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Embryology: Department of Anatomy & Histology, Khulna Agricultural University
Embryology: Department of Anatomy & Histology, Khulna Agricultural University
Embryology
Father of Embryology: Aristotle
Embryology is the study of embryo and fetus. It is the science that deals with the
origin and development of an individual organism which may be broadly
designated as embryo. It is the study of prenatal development of an individual
organism.
Or
Embryology is the branch of biological science which deals with the study of the
formation of zygote and its gradual development into an independent organism,
either in uterus or in egg shell, upto its individual existence.
Or
Embryology is the study of the growth and development of the embryo and fetus
of an organism from fertilization to extra-uterine life or conception to birth.
Objectives/Scope/Importance of Embryology:
1. To know the origin & development of germ cell.
2. To explain the mode of development from a zygote into an independent
individual.
3. To understand the normal & abnormal development of embryo.
4. To determine the sex of a new individual.
5. To understand the evolution.
6. To know the teratological defects.
7. To apply the knowledge of embryology towards pathology, immunology and
other applied subjects.
8. To increase the conception rate of animal.
9. In commercial purpose- cloning of animals and its marketing.
Methods of study:
1. Dissection: Gross dissection of embryo at different age.
2. Microscopic section: At different age of animal embryo.
3. Descriptive study: Materials, study, section and photography.
4. Experimental study: By using various drugs, chemicals, physical or
physiological means.
a) Genetic factors:
The "gene architecture" of its chromosomes is responsible for the control of
embryonic development.
b) Environmental factor:
Temperature (hypothermia, hyperthermia )
Anoxia
Adverse climate etc.
• Fetal insulin
• Fetal glucocorticoid
Further growth factors exist that affect the proliferation, differentiation and
maturation of the cells. They play an important role in embryogenesis. They are:
Periods:
Embryonic Period — defined as the time from fertilization to the earliest
(primordial) stages of organ development (about 30 days in dog, cat, sheep, pig;
almost 60 days in horse, cattle, human).
Fetal Period — It is the time between the embryonic period and parturition (the
end of gestation), during which organs grow and begin to function.
EMBRYOGENESIS
Embryogenesis is the process by which the embryo is formed and develops, until
it develops into a fetus. Embryogenesis includes the following phases:
1. Gametogenesis
2. Fertilization
3. Cleavage & Blastulation
4. Gastrulation
5. Organogenesis
6. Growth &
7. Differentiation
GAMETOGENESIS
Male or female reproductive germ cells are known as gamete. Gametogenesis is
the process of production of gamete in the gonads (i.e. testis in male and ovary in
female). Gametogenesis is two types: Spermatogenesis & Oogenesis.
Spermatogenesis
Spermatogenesis is the process by which spermatozoa is produced. This process takes
place in the seminiferous tubules of testis of male animals and the spermatozoa is
developed from the spermatogonium. In other word, the sequence of events of
formation of spermatozoa is referred to as spermatogenesis.
A-spermatogonium
B-spermatogonium
Primary spermatocyte
Secondary spermatocyte
Spermatid
Sperm cell (spermatozoon)
a) Spermatocytogenesis,
b) Meiosis &
c) Spermiogenesis.
Mitosis of the
16 days Up to the primary spermatocytes
spermatogonia
Total - 64 days
Spermatocytogenesis
Type A spermatogonia are the largest spermatogonia and share an extensive contact
area with the tubular basal lamina. They possess prominent nuclei with prominent
nucleoli and a pale or cloudy appearance. Intermediate spermatogonia are in between
type A & B spermatogonia. Type B spermatogonia have spherical nuclei containing
numerous chromatin particles and less prominent nucleoli. The mitotic division of
Type B spermatogonia results in the formation of primary spermatocytes.
Meiosis
During meiosis, two successive nuclear divisions occur (1 st & 2nd maturation division),
resulting the formation of four haploid spermatids from one primary spermatocyte.
The first maturation division proceed following the phases: the prophase (subdivided
into leptotene, zygotene, pachytene, diplotene & diakinesis), metaphase, anaphase
& telophase and resulting the formation of two secondary spermatocytes from one
primary spermatocyte. Each secondary spermatocyte possesses only half the number
(haploid) of chromosomes but have two chromatids each. Secondary spermatocytes are
short-lived, intermediate in size between primary spermatocytes and spermatids.
