Pleural cavity
extending backwards and
forwards into body walll

Fig. 14.29 Schemes to show how the pleural cavity expands into the body w a l l .

(Figs 14.30D, E, F). As illustrated in Figs 14.30B, in this membrane. These are the right
C, the two halves of the peritoneal cavity remain and left, pneumato-enteric recesses
separate in the cranial part of the abdomen. (Fig. 14.32B). The left recess soon
The attachment of the mesentery of the primitive disappears. The right recess enlarges and
gut on the posterior abdominal wall is at first in opens into the peritoneal cavity
the midline (Fig. 14.31). As a result of changes (Fig. 14.32C). The cavity of this recess now
involving the rotation of the gut and as a result of enlarges considerably and extends to the
some parts of the gut becoming retroperitoneal, left to form the part of the lesser sac that
the line of attachment of the mesentery becomes lies behind the stomach (Fig. 14.32D). It
complicated (Fig. 14.31). The peritoneal cavity, also extends cranially, on the right side of
therefore, comes to be subdivided into a number the oesophagus and behind the liver
of pockets that are partially separated by folds of (Fig. 14.33). Subsequently, with the
peritoneum. establishment of the diaphragm, the
uppermost part of this cranial extension
Development of Lesser Sac comes to lie above the diaphragm, where
it gives rise to the infracardiac bursa. The
Three distinct processes are involved in the part of the cranial extension that remains
formation of the lesser sac of peritoneum. These below the diaphragm (and behind the
may be considered one by one. liver) forms the superior recess of the lesser
(a) The dorsal mesogastrium that connects the sac.
stomach to the posterior wall of the (b) While the right pneumato-enteric recess
extends to the left, the stomach changes
abdomen is, initially, a thick membrane
its orientation, so that its posterior border
(Fig. 14.32A). Two small cavities appear
Human Embryology

Intra-embryonic
Ectoderm coelom

»( n EEEnnrJEEEEHiB
••IM—

Dorsal
mesogastrium

Ventral
mesogastrium

Fig. 14.30 Schemes to illustrate w h y the foregut has a ventral mesentery (A to C) but the midgut and hindgut
d o not. First refer t o legend t o Fig. 13.5. Figures (A) t o (C) represent the result of lateral f o l d i n g of
the embryonic disc in the region that w i l l form the upper part of the abdomen. As the disc folds
the t w o halves of the intra-embryonic coelom also undergo folding and meet in the middle line
ventral to the gut. From (C) it w i l l be clear h o w the dorsal and ventral mesogastriums are formed.
Figures (D) to (F) show the result of folding in the lower part of the abdomen. In (D) note that, at
this level, each half of the intra-embryonic coelom is open laterally. The result of folding is seen in
(F) from w h i c h it w i l l be clear w h y there is no ventral mesentery here.

(to which the dorsal mesogastrium was the lesser sac. It is continuous with the part
attached}, now faces to the left. This border of the lesser sac lying behind the stomach
forms the greater curvature. The ventral (derived from the right pneumato-enteric
border (to which the ventral mesogastrium recess: N in Fig. 14.34C).
was attached), now comes to face to the (c) With the altered orientation of the stomach,
r i g h t a n d forms t h e lesser c u r v a t u r e the dorsal mesogastrium, which is attached
(Fig. 14.34). The ventral mesogastrium t o t h e g r e a t e r c u r v a t u r e , m a y be
may now be called the lesser omentum. subdivided into t w o parts A and B, as
As a result of this change in the orientation shown in Fig. 14.35. If we trace these two
of the stomach, a part of the peritoneal p a r t s t o the posterior a b d o m i n a l wall
cavity c o m e s t o lie b e h i n d rhe lesser (Fig. 14.35) we find that the attachment
omentum (M in Fig. 14.34C). This part of of the mesogastrium on this wall can also
the peritoneal cavity now forms part of be divided into two corresponding parts.
Liver, Pancreas, Spleen; Respiratory System; Body Cavities

Stomach (
) (
Duodenum U,

Jejunum
and ileum
Ascending
colon
Transverse
colon
Descending
colon
Pelvic colon
Rectum

Fig. 14.31 Peritoneal relations of the gut. In (A) the gut is shown w h e n it is a simple m i d l i n e tube. In (B) the
dorsal w a l l of the abdomen is shown to indicate the m i d l i n e attachment of the dorsal mesentery.
The oesophagus and rectum are seen passing through the w a l l . In (C) it is shown that alternate
segments (3, 5, 7, 9) become retro-peritonea I w h i l e the segments 2, 4, 5 and 8 retain their
mesentery, (D) shows the ultimate disposition of these segments o n the posterior abdominal w a l l .
1 represents the ventral mesogastrium; 2, the dorsal mesogastrium; 3, the d u o d e n u m ; 4 , the
mesentery of the j e j u n u m and i l e u m ; 5, the ascending c o l o n ; 6, the transverse mesocolon; 7, the
descending c o l o n ; 8, the pelvic mesocolon and 9, the rectum.

a
—Dorsal
~"~-
Recess
\ _

y K
mesogastrium extends to

f
opens into •— left behind
Pneumato- -^ peritoneal stomach.
\ enteric ~ cavity.
\ recesses

IV - IL ft
\y ii (( )) 1:1 VI
^Stomach •*"
A B c 0
Fig. 14.32 Development of the lesser sac. Formation of pneumato-enteric recesses.

Part A passes from the stomach to the (Fig. 14.35 ) and forms the greater
spleen as the gastro-splenic ligament, and omentum. The greater omentum undergoes
from the spleen to the left kidney as the enlargement with the result that it comes
lienorenal ligament. It, therefore, forms to increasingly project below the level of
the left margin of the lesser sac. Part B the stomach and becomes folded on itself.
passes from the lower border of the The space within this fold forms the lower
stomach to the posterior abdominal wall part of the lesser sac (Fig. 14.36).
Human Embryology

Intracardiac bursa Diaphragm

Right pneumato
- enteric

Fig. 14.33 Development of the lesser sac. Extensions of the right pneumato-enteric recess. Note the
extension above the level of the diaphragm in (C).

Ventral mesogastrium

Stomach
Kidney

mesogastrium Stomach

Fig. 14.34 Schemes to explain formation of the lesser sac. The dorsal and ventral mesogastriums are shown in
(A). Note that the ventral mesogastrium has a free border facing d o w n w a r d s and forwards, if a
section is cut in the plane XY, the appearance seen is illustrated in (B). Subsequently the original
ventral border of the stomach comes to lie on the right side (C). Two parts of the lesser sac labelled
M and N are s h o w n . N is derived from the right pneumato-enteric recess w h i l e M is part of the
peritoneal cavity that comes t o lie behind the ventral mesogastrium {which is now the lesser omentum).
Liver, Pancreas, Spleen; Respiratory System; Body Cavities

PartB

Pyloric end Q

Fig. 14.35 Parts of the dorsal mesogastrium. Part A forms the gastrosplenic and lienorenal ligaments as
shown in Fig. 14.7. Part B elongates to form the greater o m e n t u m . The attachment of these parts
to the stomach is shown in B and to the posterior abdominal w a l l in C.

Fig. 14.36 Development of the lesser sac: D o w n w a r d extension of the sac by elongation and folding of the
greater o m e n t u m . The derivation of the parts numbered in (C) is: (1) f r o m cranial extension of
pneumato-enteric recess; (2) part of peritoneal cavity that comes to lie behind ventral
mesogastrium; (3) right pneumato-enteric recess; (4) cavity produced by elongation and folding of
greater o m e n t u m o n itself.

DIAPHRAGM and pleural cavities are above (or cranial to) it,
whereas the peritoneal cavity is caudal to it. The
Introduction
development of the diaphragm is, therefore,
The diaphragm is a partition that separates the intimately related to the development of these
thoracic and abdominal cavities. The pericardial cavities
Human Embryology

Spinal cord
Dorsal mesentery
of oesophagus

Pleuro-peritoneal
opening
Oesophagus

m transversum

Fig. 14.37 Development of the diaphragm. Pleuro-peritoneal canals and their closure. Note the other
structures in relation to these canals.

The formation of the septum transversum has The diaphragm is, therefore, formed from the
heen considered in Chapter 5. We have seen that following components (Fig. 14.39).
the liver develops in its caudal part. Its cranial
part helps to form the diaphragm. Reference to (a) Septum transversum
Figs 5.12, 5.13 and 5.14 will show that after the (b) Pleuro-peritoneal membranes
establishment of the head fold, the septum (c) Ventral and dorsal mesenteries of
transversum forms a mesodermal mass lying oesophagus
caudal to the pericardial cavity. It, therefore, (d) Mesoderm of body wall, including the
separates the pericardial and peritoneal cavities. mesoderm around the dorsal aorta
Posterior to the septum transversum, however, the
pleural and peritoneal cavities communicate There is, however, considerable controversy as
through the pleuro-peritoneal canals that lie on to how much of the diaphragm is formed from
either side of the oesophagus (Fig. 14.37). The each of the constituents. According to some
partition between the thorax and the abdomen is workers, the septum transversum forms only
completed when the pleuro-peritoneal canals are the central tendon, while according to others it
closed by the formation of the pleuro-peritoneal gives rise to almost the whole of the costal and
membranes (Fig. 14.37). sternal parts of the diaphragm. The crura of
the diaphragm are formed from the mesoderm
of the posterior abdominal wall, as a result of
Development of the Diaphragm the downward extension of the pleural cavities
It may be recalled (Fig. 14.29) that as the pleural into this region (Fig. 14.29).
cavities increase in size, they do so at the expense The nerve supply of the diaphragm from the
of the body wall, with the result that the thorax as third, fourth and fifth cervical nerves (through
a whole also expands. Simultaneously, the the phrenic nerve) shows that the diaphragm
diaphragm has also to enlarge and this enlargement has undergone great migration in a caudal
takes place at the expense of the body wall direction during development. (This descent is
(Fig. 14.38). caused by elongation of the neck, descent of
Liver, Pancreas, Spleen; Respiratory System; Body Cavities

P leu ro peri to real


'' canal
Peritoneal cavity

Pleural cavity

Contribution from
body wall
Pleuroperitoneal
membrane

Fig. 14.38 Development of the diaphragm: Schemes to show h o w expansion of the pleural cavities into the
body w a l l causes the w a l l to f o r m part of the d i a p h r a g m .

the heart, and expansion of the pleural cavities.) contents may pass through these gaps to
The sensory innervation of the peripheral parts produce diaphragmatic hernias.
of the diaphragm by the intercostal nerves is Diaphragmatic hernias may be
an evidence of the contribution made by the (Fig. 14.40):
body wall to the muscle.
(a) Posterolateral: due to failure of a
pleuro-peritoneal canal to close.
Anomalies of the Diaphragm
(b) Posterior: due to failure of the
1. Parts of the diaphragm may fail to develop development of the crura.
resulting in gaps in the muscle. Abdominal (c) Retrosternal: due to an abnormally

Pleuro-peritoneal
membrane
Dorsal mesentery &
Ventral mesentery
of oesophagus

Body wall

Fig. 14.39 Components from w h i c h the diaphragm is formed. These are


labelled on the right side of the figure.
Human Embryology

3. The muscle may be thin and aponeurotic


and may bulge upwards into the thorax.
The bulging may be unilateral or may be

w
^^^ ^ ^ R Y\ Posterolateral confined to a small area. This condition
is called congenital eventration of the
diaphragm.
Vc-/ ^Wji- central

Posterior \ \ ^^ TIMETABLE OF SOME EVENTS


DESCRIBED IN THIS CHAPTER
Retrosternal
Age Developmental Events
Fig 14.40 Sites of congenital defects in the
diaphragm. A b d o m i n a l contents may
3 weeks The hepatic bud appears. The
herniate through them resulting in
pancreatic bud also appears soon
diaphragmatic hernias. thereafter.
4 weeks The septum transversum is
established.
large gap between the sternal and The tracheobronchial diverticulum
costal parts of the muscle. is formed.
(d) Central: through the dome of the
diaphragm. Occasionally one entire 6 week The diaphragm descends to the
thoracic level.
half (usually the left) of the diaphragm
m a y be absent. 7 weeks Fusion of dorsal and ventral
2. Very rarely, an accessory diaphragm may pancreas.
be present in the thoracic cavity. When 3 months Bile formation begins.
present, it partially subdivides the lung into Pancreatic islets are formed.
two parts. 5 months Insulin secretion begins.
Chapter

15 Cardiovascular System

HIGHLIGHTS from the floor of the p r i m i t i v e ventricle.


(2) The bulbar septum that divides the conus
• T h e heart develops from s p l a n c h n o p l e u r i c into two parts. (3) The gap left between these
mesoderm related to that part of the intra- two is filled by proliferation of atrio-ventricular
embryonic coelom that forms the pericardial cushions that are formed in the atrio-ventricular
cavity. This mesoderm is the cardiogenic area. canal (Fig. 15.11).
• Two endothelial heart tubes (right and left) • The truncus arteriosus is continuous with the
appear and fuse to form one tube. This tube aortic sac (Fig. 15.27). This sac has right and
has a venous end and an arterial end (Figs 15.1, left horns. Each horn is continuous with six
15.2). pharyngeal (or aortic) arch arteries. These
• A series of dilatations appear on this tube arteries join the dorsal aorta (right or left). The
(Fig. 1 5 . 3 ) . T h e s e are (1) bulbus cordis, first, second and fifth arch arteries disappear.
(2) ventricle, (3) atrium, and (4) sinus venosus. The caudal parts of the right and left dorsal
a o r t a e fuse to form o n e m e d i a n vessel
• Further subdivisions are named as follows (Fig. 15.28).
(Fig. 15.3). The bulbus cordis consists of a
proximal one-third (which is dilated), a middle • The ascending aorta and pulmonary trunk
one-third called the conus and a distal one- are formed from the t r u n c u s a r t e r i o s u s
third called the truncus arteriosus. The narrow (Fig. 15.28B).
part connecting atrium and ventricle is the • The arch of the aorta is formed by the aortic
atria-ventricular canal. The sinus venosus has sac, its left horn and the left fourth arch artery
right and left horns. (Fig. 15.29A).
• The right and left atria of the heart are formed • The descending aorta is formed partly from
by p a r t i t i o n of the primitive a t r i u m . This the left dorsal aorta and partly from the fused
partition is formed by the septum primum and median vessel (Fig. 15.29B).
the septum secundum (Fig. 15.6). A valvular
• The brachiocephalic artery is formed from the
passage, the foramen ovale, is present between
right horn of the aortic sac (Fig. 15.29C)
these two septa. It allows flow of blood from
the right atrium to the left atrium. • The common carotid artery is derived from
part of the third arch artery (Fig. 15.30B).
• The dilated proximal one-third of the bulbus
cordis, the conus, and the primitive ventricle • The pulmonary artery is derived from the sixth
unite to form one chamber. This is partitioned arch artery (Fig. 15.31B).
to form right and left ventricles. This partition • The arteries to the gut are formed from ventral
is m a d e u p of the f o l l o w i n g . (1) T h e s p l a n c h n i c b r a n c h e s of the d o r s a l a o r t a
Interventricular septum that grows upwards (Fig. 15.36).
Human Embryology

• The renal, suprarenal and gonadal arteries are the main facts are presented first. Details are
formed from lateral splanchnic branches of the presented later.
dorsal aorta.
• Arteries to the body wall and limbs are derived Introduction
from dorsolateral (somatic intersegmental)
branches of the aorta. The heart (like all blood vessels) is mesodermal
in origin. It is formed from splanchnopleuric
• The left subclavian artery is derived from part
of the seventh cervical intersegmental artery. m e s o d e r m lying i m m e d i a t e l y c r a n i a l to the
The right subclavian artery is formed partly prochordal plate. This mesoderm constitutes the
from the seventh cervical intersegmental artery cardiogenic area. It is closely related to the
and partly from the right fourth arch artery. pericardial cavity (which is derived from part of
the i n t r a - e m b r y o n i c c o e l o m ) . For a g o o d
• The portal vein is formed from right and left
understanding of the relationship between the
vitelline veins and anastomoses between them
developing heart tube and the pericardial cavity
(Fig. 15.42).
students are advised to study Figs 5.11 - 5.14.
• The superior vena cava is derived from part of The heart is at first seen in the form of right
the right anterior cardinal vein and from the
and left endothelial heart tubes (Fig. 15.1A) that
right common cardinal vein.
soon fuse with each other. The single tube thus
• The inferior vena cava receives contributions formed shows a series of dilatations. These are:
from several veins (and anastomoses between
them). These are the right posterior cardinal 1. Bulbus cordis
vein, the right s u b c a r d i n a l vein, the right 2. Ventricle (We will refer to it as the primitive
supracardinal vein and the right hepatocardiac ventricle.)
channel. 3. Atrium (We will refer to it as the primitive
atrium or atrial chamber.)
THE HEART 4. Sinus venosus.

Development of the Heart: Main Facts The ventricle and atrium are connected by a
narrow atrio-ventricular canal. The sinus venosus
The development of heart is complex. To avoid has prolongations that are referred to as its right
confusion that may be caused by numerous details, and left horns.

f5^

U Sinus venosus

Fig. 15.1 (A) Right and left heart tubes. (B) to (D) Progressive fusion of tubes from cranial to caudal end.
Fusion of sinus venosus is partial.
Cardiovascular System

1st arch artery - i — Atrium

Ventral aorta — ^ — Sinus venosus

Aortic sac \ V _ _ Common


r - - cardinal vein

Umbilical vein

Fig. 15.2 (A) Arterial end and (B) Venous end of heart tube.

Distal 1/3= Forms ascending aorta &


Truncus arteriosus pulmonary trunk

Middle 1/3= Forms outflow tracts of


Conus right and left ventricles

Proximal 1/3= — Forms primitive right ventricle


ftrabeculated part)

Buibo-ventricular sulcus
Forms primitive left ventricle
(trabeculated part)

Atno-ventncutar
canal

\ Primitive atrium is
f— partitioned to form
right and left atria.

Body of sinus venosus is


absorbed into right atrium
and forms smooth part.
SINUS VENOSUS ^ / Body
Left horn of sinus venosus
Right horn of sinus ^ retrogresses. The remnant
venosus is absorbed Right Left forms part of coronary sinus.
into right atrium. horn horn

Right common cardinal


vein forms part of
m Vu Right vitelline vein forms part
superior vena cava. of inferior vena cava.
Right umbilical vein disappears.

Fig. 15.3 M a i n subdivisions of heart tube and their fate.


Human Embryology

Outline of orifice Outline of orifice


as seen in (B) as seen in (A)
Atrium - ^
<~-~--- \.

A
(
^r)
^—\y Sinuatrial orifice
v*- S> J -—' c
Septum spurium
/ s Left venous valve
/— N

. (
SL)
x \—, y
Outline of orifice V
X s
) .

as seen in (A) Right venous valve

Fig. 15.4 Changes in the sinuatrial orifice. Note that firstly, the centrally placed orifice shifts to the right.
Secondly, the orifice that is at first transversely orientated becomes vertical. Dotted lines in (B) and
(C) indicate the outline of the opening in Fig. 15.3 show how the change occurs.

The bulbus cordis lies at the arterial end of the opening. Gradually the opening becomes
heart. It is divisible into three parts, i.e. proximal, narrow and shifts to the right. Finally, it
middle and distal. T h e p r o x i m a l one-third is becomes a n a r r o w slit. The slit has right
dilated and does not have any special name; the and left margins called the right and left
middle one-third is called the conus and the distal venous valves. Cranially, these two valves
o n e - t h i r d is c a l l e d t h e truncus arteriosus fuse to form a structure called the septum
(Figs 15.2A and 15.3). The truncus arteriosus is spurium (Fig. 15.4).
continuous distally with the aortic sac. The aortic
sac is continuous with right and left pharyngeal
arch arteries. These arteries arch backwards to
become continuous with the right and left dorsal
aortae.
A * * c

Dorsal and ventral


The sinus venosus lies at the venous end of the A.V cushions appear.
Round A.V. canal
heart. It has right and left horns. O n e vitelline
vein (from the yolk sac), one umbilical vein (from
B ^ ^ ^ ^
the placenta) and one common cardinal vein (from
the body wall) join each horn of the sinus venosus.
The fate of the various parts of the heart tube is
summarized in Fig. 15.3. 1
#9?
Separate right and left A.V. orifices
created by fusion of cushions
Canal becomes
transverse
Formation of Atria
Fig. 15.5 Schemes to show how the A.V. canal
1. The sinus venosus and the primitive atrial undergoes division into right and left
chamber are at first connected by a wide orifices.
Cardiovascular System

2. The atrioventricular canal divides into right This blood has t o reach the left atrium,
and left halves as follows (Fig. 15.5). Two and for this p u r p o s e a c o m m u n i c a t i o n
thickenings, the atrio-ventricular cushions between right and left atria is essential.
appear on its dorsal and ventral walls. They Before the septum primum reaches and fuses
grow towards each other and fuse. The fused with the septum intermedium, blood flows
cushions form the septum intermedium through the gap between them. This gap is
(Fig. 15.6). the foramen primum. Before the foramen
primum can be closed it is essential that
Formation of Interatrial Septum another path for flow of blood be created.
This is achieved by breaking d o w n of the
The atrial chamber undergoes division into right
upper part of the septum primum. The new
and left halves by formation of t w o septa (that
gap is the foramen secundum. The septum
later fuse)(Fig. 15.6).
primum now has a free upper edge.
(b) The septum secundum grows down from the
(a) The s^ptam pn'mwrn arises from the roof of
the atrium, to the left of the septum spurium. roof the atrial chamber, to the right of the
It grows d o w n w a r d s t o w a r d s the atrio- septum primum. As it grows it comes to
ventricular canal and ultimately fuses with overlap the free upper edge of the septum
the septum intermedium. primum. Once the two septa overlap, blood
However, note the following carefully. has to flow through the interval between
T h r o u g h o u t fetal life oxygenated blood the septa. This gap is the foramen ovale. It
reaches the right atrium from the placenta. is a valvular aperture that allows blood to

Septum Opening of B
pulmonary vein
im /

Sinuatrial_
orifice

Common atrial
chamber

Foramen secundum
(arrow)

3D
Septum
secundum

Septum primum
(used to A.V.
cushions Right atrium

Fig. 15.6 Formation of interatrial septum. (A) Septum p r i m u m appears. (B) Septum p r i m u m grows towards
fused A.V. cushions. The gap between them is the foramen p r i m u m . ( Q Septum p r i m u m fuses
w i t h A.V. cushions. At the same t i m e the upper part of the septum p r i m u m degenerates to form
the foramen secundum. The septum secundum is formed to the right of the septum p r i m u m .
(D) Septum secundum overlaps the free edge of septum p r i m u m . Blood n o w flows f r o m right to
left through the o b l i q u e cleft between the t w o septa.
Human Embryology

flow from right to left, but not from left to and the venae cavae are seen opening into
right. the atrium.
After birth of the baby the left atrium starts The right margin of the original sinuatrial
receiving oxygenated blood from the lungs orifice (i.e. the right venous valve) expands
and there is no need for flow of blood from very greatly and divides into three parts
right atrium to left atrium. The foramen which form the crista terminalis (Fig. 15.8),
ovale is, therefore, obliterated by fusion of the valve of the inferior vena cava and the
the septum primum and septum secundum. valve of the coronary sinus. N o t e that the
In terms of adult anatomy, the annulus crista terminalis lies at the junction of
ovalis represents the lower free edge of the the part of the right atrium derived from
septum secundum while the fossa ovalis the sinus venosus (sinus venarum) and the
represents the septum primum. atrium proper.

DEVELOPMENT OF RIGHT ATRIUM DEVELOPMENT OF LEFT ATRIUM


(a) As described above, the main part of the The left atrium is derived from:
right atrium is derived from the right-half
(a) Left half of the primitive atrial chamber
of the primitive atrium.
(b) T h e s i n u s v e n o s u s is
a b s o r b e d into the right
R. horn of sinus venosus
atrium by great enlarge-

JsC^
common cardinal v.
m e n t of t h e s i n u a t r i a l
orifice (Figs 15.7, 15.8}. R common cardinal v. —
(c) The right half of the atrio-
ventricular canal is also
a b s o r b e d into the right
atrium. Sinuatrial orifice
guarded by right Atrial septum
and left venous
Some relevant facts about the valves
sinus venosus (and its tributaries!
may be noted at this stage. Right atnu
Coronary sinus formed
1. The left horn of the sinus Sinus venosus from L horn of sinus
Terminal part of absorbed into venosus & L common
venosus remains very small. inferior vena cava right atrium cardinal vein
It b e c o m e s p a r t of the from R. vitelline vein
coronary sinus (Fig. 15.7).
2. T h e right common
L venous valve fused
cardinal vein becomes part Superior vena cava with atrial septum
of the superior vena cava. from R common -
cardinal vein
3. T h e right vitelline vein Right venous valve
forms the terminal part of forming crista terminalis
the inferior vena cava.
After absorption of the Right atrium
sinus venosus into the right
atrium the coronary sinus. Fig. 15.7 Incorporation of sinus venosus into the right atrium.
Cardiovascular System

(b) Left half of the atrio-ventricular canal DEVELOPMENT OF VENTRICLES


(c) Absorbed proximal parts of the pulmonary
veins (see below). Fate of Bulbus Cordis
We have seen that the bulbus cordis is divisible
Absorption of Pulmonary Veins into three parts, i.e. proximal, middle (conus) and
At the time when the septum p r i m u m is just distal (truncus arteriosus).
beginning to form (Fig. 15.6A), a single pulmonary A spiral septum appears within the truncus
vein opens into the left half of the atrium. When arteriosus and subdivides it into the ascending
traced away from the heart (Fig. 15.9), the vein aorta and the pulmonary trunk. It is formed by
divides into a right and a left branch each of which union of right superior and left inferior truncus
again bifurcates, to drain the corresponding lung swellings or cushions. Fusion of these cushions
bud. takes place in such a manner that at its lower end,
Gradually, the parts of the pulmonary veins the pulmonary trunk lies ventral to the aorta, but
nearest to the left atrium are absorbed into the as it is traced upwards it comes to lie on its left
atrium, with the result four separate veins, two side. This is because of the orientation of the spiral
from each side, come to open into it (Fig. 15.9). septum.

Opening of sinus
venosus into
^L right atrium

Right
venous
valve
V^^Left
^ V venous
W valve

Crista terminalis

Opening of
' coronary sinus

•^ * Valve of
Superior- \ coronary sinus
limbic band inf. limbic band
Opening of inf vena cava Valve of inf. vena cava

Fig. 15.8 Fate of the right and left venous valves. The right venous valve expands greatly and forms the
crista terminalis, the valve of the inferior vena cava and the valve of the coronary sinus. The left
venous valve remains small and fuses with the interatrial septum.
Human Embryology

The conus forms the outflow tracts (smooth


parts) of both the right and left ventricles. The
\ »z Pulmonary vein proximal one-third of the bulbus cordis merges
with the cavity of the primitive ventricle.
A (f^ ~ ^ < x ^ L e f t atrium

Important Note
Wall of atrium derived
,» / ., from absorbed vein Please note that in the seventh edition of this
book the bulbus cordis was described as being
s<& ^ ^ ^ ^ % > ^ ^ - L . pulmonary vein divided into t w o parts, i.e. a distal part the
B
C 1 truncus arteriosus, and a proximal part the
L. pulmonary veins conus which was absorbed into the primitive
ventricle, and later formed the smooth outflow
sN>\ parts of both right and left ventricles, it is n o w

Y S recognized that the outflow parts are formed


by the middle one-third of the bulbus cordis
only, and it is this part that is the conus. The
Fig. 15.9 Absorption of pulmonary veins into the proximal one-third of the bulbus cordis takes
left atrium. At first only one vein from the part in forming the trabeculated part of the right
lungs enters the left atrium. The proximal ventricle (Fig. 15.3).
part of the vein is gradually absorbed
and is incorporated into the w a l l of the
atrium. As a result of continued absorp- The right and left ventricles are formed by
tion of tributaries, four veins {two right partitioning of this chamber. From Fig. 15.10, note
and [ w o left) finally open into the atrium. that the bulbo-ventricular cavitv consists of:

X
Truncus Pulmonary trunk Aorta
^ arteriosus

A1
Fused atrio-
ventricular
cushions — Spiral septum
\ Atrium
I 2 1 _ Fused A.v.
Right cushions
atrioventricular /
orifice \ 7 7 \ , Left A.v.
v \ orifice

Common ^ ^ ^ ^ S \
ventricle ^ ! w" x^

Y
A B
Fig. 15.10 (A) Two parts of the ventricular chamber. Part 1 lies anterior to the atrio-ventricular orifice. Part 2
is conical and lies higher up. (B) This is a section across the ventricle in the plane XY, shown in
(A). Sections in the plane indicated by the arrow in (A) are shown in Fig. 15.17.
Cardiovascular System

(a) a dilated lower part (1 in Figure) t h a t 1/3 of bulbus cordis have merged into the primitive
communicates with the atria; and ventricle has to be subdivided into right and left
(b) a conical upper part (2 in Figure) commu- halves in such a way that:
nicating with the truncus arteriosus. Part
' 1 ' is derived from the proximal one-third fa) e a c h half c o m m u n i c a t e s w i t h t h e
of the bulbus cordis a n d the primitive corresponding atrium, and
ventricle, while part ' 2 ' is from the conus. (b) the right ventricle opens into the pulmonary
trunk and the left ventricle into the aorta.

Formation of Interventricular Septum This subdivision takes place as follows:

T h e cavity formed after the conus and proximal 1. A septum, called the interventricular septum.

Proliferation from
A.V. cushions Interventricular septum

Pulmonary trunk and


aorta from truncus arteriosus

Trabeculated part
of left ventricle
from primitive
ventricle

Fig. 15.11 Two stages in the formation of the ventricular septum. (B) and (D) correspond to (A) and (C)
respectively. (A) Bulbar septum grows d o w n from above, and interventricular septum grows
upwards from below. (C) and (D) The gap between the bulbar septum and the interventricular
septum is filled in by proliferation from A.V. cushions. For explanation of orientation of these
figures see Legend to Fig. 15.10.
Human Embryology

grows upwards from the floor of the bul no- form small masses of angioblastic tissue. This
ventricular cavity and divides the lower angioblastic tissue gives rise to endothelium and
dilated part of this cavity into right and left also to blood cells. The first blood vessels are
halves (Fig. 15.11A). It meets the fused atrio-
ventricular cushions (septum inter-
medium) and partially fuses with Amniotic cavity
them (Fig. 15.11C). On the external Pericardial cavity in
surface of the heart, the site of cardiogenic area
formation of the interventricular
septum corresponds to the bulbo-
ventricular sulcus (Fig. 15.14A).
2. Two ridges, termed the right and left
bulbar ridges, arise in the wall of the Splanchnopleuric
bulbo-ventricular cavity (in the part mesoderm
derived from the conus). These ridges
grow towards each other and fuse to Foregut
form a bulbar septum (Figs 15.11A,
B). The bulbar septum grows down- Splanchnopleuric
wards towards the interventricular mesoderm
septum but does not quite reach it,
with the result that a gap is left B
between the two.
3. The gap between the upper edge of
the interventricular septum and the
lower edge of the bulbar septum, is
filled by proliferation of tissue from Fused heart tubes
the atrio-ventricular cushions
(Fig. 15.1 ID).

Further Details about the


Development of the Heart
• Foregut
Introduction
Parietal serous Dorsal
The internal surfaces of the heart and of Pericardium mesogasium
all blood vessels are lined by a layer of Heart tube
flattened cells called endothelium. The (Endocardium)
endothelium is supported, on the outside,
by varying amount of muscle, and Myocardium
Pericardial"
connective tissue. All components of the cavity
Epicardium
heart and blood vessels, i.e. endothelium,
muscle and connective tissue are of
Fig. 15.12 Relationship of heart tubes to pericardial cavity:
mesodermal origin. Very early in the life
(A) Before formation of head fold. <B) After
of the embryo, mesenchyme differentiates, formation of head f o l d . (C) and (D) show the
over the yolk sac, in the connecting stalk process of invagination of the pericardial cavity
and in the body of the embryo itself, to by the single heart tube.
Cardiovascular System

Mesocardium Hole in
mesocardium

Truncus arteriosus Transverse sinus


of pericardium
\

Bulbo-ventricular Lower 1/3 of


loop bulbus cordis

Fig. 1 5 . 1 3 Schemes to show the f o l l o w i n g . (A) Heart tube suspended by mesocardium. {Bl Appearance of a
hole in mesocardium. (C) Disappearance of mesocardium resulting in formation of transverse
sinus of pericardium, in Figures (B) to (D) note (1) gradual freeing of heart tube from septum
transversum, and (2) folding of heart tube.

Truncus Opening of atrium


arteriosus

Lower 1 /3
of bulbus
cordis

B
Fig. 15.14 Scheme to show incorporation of conus (and proximal dilated part of bulbus cordis) into the
ventricle by disappearance of the bulbo-ventricular sulcus. Note that the opening of atrium into
ventricle gradually shifts to the centre of the posterior w a l l of the c o m m o n bulbo-ventricular
chamber. The part labelled 'conus' includes the dilated part of the bulbus cordis.
Human Embryology

derived from this e n d o t h e l i u m . T h e vessels heart come to lie dorsal to the pericardial cavity
rapidly proliferate in n u m b e r a n d become and ventral to the foregut (Fig. 5.13).
interconnected to form a vascular system. Soon We have seen that the endothelial heart tube
thereafter, a primitive heart begins to p u m p is derived from the splanchnopleuric mesoderm
blood through this network of vessels with the related to the pericardial cavity (Fig. 15.12A).
result that nutrition from the placenta and yolk After the formation of the head fold, this tube
sac can be m a d e available to the g r o w i n g lies dorsal to the pericardial cavity and ventral
embryo. The heart is, therefore, the first organ to the foregut (Fig. 15.12B). The tube n o w
of the body to start functioning. mvaginates the pericardial sac from the dorsal
We have seen that the pericardial cavity is side. As it d o e s s o , t h e s p l a n c h n o p l e u r i c
formed from the cranial, midline, part of the m e s o d e r m l i n i n g t h e d o r s a l side of t h e
intra-embryonic coelom (Figs 5.6, 5.11). With pericardial cavity proliferates to form a thick
the formation of the coelom, the intra-embryonic layer called the myoepicardial mantle (or
m e s o d e r m of t h e r e g i o n s p l i t s i n t o a epimyocardial mantle) (Figs 15.12C, D). When
somatopleuric layer adjoining the ectoderm (in the invagination is complete, the myoepicardial
the roof of the p e r i c a r d i a l cavity) a n d a mantle completely surrounds the heart tube. It
splanchnopleuric layer adjoining the endoderm gives rise to the cardiac muscle (myocardium)
(Fig. 5.6) a n d f o r m i n g t h e f l o o r of t h e and also to the visceral layer of pericardium
pericardial cavity. The heart develops from (epicardium). The parietal layer of pericardium
angioblastic tissue t h a t arises from this is derived from somatopleuric mesoderm.
s p l a n c h n o p l e u r i c m e s o d e r m , w h i c h is,
therefore, called the cardiogenic area. With the Exterior of the Heart
e s t a b l i s h m e n t of t h e h e a d f o l d , t h e The heart tube is, for some time, suspended
splanchnopleuric mesoderm and the developing from the dorsal wall of the pericardial cavity

Bulbo-ventricular
sulcus

Conus \ / " ~

A l~ venosus
Lower 1/3 of
butbus cordis

Fig. 15.15 Stages in establishment of external form of the heart.