Spermiogenesis (Spermatohistogenesis)
Spermiogenesis is divided into Golgi phase, cap phase, acrosomal phase, and
maturation phase. In routine histologic sections, the Golgi & cap phases are
characterized by spherical nucleus and the acrosomal & maturation phases are
characterized by elongated nucleus.
Golgi phase: During the Golgi phase, proacrosomal granules appear in the Golgi
vesicles.
Cap phase: During the cap phase, the acrosomal vesicle grows and forms the head cap
that covers the anterior two-thirds of the nucleus.
Acrosomal phase: During this phase,
- The nucleus & cell body start to elongate in a cranio-caudal direction.
- Formation of acrosomal cap.
Maturation phase: During this phase,
- Nuclear condensation is completed.
- Excess cytoplasm is detached as residual body.
Spermination
Spermination is the release of the spermatozoa from and/ between the Sertoli cells into
the lumen of the seminiferous tubules.
Intermediate spermatogonia
Type B spermatogonia
Primary spermatocytes
Secondary spermatocytes
Spermatids
Spermatozoa
Spermatozoa
A spermatozoa is a slender, motile flagellate body, the male germ cell. It has been
estimated that about 60,000 spermatozoa contained within 1mm3 seminal fluid or
200-600 million in a single ejaculation. The concentration of spermatozoa varies
greatly among different species. Spermatozoa vary in length between
approximately 60 μm (boars, stallions) to 75 μm (ruminants). With the light
microscope, the spermatozoon seems to consist essentially of two portions: the
head & the tail (flagellum, use to swim through liquids). With the electron
microscope, the tail is seen to be subdivided further into neck, middle piece,
principle piece, and end piece.
Head: The shape of the nucleus determines the shape of the head of the
spermatozoon, which is species-dependent and subject to great variations. The
anterior pole of the nucleus is covered by the acrosomal cap, with an outer & inner
acrosomal membrane that fuse at the caudal end. The acrosomal cap contains
several hydrolytic & proteiolytic enzymes (eg. acrosin), which are become free
during the acrosomal reaction of capacitated spermatozoa in the uterine tube.
Acrosomal enzymes are needed for the penetration of the zona pellucida during
fertilization. The caudal end of the acrosome is characterized by a narrowing of the
cap and condensation of its contents. The base of the nucleus is surrounded by the
postacrosomal sheath, which consists of fibrous proteins rich in sulfur. The plasma
membrane of the post postacrosomal head region contains receptor molecules
necessary for the recognition of homologous oocyte. At the caudal surface of the
sperm head, the nuclear envelop lines an implantation groove where the tail is
inserted in a joint manner.
Neck: The neck is a relatively short & narrow structure between the head &
middle piece (connecting piece). It consists of a centrally located centriole and nine
peripheral longitudinally oriented coarse fibers continuous with the outer fibers of
the middle piece.
Middle piece: The core of the middle piece has the characteristic structure of a
flagellum: two central microtubules & nine peripheral doublets (microtubules)
making up the axial filament complex. They are surrounded by nine longitudinally
oriented outer fibers that are connected to the fibers of the connecting piece. These,
in turn, are surrounded by the mitochondria (essential for the sperm to successfully
survive on its journey to the female egg) in a helical arrangement. A ring-shaped
thickening of the plasma membrane (annulus) of the middle piece marks the limit
between the middle piece and the principle piece.
Principle piece: The principle piece is the longest portion of the spermatozoon.
The axial filament complex has a structure identical to that of the middle piece and
is surrounded by the continuing outer fibers of the middle piece. The fibers are
subject to variation in size & shape and gradually taper toward the end of the
principle piece.
End piece: The termination of the fibrous sheath marks the beginning of the end
piece, which contains only the axial filament complex. Proximally, in the end
piece, this complex has its characteristic nine peripheral doublets; distally, these
doublets gradually become reduced to singlets and terminate at various levels.
A sperm cell contains certain elements that allow it to remain "alive" for a short
period of time. These elements are various, but the most important are a plasma
membrane (effectively sealing the cell, allowing only certain things to come
through) and mitochondria (energy-producing organelles).It can only survive in the
male reproductive tract, in the female reproductive tract (only when certain
conditions are present) and in certain artificial environments.
Acrosome: The acrosome is a membrane covering the anterior 2/3 of the sperm
nucleus. It contains several hydrolytic & proteolytic enzymes, which are become
free during the acrosomal reaction of capacitated spermatozoa in the uterine tube.
Acrosomal enzymes are needed for the penetration of the zona pellucida during
fertilization. The enzymes include neuroaminidase, hyaluronidase, aryl
phosphatase, aryl sulfatase, acrosin.
Oogenesis:
Oogenesis is the process by which the female gamete or ovum or the female sex cell or
the female germ cell is produced. It is the female form of gametogenesis. Oogenesis
consists of several sub-processes: oocytogenesis, ootidogenesis, and finally maturation
to form an ovum (oogenesis proper).
Primordial Germ cell multiply mitotically resulting increased in number and become
ooginia and some of these oogonia go in the process of oogenesis.
Oocytogenesis
Oogenesis starts with the process of developing oogonia, which occurs via the
transformation of primordial follicles into primary oocytes, a process called
oocytogenesis. Oocytogenesis is completed either before or shortly after birth.
Ootidogenesis
The succeeding phase of ootidogenesis by which the primary oocyte develops into
an ootid by the process of meiosis.
However, although this process begins at prenatal age, it stops at prophase I. In late
fetal life, all oocytes, still primary oocytes, have halted at this stage of
development, called the dictyate. After menarche, these cells then continue to
develop, although only a few do so in every estrus / menstrual cycle.
Meiosis I
Meiosis I of ootidogenesis begins during embryonic development, but halts in the
diplotene stage of prophase I until puberty. (For those primary oocytes that
continue to develop in each estrus / menstrual cycle, however, synapsis occurs and
tetrads form, enabling Chromosomal crossover to occur.) As a result of meiosis I,
the primary oocyte has developed into the secondary oocyte and the first polar
body.
Meiosis II
Immediately after meiosis I, the haploid secondary oocyte initiates meiosis II.
However, this process is also halted at the metaphase II stage until fertilization, if
such should ever occur. When meiosis II has completed, an ootid and another polar
body have now been created.
Fig: Oogenesis
Spermatogenesis Oogenesis
6) Four spermatozoa are produced from 6) Only a single ovum is produced from
each primary sprematocyte. each primary oocyte.
7) Polocytes / polar bodies are not formed. 7) Polocytes are formed so that mature egg
may have haploid number of
chromosomes.
During further development, the granulosa cells increase their FSH receptors and
express LH receptors, while the Theca cells increase their LH receptors. This surge in
hormone reception results in ovulation. After ovulation, if fertilization occurs, the
corpus luteum secretes progesterone that supports the further growth of the
endometrium. If, however, fertilization does not take place, then the hormone levels
drop, the corpus luteum breaks down, no longer secreting progesterone, so that the
endometrium sloughs off producing menstruation.
FERTILIZATION
Fertilization (also known as 'conception', 'fecundation' and 'syngamy'), is the fusion of
gametes to produce a new organism. In animals, the process involves a sperm fusing
with an ovum, which eventually leads to the development of an embryo. The fertilized
egg cell is known as the zygote.
Depending on the animal species, there are two types of fertilization: External
Fertilization & Internal Fertilization
External Fertilization
External fertilization occurs mostly in wet environments and requires both the male
and the female to release their gametes into their surroundings (usually water). An
advantage of external fertilization is that it results in the production of a large number
of offspring. One disadvantage is that environmental hazards such as predators greatly
reduce the chance of surviving into adulthood. Amphibians and fish are examples of
animals that reproduce this way.
Internal Fertilization
Internal fertilization occurs within the body of the female animals. Animals that use
internal fertilization specialize in the protection of the developing egg. For example,
reptiles and birds secrete eggs that are covered by a protective shell that is resistant to
water loss and damage. Mammals take this idea of protection a step further by
allowing the embryo to develop within the mother. This extra protection increases the
chances of survival because mom supplies everything that the embryo needs. In fact,
most mammalian mothers continue to care for their young for several years after birth.
Fertilization is a chain of events. Interruption of any step in the chain will almost
certainly cause fertilization failure. The chain begins with a group of changes affecting
the sperm, which prepares them for the task ahead.
Process of fertilization:
In overview, fertilization can be described as the following steps:
a) Transport of spermatozoa
About 200-300 million of spermatozoa deposit along with male accessory sex
glandular fluid in the female reproductive tract. Among them 300-500 reach the site of
fertilization, but only one is reached for fertilization and others help in barrier
penetration. By this time, sperm become capacitated and only the capacitated sperm
pass through the corona radiata and undergoes the acrosome reaction.