Cardiovascular System

by two layers of pericardium that constitute the sinus. Simultaneously, the left horn of the
the dorsal mesocardium (Figs 15.12D, 15.13A). sinus venosus and its tributaries become much
This mesocardium soon disappears and the reduced in size, and the left horn now appears
heart tube lies free within the pericardial sac, to be just another tributary of the right half of
suspended by its two ends (Figs 15.13B, C). the sinus venosus (Fig. 15.16C). Two important
However, at this stage the caudal part of the results of these changes are that:
heart tube (atrium, sinus venosus) is embedded
within the substance of the septum rransversum. (a) The sinuatrial orifice, which was at first
The part of the heart tube lying within the situated in the middle of the posterior
pericardial cavity is thus made up of bulbus aspect of the atrial chamber, now comes
cordis and ventricle. to lie on the right side (Fig. 15.4A - C).
(b) The orifice, which was at first transverse,
This part of the tube g r o w s rapidly and,
now becomes vertical. We have already
therefore, becomes folded on itself to form a
seen t h a t t h e m a r g i n s of t h i s orifice
'LP shaped bulbo-ventricular loop {Fig. 15.13C).
form the right a n d left venous valves
Subsequently, as the atrium and sinus venosus
( F i g s l 5 . 1 6 C , 15.4D).
are freed from the septum transversum, they
come to lie behind and above the ventricle, and
the heart tube is now 'S' shaped (Fig. 15.13D). Some Facts about the Interatrial Septum
At this stage, the bulbus cordis and ventricle N o t e the following additional facts about the
are separated by a deep bulbo-ventricular sulcus formation of the interatrial septum.
(Figs 15.13D, 15.14). This sulcus gradually
becomes s h a l l o w e r so t h a t the c o n u s , the
p r o x i m a l p a r t of t h e b u l b u s c o r d i s a n d ( —\ Atrium
Body of sinus
t h e v e n t r i c l e c o m e t o form o n e c h a m b e r
(Fig. 15.14), which c o m m u n i c a t e s with the
venosus _.
—L_ 1\ ^ Left horn
truncus arteriosus. The atrial chamber which Right horn -.
-><^ /-~N " i ^ Common
:
lies behind the upper part of the ventricle and —\ ( ) (~ cardinal
) /"V 0 \ vein
of the truncus arteriosus, expands; and as it A
\ Umbilical vein
does so parts of it come to project forwards on ^ Vitelline vein
either side of the truncus. As a result of these
changes the exterior of the heart assumes its
definitive shape (Fig. 15.15).
Opening from
atrium to sinus
1
\
)
J Left horn and
venosus \ < ^ „fts tributaries
shifts to right. ) V" retrogress.
Fate of Sinus Venosus
The sinus venosus and the atrial chamber are
at first in open communication with each other
Opening becomes
B 4r*
(Fig. 15.16A). However, they become partially
narrow and is i /
separated by grooves that appear on the lateral guarded by right
wall of the heart tube, at the junction of these and left venous
valves. y? = = ^ - ^ 1 8 now
t w o c h a m b e r s . T h e right g r o o v e r e m a i n s
/ [ Nr a tributary of
shallow but the left one becomes very deep -/ J right horn.
(Figs 15.16B, C) with the result that the left
part of the sinus venosus becomes completely c -\r^i •
separated from the atrial chamber. Its blood Fig. 15.16 Retrogression of the left horn of sinus
now enters the atrium through the right half of
Human-Embryology

T h e lower edge of the septum secundum the septum, rather than by active growth of the
(crista dividens) is chick and firm. In contrast, septum itself.
the upper edge of the septum primum (that forms The membranous part of the interventricular
the lower boundary of the foramen secundum) septum is divisible into an anterior part, which
is thin and mobile like a flap. When blood tends separates the right and left ventricles, and a
to flow from the right to the left atrium, this posterior part which separates the left ventricle
thin flap moves away and there is no obstruction from the r i g h t a t r i u m (also c a l l e d atrio-
to b l o o d flow. H o w e v e r , w h e n t h e r e is a ventricular septum). The anterior part is derived
tendency for blood to flow from left to right from t h e p r o l i f e r a t i o n of tissue from the
this flap comes into apposition with the septum endocardial cushions, as described above. The
secundum and closes the opening. After birth, derivation of the posterior part is shown in
the left atrium begins to receive blood from the Fig. 15.17. It will be seen that the interatrial
lungs and the pressure within this chamber and interventricular septa do not meet the atrio-
becomes greater than that in the right atrium. ventricular cushions in the same line. As a result,
This causes a closure of the foramen ovale, a part of these c u s h i o n s separates the left
which is soon permanently obliterated by fusion ventricle from the right atrium. This part of
of the two flaps. the atrio-ventricular cushions forms the posterior
part of the membranous septum.
Some Additional Facts about the
Interventricular Septum
Valves of the Heart
The interventricular septum is probably formed
more by d o w n w a r d enlargement of the right The mitral and tricuspid valves are formed by
and left ventricular cavities on either side of proliferation of connective tissue under the

Septum secundum

Septum primum
Fused A.V. cushions
(Septum intermedium)

Proliferation from
A.V. cushions joining

Interventricular
septum
Two components that
form the membranous
part of interventricular
septum

Fig. 15.17 In (A) note that the interatrial and interventricular septa d o not meet the atrio-ventricular cushions
in the same plane. In (B) note that the membranous part of the interventricular septum is made up
of (i) the original. A.V. cushion between the attachment of the interatrial and interventricular septa,
and (ii) the endocardial proliferation from these cushions. The first part separates the left ventricle
from the right atrium w h i l e the second part separates the t w o ventricles. The tricuspid valve is
attached to the membranous septum at the junction of these parts. These figures are sections in
the plane indicated by an arrow in Fig. 15.10A.
Cardiovascular System

endocardium of the left and right atrio- After fusion of the two tubes, it lies in the sinus
ventricular canals. venosus. When the sinus venosus is incorporated
The pulmonary and aortic valves are derived into the right atrium, it comes to lie near the
from endocardial cushions that are formed at opening of the superior vena cava.
the junction of the truncus arteriosus and the The atrio-ventricular node and the atrio-
conus (Fig. 15.18A}. Two cushions, right and ventricular bundle form in the left wall of the
left, appear in the wall of the conus. They grow sinus venosus, and in the atrio-ventricular canal.
and fuse with each other (Fig. 15.18B). With After the sinus venosus is absorbed into the right
the separation of the aortic and pulmonary atrium, the atrio-ventricular node comes to lie
openings, the right and left cushions are each near the interatrial septum.
subdivided into two parts, one part going to
each orifice (Fig. 15.18C). Simultaneously, two PERICARDIAL CAVITY
more cushions, anterior and posterior appear.
As a result, the aortic and pulmonary openings We have already noted several important facts
each have three cushions, from which three about the development of the pericardial cavity,
cusps of the corresponding valve develop. and these may be briefly recapitulated as
follows:
The pulmonary valve is at first ventral to
the aortic valve (Fig. 15.18C). Subsequently, 1. The pericardial cavity is a derivative of
there is a rotation so that the pulmonary valve the part of the intra-embryonic coelom that
comes to lie ventral and to the left of the aortic lies in the midline, cranial to the
valve (Fig. 15.18D). It is only after this rotation prochordal plate (Fig. 5.11).
that the cusps acquire their definitive 2. After the formation of the head fold, the
relationships (Pulmonary trunk: one posterior, pericardial cavity comes to lie on the
two anterior; Aorta: one anterior, two posterior). ventral side of the body of the embryo
(Fig. 5.14).
Conducting System of the Heart
3. The heart tube invaginates the
At the stage when there are two heart tubes, a pericardial sac from the dorsal aspect
pacemaker (which later forms the sinoatrial (Figsl5.12C,D).
node) lies in the caudal part of the left tube. 4. The parietal layer of the serous

A B
Posterior

I I <&

Pulmonary
valve

Fig. 15.18 Formation of aortic and pulmonary valves. Note that the vessels undergo an anticlockwise
rotation (compare axis XY in (C) and (D)). It is only after this rotation that the cusps of the aortic
and pulmonary valves acquire their definitive position.
Human Embryology

pericardium and the fibrous pericardium, After disappearance of the dorsal


are derived from the somatopleuric mesocardium, the visceral and parietal
mesoderm lining the ventral side of the layers of pericardium are in continuity only
pericardial cavity (Figs 15.12A, D). at the arterial and venous ends of the heart
The visceral serous pericardium is derived tube (Figs 15.19A, B, D, E).
from the splanchnopleuric mesoderm With the folding of the heart tube, rhe
lining the dorsal side of the pericardial arterial and venous ends come closer to
cavity (Fig. 15.12D). each other. The space between them
The heart tube is initially suspended within becomes the transverse sinus of
the pericardial cavity by the dorsal pericardium (Figs 15.19C, F).
mesocardium, which soon disappears A number of blood vessels are formed at
(Fig. 15.13). the two ends of the heart tube. At the
We may now consider certain additional arterial end, these are the aorta and the
facts. pulmonary trunk. At the venous end, these

- Arterial end of heart tube -


Mesocardium

Venous end
Line of reflection of
' of heart tube
visceral pericardium

- Transverse sinus -

- Truncus arteriosus

E
Sinus venosus

- Transverse sinus -

•<©
Fig. 15.19 Schemes showing the relationship of the heart tube to the pericardial sac. (A), (B) and (C) are
lateral views w h i l e (D), (E), (F) show the dorsal aspect of the interior of the pericardial sac at
corresponding stages. Disappearance of the mesocardium leads to formation of the transverse
sinus of pericardium. Note that with the folding of the heart tube, the arterial and venous ends of
the heart tube are brought closer together, and the transverse sinus comes to lie between them.
Cardiovascular System

are the superior vena cava, inferior vena (b) Ectopia cordis: T h e heart lies exposed, o n
cava and four pulmonary veins (Fig. 15.20A). the front of the chest, and can be seen from
10. The definitive reflections of the pericar- the outside, due to defective development
dium are formed merely by rearrangement of the chest wall.
of these vessels, as shown in Fig. 15.20B.
Rearrangement of the veins at the venous Atresia or Stenosis
end results in the formation of an isolated Any of the orifices of the heart may have too
pouch of pericardium, in relation to the n a r r o w an opening (stenosis), or none at all
four pulmonary veins. This is the oblique (atresia). The aortic and pulmonary passages
sinus of pericardium. may also show supravalvular, or subvalvular,
stenosis (Fig. 15.23). Alternatively, the openings
Congenital Anomalies of the Heart
may be too large as a result of which the valves
Anomalies of Position become incompetent.
In pulmonary stenosis the foramen ovale and
(a) Dextrocardia: The chambers and blood
the ductus arteriosus remain patent. In aortic
vessels of the heart are reversed from side
stenosis also, the ductus arteriosus is patent and
to side, i.e. all structures that normally lie
blood flows into the aorta through it.
on the right side are on the left and vice
versa (Fig. 15.21). This may be a part of
Abnormal G r o w t h
the condition called situs inversus, in which
all o r g a n s a r e t r a n s p o s e d . W h e n There may be accessory cusps in the valves.
dextrocardia is not a part of situs inversus, Congenital tumours may be formed. The left
it is usually accompanied by anomalies a t r i u m m a y be p a r t i a l l y s u b d i v i d e d by a
of the chambers of the heart and of the transverse septum. The myocardium may be
great vessels. poorly developed (hypoplasia).

Line of reflection of visceral


to parietal pericardium Pulmonary trunk

Pulmonary trunk

Inferior Inferior
vena cava vena cava
Pulmonary veins Parietal pericardium
B
Fig. 15.20 Scheme to show that the oblique sinus of pericardium is established by rearrangement of veins
entering the heart. The sinus is indicated by the lower arrow in (B). The upper arrow indicates the
transverse sinus.
Human Embryology

Aorta -JrT"*>~--^

Pulmonary
£P
^f/(
\
Vv
^v [
/ Superior
vena cava

trunk \/\

Apex ^ • " \ s ^ •> I Inferior Fig. 15,23 Types of aortic stenosis. (A) Valvular.
vena cava (B) Supravalvular. (C) Infravalvular.

Fig. 15.21 Dextrocardia. The chambers and large cushions, as a result of which the fora-
blood vessels show right-left reversal. men primum persists (Fig. 15.24A).
This osteum primum defect can
also be caused by defective formation
of atrio-ventricular endocardial
cushions.
(b) The septum secundum may fail to
develop as a result of which the
foramen secundum remains wide
open [osteum secundum defect;
Fig. 15.24B).
(c) The septum primum and secundum
may develop normally but the oblique
valvular passage between them may
remain patent {patent foramen ovale;
Fig. 15.24C). The patency is signifi-
cant only if there is shunt of blood
Fig. 15.22 Patent truncus arteriosus. The ascending through it. In many cases a probe can
aorta and pulmonary trunk are be passed through the oblique slit
represented by a single channel that (probe patency) but there is no shunt.
opens into both ventricles. (d) Occasionally, there is premature
closure of the foramen ovale (i.e.
before birth). As a result, the right
Defective Formation of Septa
atrium and ventricle undergo great
This results in the formation of abnormal hypertrophy, while the left side of the
passages. heart is underdeveloped.
2. Interventricular septal defects: These may
1. Interatrial septal defects: These may be of
be seen either in the membranous or in
three types.
the muscular part of the septum
(a) The septum primum may fail to reach (Fig. 15.24D). These are the most common
the atrio-ventricular endocardial congenital anomalies of the heart.
Cardiovascular System

Fig. 15.24 Septal defects: (A) Septum primum defect, (B) Septum secundum defect, (C) Patent foramen ovale,
(D) Interventricular septum defect.

3. Defects of the spiral septum: The spiral Combined Defects


septum may n o t be formed at all. This
c o n d i t i o n is c a l l e d patent truncus Two or more of the defects may coexist. One
arteriosus (Fig. 15.22). Partial absence of classically recognized condition of this type is
the s e p t u m leads to c o m m u n i c a t i o n s k n o w n as Fallot's tetrology (Fig. 15.25). It
(shunts) between the aorta and the consists of the following:
pulmonary trunk.
4. Atrio-ventricular canal defect or persistent 1. Interventricular septal defect;
atrio-ventricular canal: Defective 2. Aorta overriding the free upper edge of the
formation of the atrio-ventricular cushions ventricular septum;
may lead to a condition in which all four 3. Pulmonary stenosis;
c h a m b e r s of t h e h e a r t m a y i n t e r - 4. Hypertrophy of the right ventricle.
communicate. The interatrial and
interventricular septa are incomplete (as Other Defects
the normal contributions to these septa
from the endocardial cushions are lacking). 1. T h e p e r i c a r d i u m may be p a r t i a l l y or
completely absent.
2. There may be congenital defects in the
If fusion of endocardial cushions is too far
conducting system of the heart.
to the right, it causes tricuspid atresia. As such
c u s h i o n s a r e n o t in a l i g n m e n t w i t h t h e
Anomalies of Relationship of Chambers
interventricular septum, the upper part of the
latter is defective. With tricuspid atresia there to Great Vessels
is increased pressure in the right atrium, as a 1. Transposition of great vessels: The aorta
result of which the foramen ovale fails to close. arises from the right ventricle and the
Defective formation of septa, if marked, can pulmonary trunk from the left ventricle.
lead t o a two-chambered heart {Cor biloculare) 2. Taussig-Bing syndrome: T h e aorta arises
in which there is one c o m m o n ventricle and from the right ventricle; and the pulmonary
one c o m m o n atrium. Alternatively, a three- trunk overrides both the right and left
chambered heart {Cortriloculare) may be seen; ventricles, there being an interventricular
it may consist of a single ventricle with two septal defect.
atria or of a single atrium with two ventricles 3. The superior or inferior vena cava may
[Cor triloculare biventriculare). end in the left atrium.
Human Embryology

to the foregut {ventral aorta), an arched portion


lying in the first pharyngeal arch, and a dorsal
portion lying dorsal to the gut {dorsal aorta)
(Fig. 15.26A).
After the fusion of the two endocardial tubes,
1. Narrow pulmonary the two ventral aortae partially fuse to form the
trunk
aorticsac, the unfused parts remaining as the right
2. Aorta overriding and left horns of the sac (Fig. 15.26B).
ventricular septum
Successive arterial arches now appear in the
3. Interventricular second to sixth pharyngeal arches, each being
septal defect
4. Hypertropy of wall
connected ventrally to the right or left horn of the
of right ventricle aortic sac and dorsally to the dorsal aorta
(Fig. 15.27). The major arteries of the head and
neck, and of the thorax, are derived from these
arches as follows:
Fig. 15.25 Four features that constitute Fallot's
tetrology.
1. The greater part of the first and second arch
arteries disappear (Fig. 15.28A).
4. The pulmonary veins may end in the right
atrium or in one of its tributaries.
In adult life, the first arch artery is represented
by the maxillary artery. The second arch artery
THE ARTERIES persists for some part of fetal life as the stapedial
artery: it may contribute to the formation of
Pharyngeal Arch Arteries
the external carotid artery.
and their Fate
The first arteries to appear in the embryo are the 2. The fifth arch artery also disappears
right and left primitive aortae. They are continuous (Fig. 1.5.28A).
with the two endocardial heart tubes. Each 3. The aortic sac is, therefore, now connected
primitive aorta consists of a portion lying ventral only with the arteries of the third, fourth

Right and left horns


ot aortic sac
1st aortic arch 1st aortic arch

Aortic sac formed by


fusion of ventral aortae

-Truncus arteriosus

Single heart tube

Fig. 15.26 Relation of first aortic arch to heart tubes. (A) Before fusion of heart tubes. (B) After fusion.
Cardiovascular System

(b) The portion of the dorsal aorta, which


First arch lies between the attachment of the third
Second arch \ and fourth arch arteries [ductus
caroticus), disappears on both sides
\ Third arch //< (Fig. 15.28B).
\ \ Fourth / / x (c) The portion of the right dorsal aorta,
\ \ \ arch ~~~J/// w h i c h lies b e t w e e n t h e p o i n t of
attachment of the fourth arch artery and
the point of fusion of the two dorsal
aortae, disappears (Fig. 15.28B).
(d) Each sixth arch artery gives off an
^ X ^ Fifth artery to the developing lung bud. On
f<l arch the right side, the portion of the sixth
Aortic -^b
sac \ \ . ^FSixth arch artery that lies between this bud
^ ^ F arch and the dorsal aorta, disappears. On
Truncus . W Left horn of
the left side, this part remains patent
. arteriosus A * aortic sac
Right dorsal -*^ and forms the ductus arteriosus. The
aorta ducts arteriosus carries most of the
blood from the right ventricle to the
dorsal aorta. It is obliterated after birth
a n d is then seen as the ligamentutn
Fig. 15.27 Aortic arches. Each arch connects the
aortic sac to the dorsal aorta. Note that arteriosum.
actually all arches are never present at (e) Each third arch artery gives off a bud
tfit- V i i n r tiriif. Tilt- liiM <IIKI M-< UIKI that grows cranially to form the external
arches have retrogressed by the time the carotid artery (Figs 15.29A, 15.31 A}.
sixth appears. (f) The dorsal aortae give off a series of
lateral intersegmental branches to the
and sixth arches. The third and fourth arch body wall. One of these, the seventh
arteries open into the ventral part, and the cervical intersegmental artery supplies
sixth arch artery into the dorsal part, of the the u p p e r limb bud. It c o m e s t o be
aortic sac. T h e spiral s e p t u m , which is attached to the dorsal aorta near the
formed in the truncus arteriosus, extends into attachment of the fourth arch artery
the aortic sac; and it fuses with its posterior (Fig. 15.29A).
wall in such a way t h a t blood from the
pulmonary trunk passes only into the sixth The development of the main arteries can now
arch artery, while that from the ascending be summarized as follows:
aorta passes into the third and fourth arch
arteries (Fig. 15.28B). (a) The ascending aorta and the pulmonary
4. Several c h a n g e s n o w take place in the trunk are formed from the truncus arteriosus
arterial arches to produce the adult pattern (Fig. 15.28B).
as follows: (b) The arch of the aorta is derived from the
ventral part of the aortic sac (1), its left horn
(a) The two dorsal aortae grow cranially, (2), a n d t h e left f o u r t h a r c h a r t e r y (3)
beyond the point of attachment of the (Fig. 15.29A).
first arch artery (Fig. 15.28B). (c) The descending aorta is derived from the
Human Embryology

First & second arch


arteries disappear.