Sperm Capacitation
Freshly ejaculated sperm are unable or poorly able to fertilize. Rather, they must first
undergo a series of changes known collectively as capacitation. Capacitation is the
process by which sperm becomes capable of fertilizing ovum after it reaches the
ampulla of the uterine tube. Capacitation is associated with removal of adherent
seminal plasma proteins, reorganization of plasma membrane lipids and proteins. It
also seems to involve an influx of extracellular calcium, increase in cyclic AMP, and
decrease in intracellular pH. Sperm that have undergone capacitation are said to
become hyperactivated, and display hyperactivated motility. Only the capacitated
sperm pass through the corona radiate and undergoes the acrosome reaction.
b) Barrier penetration
c) Sperm-Oocyte Binding
Once a sperm penetrates the zona pellucida, it binds to and fuses with the plasma
membrane of the oocyte (oolema). Binding occurs at the posterior (post-acrosomal)
region of the sperm head. A sperm glycoprotein called fertilin, which binds to a
protein in the oocyte plasma membrane and may also induce fusion.
Prior to fertilization, the egg is in a quiescent state, arrested in metaphase of the second
meiotic division. Upon binding of a sperm, the egg rapidly undergoes a number of
metabolic and physical changes that collectively are called egg activation. Prominent
effects include a rise in the intracellular concentration of calcium, completion of the
second meiotic division and the so-called cortical reaction.
Cortical Reaction:
The cortical reaction refers to a massive exocytosis of cortical granules seen shortly
after sperm-oocyte fusion. Cortical granules contain a mixture of enzymes, including
several proteases, which diffuse into the zona pellucida following exocytosis from the
egg. These proteases alter the structure of the zona pellucida, inducing what is known
as the zona reaction.
Zona Reaction:
The zona reaction refers to an alteration in the structure of the zona pellucida catalyzed
by proteases from cortical granules. The critical importance of the zona reaction is that
it represents the major block to polyspermy in most mammals. This effect is the result
of two measurable changes induced in the zona pellucida:
1. The zona pellucida hardens. Runner-up sperm that have not finished traversing
the zona pellucida by the time the hardening occurs are stopped in their tracks.
2. Sperm receptors in the zona pellucida are destroyed. Therefore, any sperm that
have not yet bound to the zona pellucida will no longer be able to bind and
fertilize the egg.)
The male and female pronuclei fuse together, their membranes break down, and the
two genomes condense into chromosomes, thereby reconstituting a diploid organism
(zygote).
Zygote: The body or structure formed by the union of the male and female pronuclei
due to fertilization and having the capability of mitosis is known as zygote
Results of fertilization:
Formation of zygote.
Restoration of diploid number of chromosome.
Determination of sex in the new individual.
The cleavage process is started.
Before gastrulation, the zona pellucida fully disappears and the inner cell mass become
very prominent and the trophoblast cells become flattened. When the zona pellucida is
disappeared, the blastula attaches with the endometrium which is termed as
implantation.
The trophoblast cells become differentiated into two strata: the outer stratum termed
the syncytiotrophoblast; while the inner layer, the cytotrophoblast or “Layer of
Langhans”. The cytotrophoblast cells are mitotically active but the syncytiotrophoblast
cells are not. The syncytiotrophoblasts invade the endometrium with three
characteristics: 1) they have invasive power, 2) they have ingestive character & 3) they
have digestive character. By invading, they get nutrition from the exposed glands &
blood vessels of the endometrium. Subsequently, the trophoblasts form placenta &
extra-embryonic (fetal) membranes and the inner cell mass give rise to future embryo.
Cleavage:
It refers to the initial series of mitotic divisions by which the large zygote is
fractionated into numerous “normal size” cells. each daughter cell of the cleavage
process is termed a blastomere. cleavage begins with a zygote, progresses through
compaction to a morula stage and terminates at the start of the blastocyst (blastula)
stage the first eight blastomeres are undifferentiated and have identical potential in
mammals; thereafter, blastomeres differentiate into inner & outer cells with
different missions
Morula
It is a solid ball of blastomeres within a zona pellucida (typically consisting of 16
to 64 blastomeres) blastomeres become compacted; cells on the inside differentiate
from those along the surface of the morula:
outer blastomeres become flattened and form tight junctions (reducing fluid
permeability); they develop the capacity to secrete fluid (internally); they are
destined to become trophoblasts which form the chorion & amnion (fetal
membranes) of the conceptus;
Note: As few as three inner blastomeres are sufficient to produce an entire embryo
(and adult). When a morula leaves the uterine tube and enters the uterus (uterine
horn) it is at about the 16-cell stage, around 4 to 7 days after fertilization
(depending on species). The 32-cell stage morula (5-7 days post ovulation) is ideal
for embryo transfer in cattle.