. 7th cervical
intersegmental a.

5th arch disappears

x
Recurrent laryngeal n.
Branch to lung bud
6th arch

Truncus arteriosus

Dorsal aorta between


3rd & 4th arches disappears
on both sides.

- 7th cervical inter-


segmental artery
forms left subclavian a.

Parts of right 6th aortic arch


6th aortic arch
and of right dorsal aorta Pulmonary trunk &
disappear. Ascending aorta formed
by subdivision of the
truncus arteriosus

Fig. 15.28 Fate of aortic arches: (A) Disappearance of 1 st, 2nd and 5th arches.
(B) Disappearance of ductus caroticus o n both sides, and of part of right dorsal
aorta. Part of the right 6th arch also disappears.
Cardiovascular System

Fig. 15.29 (A) The arch of the aorta is derived from (1) the aortic sac, (2) its left horn and (3) the left 4th arch
artery. (B) The descending aorta is derived from (1) the left dorsal aorta and (2) fused dorsal aortae.
(C) The brachiocephalic artery is derived from the right horn of the aortic sac.

left dorsal aorta, below the attachment of (g> As already mentioned, the external carotid
fourth arch artery (1), along with the fused artery arises as a bud from the third arch
median vessel (2) (Fig. 15.29B). artery (Fig. 15.31 A).
(d| The brachiocephalic artery is formed by the (h) The pulmonary arteries are derived from
right horn of the aortic sac (Fig. 15.29C). the part of the sixth arch arteries lying
(e) The proximal part of the right subclavian between the p u l m o n a r y t r u n k and the
artery is derived from the right fourth arch branches t o the lung buds (Fig. 15.31B).
arrery (1), the remaining p;irr of the artery
being derived from the seventh cervical As already stated, the part of the left sixth arch
intersegmental artery (2). On the left side, artery, between the branch to the lung bud and
the subclavian artery is derived entirely the aorta, forms the ductus arteriosus (Fig. 15.31C).
from the seventh cervical intersegmental
artery (3), which arises from the dorsal aorta The relationship of the main nerves of the head
opposite the attachment of the fourth arch and neck to the arteries, can be explained on
artery (Fig. 15.30A). the basis of the development of the arteries.
(f) The common carotid artery is derived, on The nerves of the pharyngeal arches are, at
either side, from part of the third arch first, lateral to the corresponding arteries. T h e
artery, proximal to the external carotid bud nerves of the first, second and third arches (V,
(Fig. 15.30B). The interna! carotid artery is VII & IX) retain their lateral positions. The
formed by the portion of the third arch disappearance of the ductus caroticus, enables
artery distal to the bud (1), along with the the nerve of the fourth arch (superior laryngeal)
original d o r s a l a o r t a c r a n i a l to the to move medially, a n d it comes to lie deep to
a t t a c h m e n t of the third arch artery (2) the main arteries of the neck.
(Fig. 15.30C). T h e nerve of the sixth a r c h ( r e c u r r e n t
As the right third and fourth arch arteries laryngeal), is at first caudal to the artery of
arise from the right horn of the aortic sac, this arch (Fig. 15.32A). With the disappearance
the common carotid and subclavian arteries of part of the sixth arch artery, on the right
become branches of the brachiocephalic side, the nerve moves cranially and comes into
artery. relationship with the right fourth arch artery
Human Embryology

Fig. 15.30 (A) The right subclavian artery is derived (1) from the right 4th arch artery and {2) from the right
7th cervical intersegmental artery. The left subclavian artery is formed only from the left 7th
cervical intersegmental artery. (Bl The c o m m o n carotid artery is derived from the proximal part of
the 3rd arch artery. (C) The internal carotid artery is derived f r o m (1) distal part of the 3rd arch
artery and (2) dorsal aorta (cranial-most part).

Fig. 15.31 (A) The external carotid artery arises as a bud f r o m the 3rd arch artery. (B) The pulmonary arteries
arise from the 6th arch arteries. (C) The ductus arteriosus is derived from part of the left 6th arch
artery.

(subclavian) (Figs 15.32B, C). O n the left side, normal arterial pattern is dependent upon the
it retains its relationship to that part of the sixth disappearance of some parts of the pharyngeal
arch which forms the ductus arteriosus. With arch a r t e r i e s . O c c a s i o n a l l y this process is
the elongation of the neck, and the descent of disturbed in that:
the heart, these nerves are dragged d o w n w a r d s
and, therefore, have to follow a recurrent course 1. Some parts that normally disappear may
back to the larynx. persist; and
2. Some p a r t s t h a t normally persist may
ANOMALOUS DEVELOPMENT OF disappear.
PHARYNGEAL ARCH ARTERIES As a result, several anomalies may be
We have seen t h a t the d e v e l o p m e n t of the produced. Some of these are as follows:
Cardiovascular System

Fig. 15.32 Relationship of the vagus and recurrent laryngeal nerves to the aortic arches. For explanation see
text.

Summary of the adult derivatives of truncus arteriosus, aortic sac and aortic arches

Adult Derivatives Embryological Structures


1. Ascending aorta Truncus arteriosus
2. Arch of aorta Aortic sac, left horn of aortic sac and left 4th arch artery
3. Descending aorta Left dorsal aorta and fused dorsal aorta
4. Brachiocephalic artery Right horn of aortic sac
5. Right subclavian artery Right 4th arch artery and 7th cervical intersegmental artery
6. Left subclavian artery Left 7th cervical intersegmental artery
7. Common carotid artery Proximal part of 3rd arch artery
8. Internal carotid artery Distal part of 3rd arch artery and cervical part of dorsal aorta.
9. External carotid artery As a bud from 3rd arch artery
10. Pulmonary trunk Truncus arteriosus
11. Pulmonary artery Part of 6th arch artery
12. Ductus arteriosus Part of left 6th arch artery between lung bud and aorta.

(a) Double aortic arch (Fig. 15.33A).The (d) The right subclavian artery may arise
arterial ring can compress the trachea as the last branch of the aortic arch
and oesophagus. (Fig. 15.33C). Such an artery runs to
(b) Right aortic arch (Fig. 1 5 . 3 3 B ) . the r i g h t b e h i n d t h e o e s o p h a g u s
Normally the aortic arch disappears (Fig. 15.34). Along with the aorta this
on the right side and persists on the artery forms an arterial ring enclosing
left. Reversal of this a r r a n g e m e n t the trachea and oesophagus. The ring
results in a right aortic arch. may press upon a n d obstruct these
(c) T h e ductus arteriosus, w h i c h is tubes. In this abnormality, the right
normally occluded soon after birth, recurrent laryngeal nerve does n o t
may remain patent [patent ductus hook around the subclavian artery. It
arteriosus). passes directly t o the l a r y n x . An
Human Embryology

subclavian artery arises


from the distal segment
w h i c h receives b l o o d
through a patent ductus
arteriosus.
(g) Some other anomalies of
the mode of origin of the
branches of the arch of
the aorta are illustrated
in Fig. 15.35.
(h) T h e aorta may show a
localized n a r r o w i n g of
its lumen, leading to par-
tial or even complete o b -
struction to blood flow.
This condition is called
coarctation of the aorta.
C o a r c t a t i o n is m o s t
frequently seen near the
attachment of the ductus
arteriosus to the aorta. It
may be (1) distal to the
attachment of the ductus
(postductal) or (2) proxi-
mal to the a t t a c h m e n t
(preductal), in w h i c h
case the right ventricle
supplies the distal part of
t h e b o d y t h r o u g h the
Persistence of
Anomalous right subclavian a. ductus caroticus ductus arteriosus.
When coarctation is postduc-
Fig. 15.33 Anomalies associated with the development of aortic
tal, numerous anastomoses
arches. The abnormal vessels are shown in red,
are established between
branches of the aorta taking
arterial ring can also be formed if the
origin above the constriction and those
dorsal aorta persists on both sides.
arising below this level. Coarctation is said
(e) The ductus caroticus may persist. As
to be a result of the process of obliteration
a result, the left internal carotid arises
of the ductus arteriosus extending into the
directly from the aortic arch, and the
aorta. It can also occur as an abnormality
right internal carotid from the sub-
in the vessel wall.
clavian (Fig. 15.33D).
(f) Interrupted aortic arch; A segment of
Development of Other Arteries
the aortic arch, may be missing. T h e
ascending aorta ends by supplying the The primitive dorsal aortae give off three groups
left c o m m o n carotid artery. The left of branches (Fig. 15.36). These are as follows:
Cardiovascular System

2. The lateral or intermediate splanchnic


arteries supply structures developing from
the intermediate mesoderm. These persist
as the renal, suprarenal, phrenic and
spermatic or ovarian arteries.
Left subclavian
3. The dorsolateral (somatic intersegmental)
artery branches run between two adjacent
segments. They retain their original
intersegmental arrangement in the thoracic
and lumbar regions where they can be
Oesophagus recognized as the intercostal and lumbar
arteries.

Each dorsolateral artery divides into a


dorsal and a ventral division. The ventral
division gives off a lateral branch that is most
conspicuous in the region of the limb buds.
Fig. 15.34 Relationship of abnormal right The dorsal division runs dorsally and
subclavian artery to the oesophagus and supplies the muscles of the back. Each dorsal
to the arch of the aorta. division gives off a spinal branch that runs
medially to supply the spinal cord.
1. The ventral splanchnic arteries supply the The branches of the dorsolateral arteries of
gut. Most of these arteries disappear but successive segments become interconnected by
three arteries, the coeliac, superior the formation of longitudinal anastomoses. In
mesenteric and inferior mesenteric, remain the neck, the dorsal branches are connected by
to supply the infradiaphragmatic part of anastomoses that are formed in three situations
the foregut, the midgut and the hindgut (Fig. 15.37).
respectively. Other remnants of these
vessels are the bronchial and oesophageal fa) Pre-costal, in front of the necks of the ribs
arteries. (or costal elements).

Fig. 15.35 Anomalies in the pattern of the main branches of the arch of the aorta. (A) Left common carotid
arising from brachiocephalic artery. (B) Left subclavian and left common carotid arising by a
common stem (left brachiocephalic). (C) Left vertebral artery arising directly from arch of aorta.
Human Embryology

Spinal branch

">Y i i Dorsal division

1 Somatic intersegmental
/ branch

^fj Ventral - ^ \
^ \ . division \

^ ^ " \ Lateral branch

/
\r
Ventral splanchnic
branches

Bronchial
Oesophageal
0 Lateral splanchnic
branches

Renal
Coeliac Gonadal
Sup. mesenteric Suprarenal
Inf. mesenteric Phrenic

Fig. 15.36 Basic branching pattern of the embryonic dorsal aorta.

Spinal branch • / Post-costal anastomoses = f > Vertebral artery

.—-•' !.•' V - - ' * Dorsal division

j \ f \ Ventral division

/ / Vre-costal Thyrocervical trunk


y / anastomoses > Ascending cervical a.
_^^ / Sup. intercostal a.

J Somatic intersegmental artery

Fig. 15.37 Sites of vertical anastomoses between branches of dorsal aorta. The fate of the
anastomoses is also shown.
Cardiovascular System

(b) Post-costal, between the costal elements of the seventh cervical intersegmental artery.
and the transverse processes. The main stem of this artery becomes the
(c) Post-transverse, behind the transverse subclavian artery. Like other dorsolateral
processes. arteries, it divides into dorsal, ventral and
lateral divisions. The dorsal division forms the
The pre-costal anastomoses persist as the stem of the vertebral artery (see below). The
thyrocervical trunk, the ascending cervical artery lateral division grows into the upper limb
and the superior intercostal artery. The post- forming the axillary and brachial arteries. The
costal anastomoses form the greater part of the ventral division forms the stem of the internal
vertebral artery. The post-transverse anasto- thoracic (mammary) artery.
moses remain as the deep cervical artery.
The ventral divisions of the somatic Development of the Vertebral Artery
intersegmental arteries are interconnected by (Fig. 15-38)
anastomoses, that are formed on the ventral 1. The first part of the artery, from its origin
aspect of the body wall, near the midline to the point of entry into the foramen
(Fig. 15.39). These form the internal thoracic, transversarium of the sixth cervical
superior epigastric and inferior epigastric vertebra, is formed by the dorsal division
arteries. of the seventh cervical intersegmental
At this stage, special mention must be made artery.

Spinal branch
Somatic
intersegmental.
branches Dorsal div.

Ventral div.

Internal carotid

3rd arch artery Post-costal


anastomoses

External carotid

7th cervical
intersegmental artery
4th arch artery

Fig. 15.38 Development of the vertebral artery. In (B) the part labelled 1 is derived from the dorsal division of
the seventh cervical intersegmental artery; 2 from the post-costal anastomoses; and 3 from the
spinal branch of the first cervical intersegmental artery.
Human Embryology

2. The vertical part (second part), lying in to the segments from which the limb buds take
the foramina transversaria, is formed from origin. These vessels form an arterial plexus.
the post-costal anastomoses between the However, each limb soon comes to have one
first to sixth cervical intersegmental axis artery that runs along the central axis of
arteries. the limb. Other arteries, that are formed as
3. The third (horizontal) part, running branches of the axis artery, or as new formations,
transversely on the arch of the atlas, is later take over a considerable part of the arterial
derived from the spina! branch of the first supply as a result of which much of the original
cervical intersegmental artery. axis artery may disappear.
The axis artery of the upper limb is formed
Development of the Internal Thoracic by the seventh cervical intersegmental artery.
Artery (Fig. 15.39) It persists as the axillary, brachial and anterior
interosseous arteries, and as the deep palmar
1. The main stem of the artery is formed by arch. The radial and ulnar arteries appear late
the ventral division of the seventh cervical in development.
intersegmental artery. The left subclavian artery represents the main
2. The vertical part of the artery (including stem of the seventh cervical intersegmental
its superior epigastric branch) is derived artery, and the proximal part of its lateral
from the ventral anastomoses between the division (Fig. 15.39). This explains the origin
ventral divisions of the thoracic inter- of the vertebral (dorsal division) and internal
segmental arteries (intercostal arteries). thoracic (ventral division) arteries from it. The
Development of the Arteries of the Limbs distal part of the right subclavian artery has a
similar origin, but its proximal part is derived,
The limbs are supplied by lateral branches of as already noted, from the right fourth aortic
the somatic intersegmental arteries, that belong arch.

7th cervical intersegmental artery Subclavian artery

/ , Dorsal division -Vertebral artery

- ~ - - x ' T ^ - * Ventral division

^ J r ^ ^ Lateral branch From ventral division


of 7th cervical
Ventral division intersegmental artery

The original thoracic


Ventral divisions •intersegmental
anastomoses • of thoracic arteries now appear
intersegmental as branches of
arteries internal thoracic
(2) artery
From ventral
anastomoses

B
Fig. 15.39 Development of the internal thoracic artery.
Cardiovascular System

The axis artery of the lower limb is derived (c) the part of the popliteal artery that lies
from the fifth lumbar intersegmental artery. It above the level of the popliteus muscle,
is seen as a branch of the internal iliac and (d) the distal part of the peroneal artery, and
runs on the dorsal aspect of the limb. The (e) part of the plantar arch.
femoral artery is a new vessel formed on the
ventral aspect of the thigh. Proximally it gets Umbilical Artery
linked above with the external iliac (which is a Before the fusion of the two dorsal aortae, the
branch of the axis artery) and below with the umbilical arteries appear as continuations of
popliteal artery. their distal ends (Fig. 15.40A). After fusion of
In the adult, the original axis artery is the dorsal aortae, they appear as lateral branches
represented by: of the single dorsal aorta (Fig. 15.40B).
Subsequently, each umbilical artery gets linked
(a) the inferior gluteal artery, up with that part of the fifth lumbar
(b) a small artery accompanying the sciatic intersegmental artery which forms the internal
nerve, iliac artery (Fig. 15.40C). The part of the

This part
disappears.

External iliac artery


Internal iliac artery
(from 5th lumbar I.S.A.)
Umbilical artery

Fig. 15.40 Development of the u m b i l i c a l artery. (A) U m b i l i c a l arteries are seen as continuations o f the right
and left dorsal aortae, before their fusion. (B) After fusion of dorsal aortae, the umbilical arteries
appear as lateral branches of the aorta. They cross the 5th lumbar intersegmental artery (I.S.A.).
(C) U m b i l i c a l arteries establish anastomoses w i t h the 5th lumbar intersegmental artery. (D) The
part of the umbilical artery between the dorsal aorta and the 5th lumbar intersegmental artery
disappears; and the umbilical artery is n o w seen as a branch of the latter. (E) The 5th lumbar
intersegmental artery forms the c o m m o n iliac and internal iliac arteries; and the umbilical is n o w
seen as a branch of the internal iliac.
Human Embryology

umbilical artery, which lies between the aorta Visceral Veins


and the anastomosis with the internal iliac,
These are:
disappears so that the umbilical artery is
now seen as a branch of the internal iliac 1. Right and left vitelline veins from the yolk
{Fig. 15.40 D,E). In post-natal life, the proximal sac. These are also called omphalome-
part of the umbilical artery becomes the senteric veins.
superior vesical artery, while its distal part is 2. Right and left umbilical veins from the
obliterated to form the medial umbilical placenta.
ligament. The umbilical and vitelline veins open into the
corresponding horn of the sinus venosus
VEINS (Fig. 15.41A). The parts of these veins that are
nearest to the heart are embedded in the septum
The main veins of the embryo may be divided transversum. These veins undergo considerable
into two groups, visceral and somatic. changes as follows:

Fig. 15.41 U m b i l i c a l and vitelline veins. (A) Note the umbilical and vitelline veins passing through the
septum transversum to reach the sinus venosus. (B) G r o w t h of liver cells w i t h i n the septum trans-
versum breaks up part of the umbilical and vitelline veins into capillaries. Blood reaching the liver
through the umbilical and vitelline veins n o w goes to the heart through the right and left hepato-
cardiac channels. ( Q Left hepatocardiac. channel disappears. (D) Right hepatocardiac channel
(which later forms part of the inferior vena cava) now drains the liver. Right umbilical vein
disappears. All b l o o d from the placenta n o w reaches the liver through the left umbilical vein.
Formation of ductus venosus short circuits this b l o o d to the right hepatocardiac channel.
Cardiovascular System

(a) With the development of the liver, in the septum (c) The right umbilical vein disappears and all
transversum, the proximal parts of the vitelline blood from the placenta n o w reaches the
and umbilical veins become broken up into developing liver through the left vein (N.B.
numerous small channels that contribute to The left vein is 'left') (Fig. 15.41D). In order
the sinusoids of the liver. These sinusoids to facilitate the passage of this blood through
drain into the sinus venosus, through the the liver, some of the sinusoids enlarge to create
persisting terminal parts of the vitelline a direct passage connecting the left umbilical
veins that are n o w called the right and left vein t o the right hepatocardiac channel.
hepatocardiac channels (Fig. 15.41B). The This passage is called the ductus venosus.
proximal parts of the umbilical veins lose (d) While these changes are occurring within
their communications with the sinus venosus. the liver, the parts of the right and left
(b) M e a n w h i l e , t h e left h o r n of the sinus vitelline that lie outside the substance of the
venosus undergoes retrogression and as a liver u n d e r g o alterations leading to the
r e s u l t t h e left h e p a t o c a r d i a c c h a n n e l formation of the portal vein.
disappears. All blood from the umbilical
and vitelline veins now enters the sinus
Development of the Portal Vein
venosus through the right hepatocardiac
channel (also called common hepatic vein). 1. The proximal parts of the two vitelline veins
This vessel later forms the cranial-most part lie o n t h e r i g h t a n d left s i d e s of t h e
of the inferior vena cava (Fig. 15.41C). developing duodenum (Fig. 15.42A).

" Liver

-Vitelline veins
Splenic vein

- Duodenum Superior
mesenteric vein

Proximal ventral anas.


Ductus venosus

Splenic and
.*- superior mesenteric
\ ^ veins

Distal ventral anas.

Fig. 15.42 Development of the portal vein. (A) Right and left vitelline veins. (B) Vitelline veins joined by three
transverse anastomoses: Cranial ventral, caudal ventral and dorsal. (C) Some of the veins dis-
appear. The portal vein is formed f r o m : 1. Part of left vitelline vein. 2. Dorsal anastomosis. 3. Part
of right vitelline vein. The cranial ventral anastomosis becomes the left branch of the portal vein.
Human Embryology

2. The veins soon become interconnected by three The veins that disappear are:
transverse anastomoses, two of which lie
(a) part of the right vitelline vein caudal to
ventral to the duodenum. The third anastomosis
the dorsal anastomosis;
lies dorsal to the duodenum, and is between
the two ventral anastomoses (Fig. 15.42B). (b) part of the left vitelline vein caudal to the
entry of the superior mesenteric and splenic
3. The superior mesenteric and splenic veins veins;
(which develop independently) join the left
(c) the caudal ventral anastomosis; and
vitelline vein, a short distance caudal to the
(d) the left vitelline vein between dorsal
dorsal anastomosis.
anastomosis and cranial ventral
4. Some parts of the vitelline veins now
anastomosis.
disappear. The portal vein and its right
and left divisions are derived from the veins The veins that persist to form the stem of the
that remain (Fig. 15.42C). portal vein are (Fig. 15.42C):

Subclavian
\
Anterior
cardinal -
Common
cardinal
x
,.' , ' s\
Posterior
cardinal -

B
Anterior, posterior Ant. cardinal v. R. & L anterior Part of anterior
& common cardinal joined by cardinal veins cardinal, common
veins formed subclavian vein joined by an cardinal & posterior
oblique anastomosis cardinal retrogress
on left side

>> A

Superior vena cava R. brachiocephalic L. brachiocephalic Internal jugulars


derived from part of from cranial part of from part of from most cranial
R ant. cardinal (1). R. ant. cardinal L. ant cardinal (1). parts of anterior
and R common and transverse cardinals
cardinal (2) s(2)

Fig. 15.43 Fate of anterior cardinal veins, and the development of major veins draining the upper part of the
body.
Cardiovascular System

(a) the left vitelline vein between the entry of 1. the right and left anterior cardinal veins that
the superior mesenteric and splenic veins d r a i n the c r a n i a l p a r t of t h e e m b r y o ,
and the dorsal anastomosis (labelled 1, in including the brain; and
the Figure); 2. the right and left posterior cardinal veins
(b) the dorsal anastomosis itself (2); and that drain the caudal part of the embryo.
(c) the right vitelline vein between the dorsal The anterior and posterior cardinal veins of
anastomosis and the cranial ventral each side join to form the corresponding common
anastomosis (3). cardinal vein (or duct ofCuvier), which open into
the c o r r e s p o n d i n g h o r n s of the sinus venosus
The cranial ventral anastomosis, and a part (Fig. 15.43A).
of t h e left v i t e l l i n e vein c r a n i a l t o t h i s
anastomosis, now form the left branch of the Fate of Anterior Cardinal and
portal vein (4), while the right vitelline vein Common Cardinal Veins
cranial to this anastomosis forms the right
The anterior cardinal veins are joined by the sub-
branch (5). The sinusoids that carry the blood
of t h e s e b r a n c h e s t o the liver s u b s t a n c e clavian veins that drain the forelimbs (Fig. 15.43B).
c o n s t i t u t e t h e venae advehentes. Those Soon thereafter, the anterior cardinal veins become
sinusoids that drain this blood to the inferior i n t e r c o n n e c t e d by a t r a n s v e r s e a n a s t o m o s i s
vena cava are called the venae revehentes and (Fig. 15.43C), proximal to their junction with the
form the tributaries of the hepatic veins. subclavian veins. The part of the left anterior
c a r d i n a l vein c a u d a l t o t h i s a n a s t o m o s i s
The left umbilical vein now ends in the left
retrogresses, and so does the left common cardinal
branch of the portal vein {Fig. 15.42D), while
(Fig. 15.43D).
the ductus venosus connects the left branch of
The Superior Vena Cava (Fig. 15.43F) is derived
the portal vein to the inferior vena cava (right
hepatocardiac channel). from:
(a) the right anterior cardinal vein, caudal to
the transverse anastomosis with the left
Somatic Veins anterior cardinal ( 1 , in the Figure); and
The earliest somatic veins are: (b) the right c o m m o n cardinal vein (2).

Anterior .
cardinal

Body wall -
J 1
Posterior
cardinal -

Fig. 15.44 Formation of the vein for the forelimb b u d . In (A) w e see veins f r o m the body w a l l draining into
the anterior and posterior cardinal veins. In (B) w e see that one of these veins lying at the level of
the limb bud enlarges to drain the l i m b .
Human Embryology

N o t e that the right horn of the sinus venosus


forms part of the right atrium, and thus the superior
vena cava comes t o open into this chamber.
The right brachiocephalic vein is derived from
the right anterior cardinal vein, between the point
of its junction with the subclavian vein and the
point of its junction with the transverse anastomosis
(Fig. 15.43F).
The left brachiocephalic vein (Fig. 15.43G} is
derived from:

(a) the part of the left anterior cardinal vein


corresponding to the right brachiocephalic
vein (1); and
(b) the transverse intercardinal anastomosis (2). Fig. 15.45 Development of the left superior
intercostal vein. The part labelled 1 is
derived from the anterior cardinal vein;
The internal jugular veins develop from the parts
and 2 from posterior cardinal vein. Note
of the anterior cardinal veins cranial to their
the intercostal veins draining into it.
junction with the subclavian veins (Fig. 15.43H).
The external jugular veins arise as secondary
channels and are not derived from the anterior (a) the left anterior cardinal vein caudal to
cardinal veins. the transverse anastomosis; and
T h e a n t e r i o r a n d p o s t e r i o r c a r d i n a l veins (b) the most cranial part of the left posterior
receive a series of intersegmental veins from the c a r d i n a l vein. T h e s e c o n d a n d t h i r d
body wall (corresponding to the intersegmental intercostal veins drain into this vein.
branches of the dorsal aortae; Fig. 15.44). The
The medial part of the coronary sinus is
subclavian veins arc formed by considerable
derived from the left horn of the sinus venosus
enlargement of one of these veins in the region of
(Fig. 15.46). The lateral part of the coronary
the upper limb bud (Fig. 15.44).
sinus is derived from the proximal part of the
left common cardinal vein. The remaining part
We have seen that the caudal part of the anterior of the left common cardinal vein persists as the
cardinal vein, and the whole of the c o m m o n oblique vein of the left atrium.
c a r d i n a l v e i n , of t h e left s i d e u n d e r g o
retrogression. The greater part of the posterior
cardinal vein of this side disappears, but a small Veins of the Abdomen
p a r t a d j o i n i n g the c o m m o n c a r d i n a l vein
persists as a small vein. We have also noted The inferior vena cava, the veins of the kidneys,
that the left horn of the sinus venosus undergoes gonads and suprarenals and the veins draining
considerable retrogression and is reduced to a the walls of the thorax and abdomen, are derived
tributary of the right horn. These retrogressing from a series of longitudinal venous channels that
veins of the left side persist into adult life as the appear in the embryo. Some of these are as follows:
left superior intercostal vein and the coronary 1. T h e posterior cardinal veins: We have
sinus which are derived as follows: already seen that at their cranial ends these
The left superior intercostal vein (Fig. 15.45) veins join the anterior cardinal veins to form
is formed by: the c o m m o n cardinal veins.
Cardiovascular System

Fig. 15.46 Derivation of the coronary sinus and related structures. 1 and 7 = right and left anterior cardinal
veins; 2 and 8 = posterior cardinal veins; 3 and 6 = c o m m o n cardinal veins; 4 and 5 = right and
left horns of sinus venosus; 9 = right vitelline vein. The fate of these structures is shown in
(B): 1 a and 3a = superior vena cava; 2a = terminal part of the azygos v e i n ; 4a = part of right
a t r i u m ; 5a and proximal half of 6a = coronary sinus; distal half of 6a = oblique vein of left atrium;
7a and 8a = left superior intercostal vein; 9a = inferior vena cava.

Near their caudal ends they receive the Cranially and caudally they communicate
veins of the lower limb bud (external iliac) with the posterior cardinal veins. The
and of the pelvis (internal iliac) subcardinals receive the veins from the
(Fig. 15.47A). The caudal ends of the two developing kidneys.
posterior cardinal veins become inter- At the level of the renal veins, the two
connected by a transverse anastomosis subcardinals become connected by a
(Fig. I5.47B). transverse intersubcardinal anastomosis
The subcardinal veins (green in Fig. 15.47) (Fig. 15.47D). The cranial part of the right
are formed in relation to the mesonephros. subcardinal vein also establishes an

Fig. 15.47 (A) Posterior cardinal veins. (B) Formation of transverse anastomosis. (C) Formation of subcardinal
veins. Note that they drain the developing kidney. (D) The t w o subcardinal veins become
interconnected.
Human Embryology

Fig. 15.48 Development of the inferior vena cava. Subcardinal veins are green, supracardinal veins are
orange, the subcardinal-hepatocardiac anastomosis is yellow, the hepatocardiac channel itself is
purple, and the supracardinal-subcardinal anastomosis is brown. The inferior vena cava receives
contributions from each of these components as indicated by the colour in (D).

anastomosis with the right hepatocardiac cardinal, and the supracardinal-subcardinal


channel (Fig. 15.48A). anastomosis).
3. T h e supracardinal veins ( a l s o c a l l e d 3. T h e right supracardinal-subcardinal
thoracolumbar veins) (orange in Fig, 15.48) anastomosis.
c o m m u n i c a t e cranially and caudally 4. The right subcardinal vein (between the
with posterior cardinal veins. They also supracardinal-subcardinal anastomosis and
communicate with the subcardinal veins the anastomosis between the subcardinal
t h r o u g h a n a s t o m o s e s which join the vein and the right hepatocardiac channel).
s u b c a r d i n a l s just below the renal veins This is the renal segment of the vena cava.
(Fig. 15.48B). 5. The subcardinal-hepatocardiac anasto-
mosis.
M a n y p a r t s of these l o n g i t u d i n a l v e n o u s 6. The right hepatocardiac channel. (This is
channels disappear (Fig. 15.48C). The veins that the hepatic segment of the vena cava.)
remain give rise to the inferior vena cava, renal
veins, veins of gonads and the suprarenal veins as The right common iliac vein is derived from
follows: the most caudal part of the right posterior
The inferior vena cava is derived from the follow- cardinal vein.
ing in caudal to cranial sequence (Fig. 15.48D): The left common iliac vein represents the
anastomosis between the two posterior cardinal
1. T h e lowest part of the right posterior veins.
cardinal vein (between its junction with the The right renal vein is a mesonephric vein
supracardinal and the anastomosis between that originally drains into the subcardinal vein
the two posterior cardinals. (Fig. 15.49A). It opens into that part of the vena
2. The lower part of the right supracardinal cava which is derived from the subcardinal vein
vein (between its junction with the posterior (Fig. 15.49B).
Cardiovascular System

Subcardinai
hepatocardiac Inferior vena
anastomosis R. suprarenal v. cava
Left subcardinal
L- suprarenal \

Vein from kidney


to subcardinal
Inter-subcardinal
anastomosis
R. gonadal v.
Supracardinal-
subcardinal
anastomosis

Fig. 15.49 Formation of renal, suprarenal and gonadal veins. The right renal vein is formed as a tributary of
the right subcardinal vein. The left renal vein is derived from (1) vein draining left kidney into left
subcardinal v e i n ; (2) part of left subcardinal vein itself; and (3) inter-subcardinal anastomosis. O n
each side, the suprarenal veins and gonadal veins represent remnants of the subcardinal veins.
From (B) it is seen w h y these veins drain, o n the right side into the inferior vena cava; and on the
left side into the left renal vein.

The left renal vein is derived from: The Azygos System of Veins
(a) the mesonephric vein that originally The veins draining the body wall at first drain
drains into the left subcardinal vein into the posterior cardinal vein (Fig. 15.50A).
(Fig. 15.49A); Their drainage is soon transferred to
(b) a small part of the left subcardinal vein; longitudinal venous channels called the veins
(c) the inter-subcardinal anastomosis. As this of the azygos line (or medial sympathetic line)
anastomosis lies in front of the aorta, the (Fig. 15.50B). Cranially these channels drain
left renal vein has a similar relationship into the posterior cardinal veins. The channels
(Fig. 15.49B). of the two sides are brought into communication
with each other by vessels that run dorsal to
The suprarenal veins are remnants of the part
the aorta (Fig. 15.50B).
of the subcardinal veins above the inter-
subcardinal anastomosis. It is clear from With the retrogression of the left common
Fig. 15.49B that the termination of the right cardinal vein, the left azygos line loses its
suprarenal vein in the inferior vena cava and communication with the posterior cardinal and
that of the left suprarenal vein in the left renal the blood of this channel now drains into the
vein, is because of their developmental origin. right azygos line through the post-aortic
The testicular or ovarian veins are remnants anastomoses. The development of the azygos
of the parts of the subcardinal veins below the system of veins can now be summarized as
inter-subcardinal anastomosis. The reason for follows (Fig. 15.50C).
the difference in the manner of termination of
the veins of the two sides, is obvious from 1. The azygos vein is formed from:
Fig. 15.49B. (a) the vein of the right azygos line; and
Human Embryology

(b) the most cranial part of the right poste- v a r i o u s veins are extremely c o m m o n .
rior cardinal vein through which it Anomalies of major veins are, however, rare.
opens into the superior vena cava (formed Some of these are as follows:
from the right c o m m o n cardinal).
1. Left superior vena cava:This is due to the
2. The vertical parts of the hemiazygos and failure of the left anterior and common
the accessory hemiazygos veins represent c a r d i n a l veins t o r e t r o g r e s s . T h e left
the left azygos line. Their horizontal parts superior vena cava opens into the right
are formed by the post-aortic anastomoses atrium through a large coronary sinus. In
between the azygos lines of the two sides. this condition, the normal (right) superior
3. The second and third left intercostal veins vena cava may be reduced in size or may
r e t a i n t h e i r c o n n e c t i o n w i t h t h e left even be absent (Fig. 15.51).
posterior cardinal vein, and are drained 2. Double inferior vena cava: Generally the
through the left superior intercostal vein. vena cava is double only below the level
4. The abdominal parts of the veins of the of the renal veins (Figs 15.52A - D).
a z y g o s line a r e r e p r e s e n t e d by t h e
(a) Both channels may be present on the
ascending lumbar veins.
right side (Fig. 15.52B). This is caused
by persistence of both the subcardinal
Anomalies of Veins
and supracardinal veins, below the
Minor anomalies in the mode of formation of level of the kidneys.

Fig. 15.50 (A) Veins from the body wall draining into anterior and posterior cardinal veins. (B) With the
formation of the azygos venous channel (Az) most of the veins of the body wall now drain into it.
(C) shows the ultimate arrangement. Note that veins from the 1st intercostal space drain into the
brachiocephalic veins directly (anterior cardinal). The veins of the left 2nd and 3rd spaces drain
into the left superior intercostal vein which is formed partly by the anterior cardinal and partly by
the posterior cardinal veins. On the right side the veins of these spaces drain into the part of the
azygos vein representing the terminal part of the right posterior cardinal.
Cardiovascular System

Right atrium
Coronary sinus

Fig. 15.51 Types of left superior vena cava. The normal pattern is shown in (A).

Fig. 15.52 Anomalies of the inferior vena cava.


(A) shows the normal pattern w h i l e (B), (C) and (D) show various types of duplication of the
infrarenal segment. In {E) the normal infrarenal segment is absent and is replaced by a vessel on
the left side. (G) shows absence of the hepatic segment of the vena cava, the b l o o d f l o w taking
place along a much enlarged vena azygos. (F) shows the corresponding normal pattern.
Human Embryology

(b) There may be an additional channel upper part of the inferior vena cava follows
on the left side (Figs 15.52C, D). the course of the azygos vein and opens
into the superior vena cava. The hepatic
Left inferior vena cava: The infrarenal part veins open into the right atrium at the
of the vena cava may be present on the usual site of the inferior vena cava
left side only (Fig. 15.52E). (Figs 15.52F, G).
Azygos continuation of inferior vena cava: Pre-ureteric vena cava: The inferior vena
The hepatic segment of the inferior vena cava normally lies posterior to the right
cava may be absent. This is due to non- ureter. Sometimes it may be anterior to
development of the anastomosis between the right ureter. The ureter then hooks
the right subcardinal vein and the right around the left side of the vena cava. This
hepatocardiac channel. In such cases the anomaly is caused when the infrarenal

Branches from aorta


to upper part of body

R pulmonary veins Left pulmonary


veins

Pulmonary trunk Aorta arising


arising from from left ventricle
right ventricle

Branches from
*— aorta to lower
part of body

v__r LOWER PART


OF BODY

Fig. 15.53 Scheme of the circulation in the human adult.