Implantation
The blastocyst is initially free in the uterine lumen (nourished by uterine glands).
Implantation of the blastocyst is a gradual process, beginning with apposition,
leading to adhesion (or invasion in the case of the human & Guinea Pig).
Approximate implantation times are: one week (human); two weeks (dog, cat,
sheep), 3-5 weeks (cattle), 3-8 weeks horse; or delayed up to 4 months (deer,
bears).
TWINS
Monozygotic:
Dizygotic: fraternal twins result when two zygotes develop “independently” during
the same pregnancy (independence can be compromised by fusion of fetal
membranes and blood supplies). It is possible for fraternal blastomeres to merge
and produce a single conceptus with two different genotypes (a chimera).
Epiblast cells proliferate along primitive streak margins and migrate through the
streak into the coelom. The migrating cells form endoderm & mesoderm layers.
Initial migrating cells join the hypoblast layer, forming embryonic endoderm
(hypoblast cells constitutes yolk sac endoderm).
The majority of migrating cells enter the coelom as primary mesenchyme and
become mesoderm. The primary mesenchyme migrates laterally and cranially
(but not along the midline region directly cranial to the primitive streak where
notochord will form). Note: Mesoderm divides into: paraxial, intermediate, and
lateral mesodermal regions.
Ectoderm produces: 1) Outer epithelium of the body: the epidermis and epidermal
derivatives of the integumentary system, including the hair follicles, nails, and the
glands that communicate with the skin surface (i.e., the sudoriferous, mammary, and
sebaceous glands). The lining of the mouth, salivary glands, nasal passageways, and
anus develop from ectoderm, as does portions of the skull, pharyngeal arches, teeth,
and endocrine system (pituitary and parts of adrenal glands); 2) Neural tube: brain,
spinal cord, neurohypophysis, optic nerve, retina, motor spinal nerve root; 3) Neural
crest: spinal dorsal root ganglia, cranial sensory ganglia etc.
Mesoderm produces the lining of the pleural, pericardial, and peritoneal cavities, the
muscular, skeletal, cardiovascular, and lymphatic systems, the kidneys and portions of
the urinary tract, the gonads and most of the reproductive tracts, and the connective
tissues that support all organ systems. Portions of the endocrine system (parts of the
adrenal glands and endocrine tissues of the reproductive system) also develop from
mesoderm.
Head mesenchyme → Cephalic connective tissue & muscle, dentine, outer layer of
eye, skull.
Endoderm produces most of the epithelium of the digestive system (except the mouth
and anus), the exocrine glands (except salivary glands), and the liver and pancreas.
Most of the respiratory system, including the epithelium (except that of the nasal
passageways) and mucous glands develop from endoderm, as does portions of the
urinary and reproductive systems (the stem cells that produce gametes). Pharyngeal
pouches (first pouch – middle ear, auditory tube, endodermal lining of tympanic
membrane ; second pouch – tonsillar recess; third pouch – thymus, thyroid; fourth
pouch – parathyroid, post bronchial body) also develop from endoderm.
GROWTH
Growth may be defined as the developmental increase in mass. After the organ
rudiments are formed they begin to grow and greatly increase their volume. Growth
results from synthesis of new protoplasm, both the cytoplasmic and nucleic. Increase
in mass is usually accompanied by cell division and also by increasing of intercellular
substances. In this way, the animal gradually achieves the size of its parents.
DIFFERENTIATION
Differentiation refers to the events by which cells and other parts become different
from one another. Differentiation dramatically changes a cell's size, shape, membrane
potential, metabolic activity, and responsiveness to signals. After growth the cells in
each rudiment become differentiated, i.e., they acquire a diversification of form and
function.
1) Morphological differentiation
Individual cells and their groups become structurally different from other cells and
groups of cells. For example, from a common starting point in generalized ectoderm,
epidermal and nerve cells acquire the distinguishing characteristics of size, shape and
internal structure.
2) Behavioral differentiation
Although the cells exhibit common basic properties like metabolism, irritability, some
cells possess some special capabilities along the general properties. For example, nerve
cells take up the responsibility to transport electrical impulses, muscles to contract,
gland cells to secrete special secretions and so on.