Cardiovascular System

BRAIN AND UPPER LIMBS

Pulmonary
arteries

Inferior A
vena cava m. ^^^^^r
UHl ^^^^ta^M

LIVER Ductus
arteriosus ^^^^B
j^ f c ^ Ductus
'•-•J^L venosus

PLACENTA
\ VjmDiiicai 1

Umbilic i\ arteries

LOWER PART OF BODY

Fig. 15.54 Scheme of the fetal circulation. The degree of deoxygenation is


shown by the intensity of shading. RA = right a t r i u m ; LA = left
atrium; RV - right ventricle; LV = left ventricle; F'O = foramen
ovale; P - pulmonary trunk; A = aorta.
Human Embryology

part of the vena cava develops from the (a} oxygenated blood from the right atrium,
subcardinal vein (which lies anterior to and
the ureter} instead of the supracardinal vein (b) a small a m o u n t of deoxygenated blood
(which lies posterior to the ureter}. from the lungs.

T h e blood in this c h a m b e r is, therefore,


FETAL CIRCULATION fairly rich in oxygen. This blood passes into
the left ventricle and then into the aorta.
The circulation in the fetus is essentially the same
Some of this oxygen-rich blood passes into
as in the adult except for the following (compare
the carotid and subclavian arteries to supply
Fig. 15.53 with Fig. 15.54).
the brain, the head and neck and the upper
extremities. The rest of it gets mixed up with
1. The source of oxygenated blood is not the
poorly oxygenated blood from the ductus
lung hut the placenta. arteriosus. The parts of the body that are
2. Oxygenated blood from the placenta comes supplied by branches of the aorta arising
t o the fetus t h r o u g h the umbilical vein,
distal t o its j u n c t i o n w i t h t h e d u c t u s
which joins the left branch of the portal vein.
arteriosus, therefore, receive blood with
A small portion of this blood passes through
only a moderate oxygen content.
the substance of the liver to the inferior vena
6. Much of the blood of the aorta is carried by
cava, but the greater part passes directly to
the umbilical arteries to the placenta where
the inferior vena cava through the ductus
it is again oxygenated and returned to the
v e n o s u s (Fig. 1 5 . 4 2 D ) . A s p h i n c t e r
heart.
mechanism in the ductus venosus controls
blood flow.
3. The oxygen-rieh blood reaching the right
Changes in the Circulation at Birth
atrium through the inferior vena cava is Soon after birth, several changes take place in the
directed by the valve of the inferior vena fetal blood vessels. These lead to the establishment
cava towards the foramen ovale. Here it is of the adult type of circulation. The changes are
divided into t w o portions by the lower as follows:
edge of the s e p t u m s e c u n d u m {crista
dividens): 1. The muscle in the wall of the umbilical
arteries contracts immediately after birth,
(a} Most of it passes through the foramen
and occludes their lumen. This prevents loss
ovale into the left atrium.
of fetal blood into the placenta.
(b) The rest of it gets mixed up with the
2. The lumen of the umbilical veins and the
blood returning to the right atrium
ductus venosus is also occluded, but this
through the superior vena cava, and
takes place a few minutes after birth, so
passes into the right ventricle.
that all fetal blood that is in the placenta
4. From the right ventricle, the blood (mostly has time to drain back to the fetus.
deoxygenated) enters the pulmonary trunk. 3. The ductus arteriosus is occluded, so that
Only a small portion of this blood reaches all blood from the right ventricle now goes
the lungs a n d passes t h r o u g h it to the to the lungs, where it is oxygenated.
left a t r i u m . T h e g r e a t e r p a r t is s h o r t - Initial closure of the ductus arteriosus is
circuited by the ductus arteriosus into the caused by contraction of muscle in the vessel
aorta. wall. Later intima proliferation obliterates
5. We have seen that the left atrium receives: the lumen.
Cardiovascular System

4. The pulmonary vessels increase in size and, lymph sacs. Traditionally, these sacs have been
consequently, a much larger volume of considered to be outgrowths from veins.
blood reaches the left atrium from the lungs. However, they are now regarded to be
As a result, the pressure inside the left predominantly independent formations from
atrium is greatly increased. Simultaneously, mesenchyme.
the pressure in the right atrium is diminished Six sacs can be recognized. The right and
because blood from the placenta no longer left jugular sacs lie near the junction of the
reaches it. The net result of these pressure posterior cardinal and subclavian veins (i.e. at
changes is that the pressure in the left the future junction between the internal jugular
atrium now exceeds that in the right atrium and subclavian veins). The right and left
causing the valve of the foramen ovale to posterior (or iliac) sacs lie around the
corresponding common iliac vein. The
retroperitoneal sac (unpaired) lies in relation
The vessels that are occluded soon after birth to the root of the mesentery. The sixth sac (again
are, in due course, replaced by fibrous tissue, and unpaired) is the cisterna chyli. It lies in the
form the following ligaments: midline, some distance caudal to the
retroperitoneal sac.
Vessel Remnant Lymphatic vessels are formed either by
extension from the sacs or may form de novo,
(a) Umbilical arteries Medial umbilical
ligaments and extend into various tissues. Ultimately all
the sacs except the cisterna chyli are invaded
(b) Left umbilical vein Ligamentum teres
by connective tissue and lymphocytes, and are
of the liver
converted into groups of lymph nodes.
(c) Ductus venosus Ligamentum The thoracic duct is derived from right and
venosum left channels that connect the cisterna chyli to
(d) Ductus arteriosus Ligamentum the corresponding jugular sac. The two channels
arteriosum anastomose across the midline. The thoracic
duct is formed from the caudal part of the right
LYMPHATIC SYSTEM channel, the anastomosis between the right and
left channels, and the cranial part of the left
The first signs of the lymphatic system are seen channel. The cranial part of the right channel
in the form of a number of endothelium lined becomes the right lymphatic duct.
Human Embryology

TIMETABLE OF SOME EVENTS DESCRIBED IN THIS CHAPTER


Age Developmental Events
3rd week Blood and vessel forming cells (angioblastic islands) appear.
The cardiogenic area, heart tubes and pericardium have formed.
4th week Heart and pericardium lie ventral
to foregut.
Subdivisions of heart tube are visible.
Heart begins to beat (becomes functional).
Formation of septa in the heart begins.
Aortic arches start appearing in cranial to caudal sequence.
Most of the first aortic arch disappears at the end of 4th week.
Veins start forming.
5th week The spiral septum is formed.
Formation of aortic arches is complete.
Lymphatic sacs form.
The cardinal, umbilical and
vitelline veins are formed.
Conduction system of heart forms.
6th week Coronary circulation is established.
Atrio-ventricular valves and papillary muscles are forming.
7th week Heart septa are completely formed.

The heart is most susceptible to teratogens between three and six weeks. It can be affected up t
the eighth week.
I
Chapter

16 Urogenital System

HIGHLIGHTS • T h e uterine tubes are derived from


paramesonephric ducts (mesoderm).
• The urogenital system is derived from the inter-
mediate mesoderm, and the primitive urogeni- • The uterus is formed from the utero-vaginal
tal sinus (UGS) which is a part of the cloaca. canal (fused right and left paramesonephric
ducts).
• The primitive UGS divides into the vesico-
urethral canal and the definitive UGS (Fig. 16.3). • External genitalia are formed from swellings
that appear around the urogenital membrane.
• The vesico-urethral canal divides into the
urinary bladder and the primitive urethra. • Gonads (testis and ovary) are derived from
coelomic epithelium covering the nephrogenic
• The definitive UGS has a pelvic part and a c o r d . Ova a n d s p e r m a t o z o a arise from
phallic part. primordial germ cells that arise in the region
• The kidneys develop from two sources. The of the yolk sac. Each testis is formed in the
excretory tubules (nephrons) are derived from l u m b a r region, a n d later descends to the
the metanephros (= lowest part of nephrogenic scrotum.
c o r d w h i c h is derived from i n t e r m e d i a t e • The duct system of the testis is derived from
mesoderm). The collecting part is formed by mesonephric tubules and from the mesonephric
ramification of the ureteric hud (which arises duct.
from the mesonephric duct).
• The ureter arises from the ureteric bud.
INTRODUCTION
• The urinary bladder is derived from the cranial
part of the vesico-urethral canal (endoderm). Two embryonic structures that play an important
The epithelium of the trigone is derived from role in the development of the urogenital system
absorbed mesonephric ducts. are the intermediate mesoderm and the cloaca.
• The female urethra is derived from the primitive These are briefly considered below.
urethra and the pelvic part of the UGS.
Intermediate Mesoderm
• In the male, the prostatic urethra corresponds
to the female urethra. The membranous urethra We have seen (Chapter 5) that the intra-embryonic
is derived from the pelvic part of UGS and the mesoderm is subdivided into:
penile urethra from the phallic part of UGS.
The terminal part is ectodermal. (a) the p a r a x i a l m e s o d e r m which becomes
• The prostate is formed by buds arising from segmented to form the somites,
the caudal part of the vesico-urethral canal and (b) the lateral plate mesoderm in which the
the pelvic part of UGS. intra-embryonic coelom appears, and
Human Embryology

Intra-embryonic (c) the intermediate mesoderm lying between


Neural groove
coelom the two (Fig. 16.1).

After the folding of the embryonic disc and the


formation of the peritoneal cavity, the intermediate
mesoderm forms a bulging on the posterior
abdominal wall - lateral to the attachment of the
dorsal mesentery of the gut. This bulging is now
Somite mesoderm called the nephrogenic cord (Fig. 16.2A). Its surface
is covered by the epithelium lining the peritoneal
Fig. 16.1 Location of intermediate mesoderm.
cavity (coelomic epithelium). It extends from the
cervical region to the sacral region of the embryo.
At varying stages of development, a number of

L Nephrogenic Paramesonephric duct


cord
Mesonephnc duct
Mesonephros
Gonad

Fig. 16.2 Nephrogenic cord (A) and structures that develop in it (B).

CLOACA

RECTUM

PRIMITIVE UROGENITAL SINUS

1
i \ sinus
Vesicourethral canal Definitive urogenita

I 1 1 i 1 1
URINARY PRIMITIVE PELVIC PHALLIC
BLADDER URETHRA PART PART
Fig. 16.3 Subdivisions of the cloaca. Also see Fig. 16.4.
Urogenital System

important structures are formed in relation to the the primitive urogenital sinus and rectum have been
nephrogenic cord on each side. considered in Chapter 13 (see Figs 13.3 and 13.4).
These are as follows (Fig. 16.2B): In further development, the primitive urogenital
sinus is subdivided into a cranial part, called the
(a) Excretory tubules associated with the vesico-urethral canal, and a caudal part, called
development of the kidney. the definitive urogenital sinus. The openings of
(h) The nephric duct which is formed in relation the mesonephric ducts (see below) lie at the junction
to the developing excretory tubules of these two subdivisions (Fig. 16.4A}. Still later,
mentioned in (a). At later stages, this the definitive urogenital sinus shows a division
becomes the mesonephric duct. into a crania! pelvic part and a caudal phallic
(c) The paramesonephric duct, which is formed part (Fig. 16.4B ).
lateral to the nephric duct. The urogenital system is derived from the
(d) The gonad (testis or ovary), which develops various structures that develop in the intermediate
from the coelomic epithelium lining the mesoderm and from the various subdivisions of
medial side of the nephrogenic cord. the cloaca, as described now.

Cloaca
DEVELOPMENT O F KIDNEYS
The formation of the cloaca and its subdivision into
The definitive human kidney arises from two
distinct sources. The excretory tubules (or
nephrons) are derived from the lowest part of the
nephrogenic cord. This part is the metanephros,
the cells of which form the metanephric blastema.
•- Vesico-urethral canal

----- Mesonephric duct

Urogenital sinus

_ Urinary bladder
Metanephros
--Primitive urethra

Pelvic pari of urogenital sinus

Phallic part of urogenital sinus

Fig. 16.5 Pronephros, mesonephros and


metanephros. Note that the mesonephric
Fig. 16.4 Subdivisions of the primitive urogenital duct opens into the cloaca; and gives off
sinus. Also see Fig. 16.3. the ureteric bud.
Human Embryology

Degenerating
Pronephricduct

duct
Fig. 16.6 Some details of developing
Glomerulus ^ ^ x ^ T ^ pronephros, mesonephros
and metanephros. The
Excretory pronephros and pronephric
mesonephric
tubule duct degenerate soon after
MESONEPHROS formation. The proximal
(Green)
part of the mesonephros
shows segmentation (in
cranio-caudal sequence).
The segments contain
functional excretory tubules
that drain into the
mesonephric duct. Most of
METANEPHROS
these tubules disappear by
the time the metanephros
Ureteric bud
forms the definitive kidney.

Metanephric tissue
/
Ureteric
bud'
#

Major calyx

Minor calyx
Ureteric bud dividing Collecting tubules
into various generations
of branches D

Fig. 16.7 Formation of collecting system of the kidney, from ramifications of the ureteric bud.
Urogenital System

The collecting part of the kidney is derived from through three stages of evolution. The most
a diverticulum called the ureteric bud, which arises primitive of these is called the pronephros. It is
from the lower part of the mesonephric duct the functioning kidney in some cyclostomes and
(Fig. 16.5). fishes. This has been succeeded in higher
vertebrates by the mesonephros, which is the
Some of the features of the development of functioning kidney of most anamniotes. The
kidney in the human embryo can be appreciated kidney of amniotes (including man) is called
only if the evolutionary history of the organ is the metanepbros.
kept in mind. The vertebrate kidney has passed During the development of the human

Metanephric vesicle

Vesicle becomes
pear shaped

Proximal convoluted
tubule

Henle's loop

\J
Fig. 16.8 Scheme to show stages in the development of the nephron.
Human Embryology

embryo, the evolutionary history of the kidney


repeats itself being a classic example of the
saying that ontogeny repeats phytogeny. The
pronephros is formed in relation to the cervical
region of the nephrogenic cord. This is followed
by appearance of the mesonephros in the
thoracolumbar region, and finally by formation
of the metanephros in the sacral region (Fig.
16.5). The human pronephros is non-functional,
and disappears soon after its formation. A
nephric duct formed in relation to the
pronephros and ending in the cloaca, however,
persists. The mesonephros consists of a series
of excretory tubules that develop in the
Fig. 16.9 Ascent of the kidney.

Fig. 16.10 Anomalies of the kidney. (A) Congenital polycystic kidney. (B) Aberrant renal arteries. (C) Lobulated
kidney. (D), (E), (F) Transposition of kidney. (G), (HI Horseshoe kidney, and (I) Pancake kidney.
Urogenital System

Mesonephric duct
Mesonephric duct
Part of bladder wall derived
from mesonephric ducts

n
Primitive urethra

Fig. 16.11 (A) Mesonephric duct opens into primitive urogenital sinus. (B) As the sinus grows the proximal
parts of mesonephric clue Is arc absorbed so [hat the mesonephric due is and ureters n o w open
separately- (C) The openings are at first close together. (D) Further absorption of ureters causes
their opening to shift upwards and laterally. The shaded area is derived from absorbed parts of
ureters and mesonephric ducts and is of mesodermal origin. II forms the trigone of the bladder
and the posterior w a l l of part of the urethra.

thoracolumbar region. These tubules drain into the minor calyces a n d c o l l e c t i n g t u b u l e s


the nephric duct which may now be called the (Fig. 16.7).
mesonephric duct. Most of the mesonephric Collecting tubules formed vary in number
t u b u l e s d i s a p p e a r , b u t s o m e of t h e m are from one to three million. The growing tip of
modified and take part in forming the ducr each of the numerous branches of the ureteric
system of the testis. bud is dilated to form an ampulla. The cells of
As the ureteric bud grows cranially towards the metanephric blastema in contact with an
the m e t a n e p h r i c blastema, its growing end ampulla u n d e r g o differentiation to form a
becomes dilated to form an ampulla. The nephron. This differentiation is induced by the
ampulla divides repeatedly. The first three to a m p u l l a . L o o s e l y a r r a n g e d cells of t h e
five generations of branches fuse to form the metanephric blastima form solid clumps in
pelvis of the kidney. The next divisions become relation to the ampulae. Each solid clump is
the major calyces while further divisions form c o n v e r t e d into a vesicle. T h e vesicle soon
Human Embryology

becomes pear shaped and opens into the


ampulla. The vesicle now becomes an 'S'
shaped tube. Its distal end comes to be
invaginated by a tuft of capillaries which form
a glomerulus. The various parts of the nephron
are derived from this ( S' shaped tube as shown
in Fig. 16.8.