3) Chemical differentiation
The individual cells become biochemically different from one another as the egg
undergoes cleavage and progressively becomes converted to blastula, gastrula and
definitive embryo.
Fetal membrane
The fetal membrane surrounds the fetus during pregnancy. Four fetal membranes
develop in a conceptus. Two arise from the trophoblast layer of the blastocyst and
are continuous with the somatopleure of the embryo. Two arise from the inner cell
mass of the blastocyst and are continuous with splanchnopleure of the embryo;
these two splanchnopleure membranes are vascular. The four fetal membranes are:
1. Chorion — forms the outer boundary of the entire conceptus (from trophoblast)
Note: The term conceptus refers to the embryo or fetus plus its fetal membranes.
Functions:
During pregnancy:
o Protects the foetus against injury.
o A medium for free foetal movement.
o Maintains the foetal temperature.
o Source for nutrition of the foetus.
o A medium for foetal excretion.
During Labour:
o The fore-bag of water helps the dilatation of the cervix during labour.
o It acts as an antiseptic for the birth canal after rupture of the
membranes.
Placenta
The placenta connects the developing fetus to the wall of the mother’s uterus
during pregnancy. It grows in the wall of the uterus and is attached to the fetus
within the uterine cavity by the umbilical cord. The placenta is formed by cells that
originate from the fetus and is therefore the first of the fetal organs to
develop. Placenta is the region(s) of apposition between uterine lining and fetal
membranes where metabolites are exchanged for sustaining pregnancy.
Diffused type placenta: The villi are uniformly distributed on the surface of
blastocyst, except at the extreme ends. Ex: Horse, pig.
Cotyledonary placenta: The villi are arranged in groups. Each group is called
cotyledon. Each cotyledon fits into caruncle of uterus. Ex: Sheep, Cow, Deer.
Zonary placenta: The villi are in the form of transverse zones. In dog, a single
girdle of villi will be present. In fox, two girdles of villi are present. The villi
penetrate into the uterine wall. Hence during parturition uterine wall is damaged.
Ex: Cat, Dog, Carnivores.
Discoidal Placenta: On the entire surface of blastocyst, the villi are in the form of
discs. When the embryo is growing it moves away from uterus hence the with look
like a disc. These villi are intimately connected with uterus. Hence during
parturition much uterine tissue is damaged. Ex: Rat, Bat, Rabbit.
According to HISTOLOGY:
Just prior to formation of the placenta, there are a total of six layers of tissue
separating maternal and fetal blood. There are three layers of fetal extraembryonic
membranes in the chorioallantoic placenta of all mammals, all of which are
components of the mature placenta:
There are also three layers on the maternal side, but the number of these layers
which are retained - that is, not destroyed in the process of placentation - varies
greatly among species. The three potential maternal layers in a placenta are:
According to number of layers of cells present between fetus and uterus blood
supply the placenta is classified into five types:
c) Endothelio chorial placenta: The chorion of the fetus will come in contact
with the endothelium of mother‘s uterus. Ex: Dog, Carnivores.
FUNCTIONS OF PLACENTA:
1. Placenta forms a physiological barrier between mother and fetus.
2. Placenta allows the diffusion of monosacharides, amino acids, hormones,
vitamins, oxygen, carbondioxide, water and other waste materials to foetus.
3. It works as an excretory organ of fetus. It releases the nitrogenous waste
materials into mother blood.
4. It works as an endocrine gland. It secretes lactogen, progesterone, etc.
hormones.
5. Perform immunological activity.
A. Placentation:
1. Cheiloschesis (cleft lip), palatoschisis (cleft palate): Cleft lip is caused by failure
of fusion of medial nasal and maxillary processes; cleft palate is caused by failure
of medial palatine processes to fuse.
2. Heterotopic polyodontia (dentigerous cyst, “ear teeth”): Primordia of enamel
organs escape to the exterior and develop tooth structures anchored on the parietal
bone or base of the ear. These cause festering problems and must be relieved
surgically.
3. Thyroglossal duct cyst: Failure of involution of thyroglossal duct is the cause. A
surgically removable fluid-filled cyst seen at birth interferes with breathing.
C. Digestive tract.
G. Body cavities.
1. Pleuroperitoneal hernia: Failure of closure of one or both pleuroperitoneal folds
results in intestinal herniation into the pleural cavity.
2. Peritoneopericardial diaphragmatic hernia
J. Musculoskeletal system.
1. Premature physeal closure (achondroplasia, chondrodysplasia).
2. Spina bifida.