Ascent of the Kidney


We have seen that the definitive human kidney
is derived from the metanephros, which lies in
the sacral region. In subsequent development
of the embryo, differential growth of the
abdominal wall causes the kidney to ascend to
the lumbar region (Fig. 16.9). The metanephros, Fig. 16.12 Anomalies of ureters. Also see
Fig. 16.10.
at first, receives its blood supply from the lateral
sacral arteries, but with its ascent, higher 3. Anomalies of Shape
branches of the aorta take over the supply. The
definitive renal artery represents the lateral (a) Horseshoe kidney: The lower poles of
splanchnic branch of the aorta at the level of the two kidneys (or sometimes the
the second lumbar segment. upper poles) may be fused. The
connecting isthmus may He either in
During the ascent, the kidneys pass through
front of, or behind, the aorta and
the fork-like interval between the right and left
inferior vena cava (Figs I6.10G, H).
umbilical arteries. If the arteries come in the
A horseshoe kidney does not ascend
way of ascent, the kidney may remain in the
higher than the level of the inferior
sacral region (see 'Anomalies of Kidneys').
mesenteric artery as the latter prevents
its higher ascent.
Rotation of the Kidney
(b) Pancake kidney: The two kidneys may
The hilum of the kidney, at first, faces form one mass, lying in the midline
anteriorly. The organ gradually rotates so that or on one side (Fig. 16.101).
the hilum comes to face medially. (c) The two kidneys may lie on one side,
Anomalies of Kidneys one above the other, the adjacent poles
being fused.
1. One or both kidneys may be absent (d) Lobulated kidney: The fetal kidney
[agenesis). The kidney may be under- is normally lobulated. This lobulation
developed {hypoplasia} or overdeveloped may persist (Fig. 16.IOC).
{hyperplasia). Adrenal tissue may be
present within the substance of the kidney. 4. Anomalies of Position
Distension of the pelvis with urine (a) The kidneys may fail to ascend. They
{hydronephrosis) may occur as a result of then lie in the sacral region.
obstruction in the urinary passages. (b) The ascent of the kidneys may be
2. Duplication: There may be an extra kidney incomplete as a result of which they
on one side. It may be separate or may be may lie opposite the lower lumbar
fused to the normal kidney (Fig. 16.12). vertebrae.
Urogenital System

(c) The kidneys may ascend too far, and (b) Incomplete rotation: T h e hilum is
m a y even be p r e s e n t w i t h i n t h e directed anteromedially.
thoracic cavity. (c) Reverse rotation: The hilum is directed
(d) Both kidneys may lie on one side of anterolaterally.
the midline. They may lie one above
6. Congenital Polycystic Kidney: Failure of
the other or side by side (Figs 16.1 OD,
the excretory tubules of the metanephros
E). The ureter of the displaced kidney
to establish contact with the collecting
crosses to the opposite side across the
tubules, leads to the formation of cysts.
midline.
Isolated cysts are c o m m o n l y seen, but
(e) Both kidneys may be displaced to the sometimes the whole kidney is a mass of
opposite side. The two ureters then such cysts (Fig. 16.10A). The cysts press
c r o s s e a c h o t h e r in t h e m i d l i n e upon normal renal tissue and destroy it.
(Fig. 16.10F).
An a l t e r n a t i v e r e c e n t view on the
5. Abnormal Rotation formation of cysts in the kidney is that
they are derived from a b n o r m a l l y
(a) Non-rotation: The hilun is directed
developed collecting tubules.
forwards.
7. Aberrant Renal Arteries: The kidney may
receive its b l o o d s u p p l y p a r t i a l l y or
e n t i r e l y , f r o m a r t e r i e s a r i s i n g at an
abnormal level (Fig. 16.10B). In the case
of non-ascent, or of incomplete ascent, the
aberrant arteries may constitute the only
supply to the organ. An aberrant artery
may be the only source of arterial blood
t o a segment of the kidney. It may press
upon the ureter and cause obstruction,
leading to hydronephrosis.
8. Multiple Anomalies: Two or more of the
above anomalies may coexist. Anomalies
of position are frequently associated with
those of rotation.

ABSORPTION OF LOWER PARTS


OF MESONEPHRIC DUCTS
INTO CLOACA
It has been noted above, that the lower ends of the
mesonephric ducts open into t h a t part of the
cloaca that forms the urogenital sinus. It has also
been seen, that the ureteric buds arise from the
mesonephric ducts, a little cranial to the cloaca
(Fig. 16.11A). The parts of the mesonephric ducts,
caudal to the origin of the ureteric buds, are
Fig. 16.13 Abnormal sites at w h i c h the ureter may absorbed into the vesico-urethral canal, with the
open. result that the mesonephric ducts and the ureteric
Human Embryology

buds now have separate openings into the cloaca DEVELOPMENT OF THE
(Fig. 16.1 IB). These openings are at first close URINARY BLADDER
together (Fig. 16.11C). However, the openings of
the ureteric buds move cranially and laterally due The epithelium of the urinary bladder develops
to continued absorption of the buds. The triangular from the cranial part of the vesico-urethral canal
area (on the dorsal wall of the vesico-urethral (endoderm). The epithelium of the trigone of the
canal) between the openings of the ureteric buds bladder is derived from the absorbed mesonephric
and those of the mesonephric ducts, is derived from ducts (mesoderm). (However, it is later overgrown
the absorbed ducts and is, therefore, of mesodermal by t h e s u r r o u n d i n g e n d o d e r m a l cells.) T h e
origin (Fig. 16.11D). muscular and serous walls of the organ are derived
from splanchnopleuric mesoderm.
DEVELOPMENT OF THE URETER The developing bladder is continuous cranially
with the allantois. It is uncertain whether the
The ureter is derived from the part of the ureteric allantois contributes to the formation of the bladder.
bud that lies between the pelvis of the kidney, and The allantois atrophies and is seen in postnatal
the vesico-urethral canal. life as a fibrous band, the urachus, extending from
the apex of the bladder to the umbilicus.
Anomalies of the Ureter Anomalies of the Urinary Bladder
1. The ureter may be partially or completely 1. T h e urinary bladder may be absent, or
duplicated (Fig. 16.12). This condition may be duplicated.
may, or m a y n o t , be a s s o c i a t e d with
2. The sphincter vesicae may be absent.
duplication of the kidney. Very rarely, there
3. The lumen of the urinary bladder may be
may be more than t w o ureters on one, or
divided into compartments by septa.
both sides. Of the two ureters one may
4. The bladder may be divided into upper
open into the urinary bladder while the
and lower c o m p a r t m e n t s (hourglass
other may open at an abnormal site (see
bladder) because of a constriction in the
below).
middle of the organ (Fig. 16.14A).
2. Instead of opening into the urinary bladder, 5. The bladder may communicate with the
the ureter may end in the prostatic urethra, rectum (Figs 13.25A, H ) .
ductus deferens, seminal vesicles or rectum,
in the male (Fig. 16.13B); a n d in the
urethra, vagina, vestibule or rectum in the
female (Fig. 16.13A).
3. The upper end of the ureter may be blind,
i.e. it is not connected to the kidney.
4. The ureter may be dilated {hydroureter)
because of obstruction to urine flow.
5. The ureter may have valves or diverticula.
6. T h e right ureter may pass behind the
inferior vena cava. It then hooks around
the left side of the vena cava; this may
result in kinking and obstruction of the
Fig. 16.14 Anomalies of the bladder. {A) Hourglass
ureter. The real defect is in the develop-
bladder. (B) Ectopia vesicae. The ureteric
ment of the vena cava as described in
Openings and the trigone are seen o n the
Chapter 15.
surface of the body.
Urogenital System

"emale urethra
Prostatic urethra

Membranous urethra

oo
Fig. 16.15 Development of urethra. (A) Primitive UGS showing opening of
mesonephric ducts. (B) Primitive UGS divided into vesico-
urethral canal and definitive UGS. Mesonephric ducts and
ureters open separately at the junction of the t w o parts.
( O Vesico-urethral canal subdivided into urinary bladder and
primitive urethra. The definitive UGS divides into pelvic and
phallic parts. (D) The female urethra is formed from the primitive
urethra and from part of the pelvic portion of UGS. The rest of
the pelvic part of UGS forms the vestibule. (E] In the male the
prostatic urethra is formed in the same w a y as the female
urethra. The membranous urethra is derived from the pelvic part
of UGS. The penile urethra is derived from the phallic part of
UGS. Red circles = openings of mesonephric ducts and ureters.
Blue = part derived from mesoderm. Green = ectoderm.
Human Embryology

6. Ectopia vesicae: The lower part of the from the mesonephric ducts and is, therefore,
anterior abdominal wall, as well as the mesodermal in origin. The female urethra may
v e n t r a l w a l l of t h e b l a d d e r , m a y be receive a slight contribution from the pelvic part
missing. As a result, the cavity of the of the urogenital sinus (Fig. 16.15).
bladder may be exposed on the surface of
the b o d y (Fig. 16.14B). This defect is
DEVELOPMENT OF THE
usually associated with epispadias. Ectopia
MALE URETHRA
vesicae is caused by failure of mesoderm
to migrate into the lower abdominal wall (a) The part of the male urethra extending from
(between umbilicus and genital tubercle). the urinary bladder up to the openings of
F a i l u r e of m i g r a t i o n m a y be d u e t o the ejaculatory ducts (original openings of
excessive d e v e l o p m e n t of the cloacal mesonephric ducts), is derived from the
membrane. The ectoderm of the anterior caudal part of the vesico-urethral canal
abdominal wall and the endoderm of the (endoderm). The posterior wall of this part
ventral wall of the urinary bladder remain is derived from the absorbed mesonephric
unsupported and thin. Their rupture leads d u c t s ( m e s o d e r m ) . (It m a y l a t e r b e
to the exposure of the cavity of the urinary overgrown by endoderm.)
bladder. (b) The rest of the prostatic urethra, and the
7. Congenital diverticula may be present. membranous urethra, are derived from the
These are found at the junction of the pelvic p a r t of the definitive u r o g e n i t a l
trigone with the rest of the bladder. sinus.
(c) The penile part of the urethra (except the
DEVELOPMENT OF THE terminal part) is derived from the epithelium
FEMALE URETHRA of the phallic part of the definitive urogeni-
tal sinus (see 'Development of Penis').
The female urethra is derived from the caudal part (d) The terminal part of the penile urethra, that
of the vesico-urethral canal (endoderm). We have lies in the glans, is derived from ectoderm
seen that the posterior wall of this canal is derived (Fig. 16.15).

Urinary bladder Median lobe


Prostatic urethra Prostatic utricle

Capsule
Stroma (muscle,
connective tissue)
Prostatic utricle
Glands opening onto
posterior wall of urethra
above ejaculatory ducts

Fig. 16.16 Mesodermal and endodermal derivatives of the prostate. The glands of the median lobe, w h i c h
open onto the posterior wall of the prostatic urethra (above the opening of the ejaculatory ducts)
are mesodermal. Figure (A) shows a transverse section above the level of the opening of
ejaculatory ducts. Figure (B) is a sagittal section.
Urogenital System

From the above it will be clear that the female


Mesenchyme of nephrogenic cord
urethra corresponds to the prostatic part of the
Coelomic
male urethra. epithelium Paramesonephhc
duct

Anomalies of the Urethra


1. There may be an obstruction to the urethra
at its junction with the bladder.
2. The urethra may show diverticula.
3. It may be duplicated in whole or in part.
4. The urethra may have abnormal commu-
nications with the rectum (Figs 13.23B, C),
vagina (Fig. 16.22) or ureter (Fig. 16.13).
5. Hypospadias and epispadias.

DEVELOPMENT OF THE PROSTATE


Fig. 16.17 Formation of paramesonephric ducts by
This gland develops from a large number oi buds invagination of c o e l o m i c epithelium.
that arise from the epithelium of the prostatic
urethra, i.e. from the caudal part of the vesico-
urethral canal and from the pelvic part of the The outer zone differentiates earlier than the
definitive urogenital sinus. These buds form the inner zone. In later life, the outer zone is
secretory epithelium of the gland. The buds that frequently the site of carcinomatous change,
arise from the mesodermal part of the prostatic while the inner zone is affecred in senile
urethra (i.e. posterior wall, above the openings of hypertrophy of the organ.
the ejaculatory ducts) form the inner glandular The muscle and connective tissue of the gland
zone of the prostate. Buds arising from the rest of are derived from the surrounding mesenchyme
the prostatic urethra (endoderm) form the outer which also forms the capsule of the gland.
glandular zone.

Gonad

uI- "1/

w
% • - « - Mesonephnc -*-mm
1 Paramesonephric mm

w
^M duct mm
\
Utero-
vaginal
canal

A B
Fig. 16.18 Formation of utero-vaginal canal by fusion of the caudal parts of paramesonephric ducts.
Human Embryology

hc:\ rgN
>// Prostatic
! utricle
r l 2
X Appendix of
3 V
testis
fpJLry.
A
0
/*C
B
Testis

Fig. 16.19 Fate of paramesonephric ducts. (A) In the female they form the uterine tubes, the uterus and part
of the vagina. (B) In the male most of the duct disappears. Remnants are seen as the appendix of
the testis and the prostatic utricle.

The secretory elements of the prostate are from the urogenital sinus form the paraurethral
rudimentary at birth. They undergo consi- glands of Skene.
derable development at puberty. The organ
undergoes progressive atrophy in old age, but PARAMESONEPHRIC DUCTS
in some men it undergoes benign hypertrophy.
The prostate may, rarely, be absent. We have seen that these ducts are present in the
intermediate mesoderm. They are formed by
Female Homologues of Prostate invagination of coelomic epithelium (Fig. 16.17).
They lie lateral to the mesonephric ducts in the
Endodermal buds, similar to those that form cranial part of the nephrogenic cord (Fig. 16.18A).
the prostate in the male, are also seen in the When traced caudally they cross to the medial
female. The buds that arise from the caudal side of the mesonephric ducts. Here the ducts of
part of the vesico-urethral canal give rise to the two sides meet and fuse in the middle line to
the urethral glands^ whereas the buds arising form the utero-vaginal canal (or uterine canal)

Fig. 16.20 Anomalies of the uterus. (A) Duplication of uterus and vagina. (B) Bicornuate uterus. (C) Septum in
uterus. (D) Unicornuate uterus.
Urogenital System

(Fig. 16.18B). The caudal end of this canal comes Anomalies of the Uterus
in contact with the dorsal wall of, the definitive
urogenital sinus. We have already seen that, in 1. The uterus may be completely, or partially,
the female, this part of the sinus give rise to the duplicated (Figs 16.20A, B). Complete
vestibule. In the female, the paramesonephric ducts duplication is referred to as uterus
give origin to the uterine tubes, the uterus and didelphys.
part of the vagina (Fig. 16.19A). 2. The lumen may be partially, or completely,
subdivided by a septum (Fig. 16.20C).
3. The entire uterus may be absent.
DEVELOPMENT OF UTERUS AND
4. One half of the uterus may be absent
UTERINE TUBES {unicornuate uterus) (Fig. 16.20D).
The epithelium of the uterus develops from the 5. The uterus may remain rudimentary.
fused paramesonephric ducts (utero-vaginal canal: 6. There may be atresia of the lumen either
1 in Fig. 16.19A). The myometrium is derived in the body or in the cervix.
from surrounding mesoderm (3). As the thickness
Anomalies of the Uterine Tubes
of the myometrium increases, the unfused
horizontal parts of the two paramesonephric ducts 1. The uterine tubes may be absent, on one
come to be partially embedded within its or both sides.
substance, and help to form the fundus of the 2. The tubes may be partially, or completely,
uterus (2). The cervix can soon be recognized as a duplicated on one or both sides.
separate region. In the fetus the cervical part is 3. There may be atresia of the tubes.
larger than the body of the uterus.
The uterine tubes develop from the unfused parts DEVELOPMENT OF VAGINA
of the paramesonephric ducts. The original points
of invagination of the ducts into the coelomic We have noted that the lower end of the utero-
epithelium remain as the abdominal openings of vaginal canal comes in close contact with the
the tubes. Fimbria are formed in this situation. dorsal wall of part of the urogenital sinus

Sinovaginal
Utero-vaginal canal bu|b
/ V

Fig. 16.21 (A) Utero-vaginal canal (mesoderm) in contact with lining of UGS (endoderm). (B) Sinovaginal
bulbs are formed by proliferation of endodermal lining. (C) Solid vaginal plate derived partly from
mesoderm of utero-vaginal canal and partly from endoderm of sinovaginal bulbs. (D) Vagina
formed by canalization of vaginal plate.
Human Embryology

(Fig. 16.21A). However, the utero-vaginal canal


and the urogenital sinus are soon separated from
each other by the formation of a solid plate of
cells called the vaginal plate. The vagina is
formed by the development of a lumen within the
vaginal plate (Fig. 16.21D). -Sacrum

The vaginal plate is formed as follows:


Endodermal cells of the urogenital sinus
proliferate to form two swellings called the
sinovaginal bulbs (Fig. 16.21B). These bulbs
soon fuse to form one mass. Most of the vaginal
plate is formed from these sinovaginal bulbs
(Fig. 16.21C). The part of the vaginal plate Vagina
near the future cervix is derived from
mesodermal cells of the utero-vaginal canal. Fig. 16.22 Vaginal fistulae are abnormal c o m m u n i -
The hymen is situated at the junction of the cations between vagina and surrounding
cavities. The fistulae are shown in solid
lower end of the vaginal plate with the
black. They may connect the vagina to
urogenital sinus. Both surfaces of the hymen
the rectum (rectovaginal fistula]; to the
are lined by endoderm. urinary bladder (vesico-vaginal fistula) or
to the urethra (uretero-vaginal fistula).

Genital
tubercle
Urogenital
membrane

Cloacal _
membrane • _\ n / "O Genital
swelling
r\
• c
Urogenital
membrane

Genital tubercle ^ ^ Labium ^ t t B k .


ma us
Primitive ^ f f V ' \ J f l
urethral told ^ m A

V
swelling [ I V I / \ -_/'
Openinq into S* —
Urogenital sinus Vestibule —Anus

Fig. 16.23 (A) Cloacal membrane. (B) Cloacal membrane divides into urogenital membrane and anal
membrane. (C) Right and left genital swellings, and a median genital tubercle appear.
(D) Urogenital membrane breaks d o w n . Its edges form the primitive urethral folds.
(E) Genital tubercle becomes the clitoris. The genital swellings become the labia majora,
and the primitive urethral folds become the labia minora.
Urogenital System

Phallic part of
urogenital sinus

Primitive urethral fold Labium minus


Labium majus
Urogenital membrane

Fig. 16.24 Development of female external genitalia. Ectoderm is shown in green


and endoderm in orange. Compare w i t h Fig. 16.23.

Anomalies of Vagina DEVELOPMENT OF


EXTERNAL GENITALIA
1. The vagina may be duplicated. This
condition is usually associated with Introduction
duplication of the uterus (Fig. 16.20A). We have seen that, with the formation of the
2. The lumen may be subdivided longi- urorectal septum, the cloacal membrane comes
tudinally, or transversely, by a septum. to be subdivided into a ventral, urogenital
3. The vagina may be absent. This condition membrane and a caudal anal membrane
may or may not be associated with absence (Figs 16.23A, B). The urogenital membrane
of the uterus. becomes elongated in a cranio-caudal direc-
4. The hymen may be imperforate. tion. The mesoderm on either side of it is soon
5. The vagina may have abnormal commu- heaped up to form two longitudinal elevations
nications with the rectum {rectovaginal
called the primitive urethral folds (Figs 16.23D,
fistula) or with the urinary bladder (vesico-
16.24A). In addition to these folds, three other eleva-
vaginal fistula) (Fig. 16.22).
tions of mesoderm are soon apparent. These are:
(a) the genital tubercle which is situated in
Paramesonephric Ducts in Male
the midline between the urogenital
The paramesonephric ducts remain rudimentary
in the male. The greater part of each duct
eventually disappears (Fig. 16.19B). The cranial
end of each duct persists as a small rounded
body attached to the testis {appendix of testis)
which may occasionally give rise to cysts. It
has generally been considered that the prostatic
utricle represents the utero-vaginal canal and
is, therefore, a homologue of the uterus.
However, it is now believed to correspond
mainly to the vagina (and possibly part of the
uterus).
Fig. 16.25 Formation of prepuce of penis.
Human Embryology

Fig. 16.26 A t o D Stages in the development of


male genitalia and of penile
urethra. In each set (A to H),
the central figure (1) shows the
genital region from the ventral
aspect; (2) and (3) are transverse
sections at the levels indicated;
and (4) is a median section
through the region. In sections
ectoderm is depicted in black
line, endoderm in red and
mesoderm in green.

Note the following. The phallus is


formed by enlargement of the
genital tubercle.
Caudal to the phallus there is a
median, longitudinal depression,
the primitive urethral groove
(PUG! bounded by primitive
urethral folds (PUF). Lateral to
these folds w e see the genital
swellings (GS). In the depth of the
primitive urethral groove there is
llit- urogenital membrane w h i c h
separates the groove from the
urogenital sinus.

B. The phallus has enlarged. The


primitive urethral groove (PUG) is
beginning to extend onto it. A
solid mass of endodermal cells

J~C derived from the urogenital sinus,


extends into the phallus. This
mass is the urethral plate (UP).

C. The primitive urethral groove is


n o w fully formed. The urethral
plate has enlarged and extends
deeper into the phallus.

D. The urogenital membrane has


broken d o w n so that endoderm
of the urogenital sinus (UGS) can
now be seen from outside.
Urogenital System

Fig. 16.26 (Continued) E to H

(First see A t o D o n previous page.)

E. Ectoderm overlying the urethral


plate has disappeared. As a result
endoderm of the plate is seen o n
the surface.

Cells in the centre of the urethral


plate now break d o w n and
convert the plate into groove that
is seen o n the surface. This is the
definitive urethral groove, and
the folds forming its edges are the
definitive urethral folds.

Solid ectodermal
Q downgrowth

The definitive urethral folds grow


towards each other and fuse to
form a median raphe. In this way
the definitive urethral groove is
converted into a tube, w h i c h is
the urethra. This process of
fusion starts caudally and
progresses cranially.

Raphe

In this figure and in (C), note that


the urethra formed as described
above does not extend into the
glans. The part of the urethra
lying in the glans is derived from
ectoderm, w h i c h first forms a
solid cord that is later canalized.

Raphe Scrotum
Human Embryology

m e m b r a n e a n d the l o w e r p a r t of t h e covering the distal p a r t of the phallus


anterior abdominal wall; and (Fig. 16.25).
(b) t h e r i g h t a n d left genital swellings (b) We h a v e seen t h a t t h e u r o g e n i t a l
(Fig. 16.23C). membrane lies in a linear groove, flanked
on either side by the primitive urethral
Development of Female External folds (Figs 16.26A to C). As the phallus
Genitalia (Figs 16.23, 16.24) grows, this groove elongates and extends
(a) The genital tubercle becomes cylindrical onto its undersurface (Fig. 16.26B). This
and forms the clitoris. groove is lined by ectoderm and is called
(b) The genital swellings enlarge to form the the primitive urethral groove.
labia majora. Their posterior ends fuse (c) From Figs 16.26C and 16.26D it will be
across the midline to form the posterior clear that the phallus is closely related to
commissure. the endodermal lining of the phallic part
(c) The urogenital membrane breaks down, of the urogenital sinus. The endodermal
so that continuity is established between cells of this lining proliferate and grow
the urogenital sinus (which forms the into the phallus, in the form of a solid plate
vestibule) and the exterior. The primitive of c e l l s c a l l e d t h e urethral plate
urethral folds now form the labia minora. (Fig. 16.26C). T h e cells of the urethral
It will be obvious that they are lined on plate are in contact with the ectodermal
the outside by ectoderm and on the inside cells lining the primitive urethral groove.
by endoderm (Fig. 16.24). (d) The urogenital m e m b r a n e soon breaks
down, so that the urogenital sinus (phallic
Development of Male External Genitalia
part) opens to the outside, in the caudal
(a) The genital tubercle becomes cylindrical p a r t of the p r i m i t i v e u r e t h r a l g r o o v e
and is now called the phallus. It undergoes (Fig. 16.26D). At the same time, the cells
great enlargement to form the penis. As forming the core of the urethral plate
the phallus g r o w s , the glans becomes degenerate, along with the ectodermal
distinguishable by the appearance of a cells lining the primitive urethral groove.
coronary sulcus. Still later, the prepuce is In this way, a deeper groove (called the
formed by reduplication of the ectoderm definitive urethral groove) lined by

Open groove
under penis Various levels
at which urethra
Half of scrotum may open

Urethral opening

Fig. 16.27 (A) Cleft scrotum. (B) Hypospadias. The urethra opens onto the ventral aspect of the penis.
Urogenital System

endodermal cells, is now formed on the prepuce, may be missing. The opening of
undersurface of the phallus (Fig. 16.26F). the prepuce may be too narrow to allow
At the base of the phallus this groove is retraction (phimosis).
continuous with the cavity of the urogenital 2. The penis may be double or bifid.
sinus (Fig. 16.26F). The margins of this 3. Rarely, the penis may lie posterior to the
groove are called the definitive urethral scrotum.
folds. 4. The urethral folds may fail to fuse,
These folds now approach and fuse with partially, or completely. When failure to
each other. The fusion begins posteriorly fuse is complete the scrotum is in two
in the region of the urogenital sinus and halves and the genitals look like those of
extends forwards onto the phallus the female (Fig. 16.27A). If the defect is
(Figs 16.26G, H). The penile urethra is confined to the anterior part of the phallus,
formed as a result of this fusion. It will the urethra opens on the undersurface of
now be apparent that the wall of the penile the penis. This condition is called
urethra is made up of: hypospadias (Fig. 16.27B).
5. The urethra sometimes opens on the dorsal
(i) the original endodermal lining of the
aspect of the penis. The condition is called
phallic part of the urogenital sinus, and
epispadias, and is usually associated with
(ii) the endodermal cells of the urethral
ectopia vesicae. In such cases it is believed
plate.
that the genital tubercle is formed caudal
The penile urethra formed in this way to the urogenital membrane instead of
extends only up to the glans penis. The being ventral to it. When the membrane
distal-most part of the urethra is of ruptures, the urogenital sinus opens cranial
ectodermal origin and is formed by to the developing penis.
canalization of a solid mass of ectodermal
Other anomalies of the penile urethra have
cells (Figs 16.26G, H).
(e) The genital swellings fuse with each other, been described earlier.
in the midline, to form the scrotal sac into
Path taken by
which the testes later descend. primordial germ cells
to reach gonad Mesentery
Prenatal Diagnosis of Sex
The sex of a baby can be determined before
birth by ultrasound examination. The penis can
be seen in a male child.
In this connection it has to be noted that in
fetuses about three to four months old, the
genital tubercle is equally developed in both
the male and female. Ultrasound examination
at this stage can be misleading as the clitoris
can be mistaken for a penis.
Yolk sac
Anomalies of M a l e External Genitalia
Fig. 16.28 Migration of primordial germ cells from
1. The entire penis may be absent.
the neighbourhood of the yolk sac to the
Alternatively the corpora cavernosa or the developing gonad.
Human Embryology

Primordial
germ celfs

Mesonephric toejifial coelomic bay


duct and tubule
Mesentery

Thickened coelomic
epithelium forming
genital ridge

Some cells of sex


cords surround
Enlarging sex cords Degenerating primordial germ cells,
mesonephric duct l\

Mesonephric duct
becomes duct of
epididymis Mesonephric Primordial follicles
, tubules forming
V—. vasa efferentia

C Rete-
O
testis

Seminiferous
tubules formed
by canalization
of sex cords Germinal epithelium

Fig. 16.29 Development of gonads. (A) Indifferent stage. (B) and (C) Testis. (D) and (E) Ovary.
Urogenital System

Anomalies of Female External Genitalia that covers the medial side of the mesonephros, of
the corresponding side (Fig. 16.29). In the region
1. The clitoris may be absent, may be bifid, where the testis is to develop, this germinal
or may be double. It may be enlarged in epithelium becomes thickened. This thickening is
hermaphroditism. called the genital ridge. The cells of the germinal
2. The labia minora may show partial fusion. epithelium proliferate and form a number of solid
3. The urethra may open on the anterior wall sex cords, that grow into the underlying
of the vagina; this is the female equivalent mesenchyme. They reach deep into the gonad and
of male hypospadias. are called medullary cords. They are soon
canalized to form the seminiferous tubules.
Meanwhile, the primordial germ cells migrate to
Primordial Germ Cells (Fig. 16.28)
the region of the developing testis and get
The cells of the ovaries and the testes, from which incorporated in the seminiferous tubules.
germ cells are formed, are believed to be
segregated early in the life of the embryo. The interstitial cells of the testis, are derived
They probably differentiate in the wall of the from sex cords that are not canalized. Some of
yolk sac and migrate to the region of the them are also derived from the surrounding
developing gonads. mesenchyme.
All spermatozoa and ova that are formed The mesenchymal cells, surrounding the
throughout the life of the individual are believed developing testis, form a dense layer of fibrous
to arise from these primordial germ cells. tissue. This is the tunica albuginea. It
Gonads do not develop as long as primordial completely separates the sex cords from the
germ cells do not reach them. These cells have an germinal epithelium and, thereafter, this
inducing erred on rhc gonad. epithelium can make no further contribution to
testicular tissue.

DEVELOPMENT OF TESTES Duct System of Testes

Hach testis develops from the coelomic epithelium, We have seen, above, that the testis develops in

Seminiferous
tubules Seminiferous
tubule

Fig. 16.30 Development of duct system of the testis. Structures derived from sex cords are shown in grey.
Human Embryology

testes, in turn, establishes contact with persisting


mesonephric tubules which form the vasa
efferentia. The cranial part of the mesonephric
duct becomes highly coiled on itself to form the
epididymis while its distal parr becomes the ductus
deferens. The seminal vesicle arises, on either side,
as a diverticulum from the lower end of the
mesonephric duct. The part of the mesonephric
duct that lies between its opening into the prostatic
urethra and the origin of this diverticulum, forms
the ejaculatory duct.

Descent of Testes
The testes develop in relation to the lumbar region
of the posterior abdominal wall. During fetal life,
theyigradually descend to the scrotum. They reach
Fig. 16.31 Descent of the testis (from the lumbar
the iliac fossa during the third month, and lie at
region to the scrotum),
the site of the deep inguinal ring up to the seventh
month of intrauterine life. They pass through the
close proximity to the mesonephros and the inguinal canal during the seventh month, and are
mesonephric duct. We have also seen that most of normally in the scrotum by the end of the eighth
the mesonephric tubules degenerate. Some of these month (Fig. 16.31).
that lie near the testis persist and, along with the
mesonephric duct, form the duct system of the testis
(Fig. 16.30). The descent of the testes is caused or assisted
by several factors. These are:
The ends of the seminiferous tubules anastomose
with one another to form the rete-testes. The rete- 1. Differential growth of the body wall.

Peritoneum
Transversus abdominis
and internal oblique m.
Skin

F. transversalis

Gubernaculum

Ext. oblique aponeurosis

Fig. 16.32 The gubernaculum, w h i c h helps in descent of the testis.


Urogenital System

Fig. 16.33 Relation of descending testis to processus vaginalis. Note that as the testis descends it
progressively invaginates the processus vaginalis.

Formation of inguinal bursa: About the c o r r e s p o n d i n g increase in l e n g t h .


sixth month of intrauterine life, the various There is thus a relative shortening of
layers of the abdominal wall, of each side, the gubemaculum and, as a result, the
show an outpouching towards the scrotum testis assumes a progressively lower
(Fig. 16.32). This p o u c h progressively position.
increases in size and depth, and eventually (b) The gubemaculum helps to dilate the
reaches the bottom of the scrotal sac. The inguinal bursa.
descending testis enters this pouch to reach (c) It provides a continuous pathway for
the scrotum. Note that the pouch is formed the descending testis.
before the testis enters it. The cavity of
Processus vaginalis: This is a diverticulum
the inguinal bursa becomes the inguinal
of the peritoneal cavity. It actively grows
canal, while the various layers of its wall
form the c o v e r i n g s of t h e t e s t i s a n d
spermatic cord.
The gubemaculum: This is a b a n d of
mesenchyme which extends from the lower
pole of the testis to the scrotum. For many
years, it was believed that descent of the
testis was caused by shortening of the
gubemaculum. However, we now k n o w
t h a t this is n o t p o s s i b l e b e c a u s e the
g u b e m a c u l u m does not contain any
c o n t r a c t i l e tissue. A c c o r d i n g t o some
authorities the g u b e m a c u l u m does not
reach the scrotum but reaches the bottom
of the inguinal bursa. In spite of this, the
g u b e m a c u l u m does play an i m p o r t a n t
part in the descent of the testis as follows: Fig. 16.34 Ectopic positions of the testis: (a) under
skin of abdomen; (b) over front of thigh;
(a) When the embryo increases in size, (c) in femoral canal; (d) under skin of
the gubemaculum does not undergo a penis; (e) in perineum.
Human Embryology

into the gubernacular mesenchyme of the the scrotum. Some interesting facts about
inguinal canal and of the scrotum (Fig. 16.32). this condition are as follows:
As the testis descends, it invaginates the
(a) The testis may complete its descent
processus vaginalis from behind. After the
descent of the testis is completed, the after birth.
processus vaginalis loses all connection (b) Spermatogenesis often fails to occur
with the peritoneal cavity and becomes the in an undescended testis.
tunica vaginalis (Fig. 16.33). (c) An undescended testis is more likely
5. The descent of the testis is greatly to develop a malignant tumour than
influenced by hormones secreted by the a normal testis.
pars anterior of the hypophysis cerebri. (d) The condition can be surgically
corrected.
Anomalies of Testis 5. Abnormal positions (Ectopia): The testis
1. The testis may be absent, on one or both may lie (Fig. 16.34):
sides. (a) Under the skin of the lower part of the
2. The testis may be duplicated. abdomen.
3. The two testes may be fused together. (b) Under the skin of the front of the thigh.
4. Anomalies of descent (Cryptorchidism): (c) In the femoral canal.
Descent of the testis may fail to occur, or (d) Under the skin of the penis.
may be incomplete. The organ may lie in (e) In the perineum behind the scrotum.
the lumbar region, in the iliac fossa, in
the inguinal canal, or in the upper part of 6. Also see hermaphroditism.

Congenital hernia Vaginal hydrocoele Funicular hernia


Congenital hydrocoele

Infantile hernia Interstitial hernia Hydrocoele of cord Infantile hydrocoele

Fig. 16.35 Anomalies of processus vaginalis. Abnormal persistence of the processus vaginalis can lead to
hernia (passage into it of abdominal contents, indicated by arrows); or hydrocoele (collection of
fluid, shown as dots). Various types of hernia and hydrocoele are s h o w n .
Urogenital System

Anomalies of Duct System of Testis


Superior
aberrant ductules
1. The seminiferous tubules may fail to estab-
lish connection with the vasa efferentia.
2. The ductus deferens may be absent, in
whole or in part, on one or both sides.
3. The ductus deferens may have no
connection with the epididymis.

Anomalies of the Processus Vaginalis


We have seen that the part of the processus
vaginalis, which extends from the deep inguinal
ring up to the tunica vaginalis, normally
disappears. This may persist in whole or in part. Seminal
vesicle
Abdominal contents may enter it to produce
various forms of inguinal hernia. Alternatively,
fluid may accumulate in it producing the
condition called hydrocoele. Various forms of
Fig. 16.36 (Al Mesonephric duct, early stage.
hernia and of hydrocoele are shown in
(B) Mesonephric duct in the male,
Fig. 16.35.
before descent of the testis.

Uterine tube

Epoophoron U{enj&

Ovary

Paradidymis

Epididymis

Testis Inf. aberrant ductules

Fig. 16.37 Some structures derived from the mesonephric ducts. (A) In the male these are the epididymis, the
ductus deferens, the seminal vesicles and ejaculatory ducts. The appendix of the epididymis is a
vestigial remnant. (B) In the female most of the duct disappears. Some remnants are seen as the
epoophoron. For complete list of derivatives of the mesonephric ducts see text.
Human Embryology

Vestigial Structures in the Region of the Testis germinal epithelium may contribute to the
ovary even in postnatal life.
A number of vestigial structures are to be seen
in the n e i g h b o u r h o o d of t h e testis. T h e i r
Descent of the Ovary
importance lies in the fact that any one of them
may enlarge to form a cyst. The ovary descends from the lumbar region,
These structures are: where it is first formed, to the true pelvis. A
g u b e r n a c u l u m f o r m s , as in the m a l e , a n d
(a) Appendix of testis (also called hydatid
extends from the ovary to the labium majus. It
of Morgagni).
becomes attached to the developing uterus at
(b) Appendix of epididymis. its junction with the uterine tube. T h e part of
(c) Superior aberrant ductules. the gubernaculum that persists between the
(d) Inferior aberrant ductules. ovary a n d the uterus becomes the (round)
(e) Paradidymis.
ligament of the ovary. The part between the
uterus and the labium majus becomes the round
DEVELOPMENT OF THE OVARY ligament of the uterus.

T h e early stages in the d e v e l o p m e n t of t h e


Anomalies of Ovary
o v a r y a r e e x a c t l y t h e s a m e as in the testis
(Figs 16.29A, C, E). 1. The ovary may be absent on one or both
sides.
1. Thecoelomic epithelium on the medial side 2. The ovary may be duplicated.
of the mesonephros becomes thickened to 3. T h e o v a r y m a y d e s c e n d i n t o t h e
form genital ridges. inguinal canal or even into the labium
2. Cords of cells (sex cords or medullary cords) majus.
proliferate from this germinal epirhelium, 4. Adrenal or thyroid tissue may be present
and grow into the underlying mesoderm. in t h e o v a r y . T h e o v a r y s o m e t i m e s
3. Primordial germ cells, which are formed in c o n t a i n s cells t h a t a r e c a p a b l e of
relation to the yolk sac, migrate to the region differentiating into various tissues like
of the developing ovary and give rise to bone, cartilage, hair, etc., and the growth
oocytes. of these cell rests can give rise to a peculiar
4. The sex cords become broken up into small t u m o u r called a teratoma.
masses. The cells of each mass surround one
primordial germ cell, or oocyte, to form a FATE OF MESONEPHRIC DUCT AND
primordial follicle. TUBULES IN THE MALE
According to some authorities the original The mesonephric ducts give rise to the following
(medullary) sex