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DEPT. ANATOMY & HISTOLOGY

EMBRYOLOGY/DEVELOPMENTAL ANATOMY
VAN 211

Dr.GIRISH. M. H.
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Reference books:

1.Developmental Anatomy – Arey L.B.

2. Patten’s foundations of embryology –Carlson
3. Medical embryology – Langman
4.Veterinary embryology-Latshaw or Mc.Geddy
5.Embryology of domestic animals – Noden & Delahunta
6. An introduction to embryology – Balinsky

Definition: Embryology is study of embryos.

Embryo: is the juvenile stage of an individual organism. i.e. stage before the birth. May be
within the eggs(birds) or uterus of mammals.
So it can be defined as “the branch of science which deals with the origin and
development of an individual organism like chick, bird, cow, dog, horse, man.
Development: is the gradual bringing to perfection both structurally and functionally.
Difference b/w :
a.embryolgy-is study of embryology stops when embryo hatches or it is born.
b. developmental anatomy: upto adult or old age. So it has broader scope compared to
embryology.
Life of animal passes through 3 stages:
1.Prenatal stage(embryonic stage): stage before birth. It includes
a. stage of ovum: stage from fertilization to development of blastocyst (before germ layer
formation) i.e. 1st Week.
b. stage of embryo: stage of gastrulation to organogenesis. i.e. 2nd week to 8th week.
cow-12days to 45 days.
c. stage of foetus: it is above 45 days to stage of parturition. In this stage organogenesis is
complete and enormous increase in the size of foetus.

2.Birth
3.Postnatal stage: stage after birth. It includes
a. New born
b. Infant
c.Childhood
d.Adult
e.Senesence
f.death
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Historical background:
1. 384-322BC, Aristotle-wrote 1st book on embryology and described the development
of chicks.
2. 16th -17th C- nothing is added
3. 1677-Antony Von Levwenhock & Hamm-observed the human sperm under
microscope for first time.
4. 1672-De Graff- observed human ovarian follicle so called Graffian follicke
5. 1827-Von Baer, described mammalian germ layers.
6. Theories during 16th – 17th C
a.Preformation theory:states that an individual in a miniature form exist in the form of
sperm or ovum.by Haertsoeks in 1694
two opinion about this-1. Miniature form of an organism present with in ovum- ib by ovist
2. Miniature form of an organism present with in sperm is by animalicules
Hastsocker was in opinion that child exit in the head of sperm called humuncules.
b.Epigenesis/epigenetic theory: proposed by Wolff in 1759. He states that development
takes place from formless beginning to complex organization.
Development is epigenesist. ie.characters are inherited i.e. preformation. Eg. Color of eyes,
hairs, shape of nose i.e. merely transformation of genes from preexisting cell to organism
7. 1864-Pasteur- states that every cell arises from subdivision of preexisting cell.
8. 1900-Dreisch- stated “many forms of daughter cell of fertilized egg when they are
separated, they develop into separate individuals”.
9. 1961-Harvey, 1672-Malphigi, contributed to description of stages of developing
chicks as seen in simple lenses.
Development: it may be a. an individual development(cow, sheep, dog etc)
b. Racial development (broader group of animals like fish, mammals, amphibian)
Development of an individual animal is Ontogeny. Whereas group of animal or racial
development is Phylogeny.
Theory states “Ontogeny recapitulate phylogeny” eg. During Development of dog or
human – they passes through various stages of fishes, amphibian, reptiles, birds and then
mammals. It is not completely true but during development shows some characters so the
theory modified as “Ontogeny is a brief and incomplete recapitulation of phylogeny”.eg.:
a. in earlier stage there is development of Gill archs present in fish get modified into some
other structure i.e. pharyngeal arch.
b.during development of kidney it passes through pronephros(function in fish, amphibian),
mesonephros(in reptail) and metanephros stages(birds and mammals)
c. Aortic arches-6 develops but some get modified in development to suit the needs.
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Scope and importance of study of Embryology:

1. Study of embryology assist the student in understanding the gross anatomical
structure in animal ie. relationship which exist in one organ with another organ. Eg.
Aorta only on left side, venacava on right side, heart is single & in thoracic cavity,
ovaries are pair. Recurrent laryngeal nerve on left side winds round arch of aorta and
right side winds right subclavian artery is due to difference in aortic arches.
Trapezius muscle supply by 11th cranial nerve. This muscle develop from cuccularis
in fishes act as elevator of gill arches in fish. Modification of this in reptails, birds and
mammals become trapezium. And in fish and amphibian only 10 cranial nerves and
11th and 12th become modified spinal nerve.
2. To understand malformation/teratology/defect in birth.-so easy to correct using
corrective measure eg. Persistent foramen ovale, hydrocephalic condition, Cyclops
condition, cleft lip, septal defects etc.
3. To understand pathology and corrective surgery.
Branches of Embryology:
1.Descriptive embryology: studies basic structural pattern of embryonic development. His
and Bern, 1890- answers questions how and when it takes place in particular time.
______Bear is the father of embryology.
2.Comparative embryology: study involves comparative study of different types of embryos
of different species. eg. Rat, chick, pig, man embryos.
3.Experimental embryology: it studies the factor which regulate or activate the developmental
process. And answeres to questions why particular structure develop in particular way.
Spemann and Roux, 1852-1924, performed no. of experiments on chicken embryos and
putforth theory called developmental mechanics.
4.Chemical embryology: study of use of different chemicals and its effects on development of
embryo by Needham.
5.Teratology: study of abnormal development or malformations.
6.Reproductive biology: deals with applied aspect of embryology in different stage of
development eg. Embryo transfer technology, conception, fertilization etc.
General features of development:
1. Cell proliferation: ie. Increase in number of cells from unicellular to multicellular
organism. (zygote→divide→multicellular organism).
2. Cell growth: increase in mass. It may be due to increase in size of cell or increase in
number of cells. During growth there is a. protoplasmic synthesis b. increase uptake
of water c. intercellular deposition d. intracellular storage eg. Fat b/w cells.
3. Differentiation: transformation of a generalized cells into a cell with specific function.
Eg. Ectoderm-skin, nervous system
Mesoderm-vascular, urinary, genital system develops.
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Two methods of differentiation:

a.Histogenesis: is process where in substance and structure of cell progressively changes

giving rise to specialized structure and tissues. Eg.Mesoderm –genital, urinary, blood,
skeletal system. Connective tissue, cartilage.

b.Morphogenesis: there is moulding of different parts or organs to give a definite shape. it

involve morphogenetic processes which exists in different part during development. It
includes 1. Cell migration i.e. delamination and cell aggregation form.(Fig.A , page 1. 2a cell
mass, 2b Cords of cells, 2c sheath of cells)

2.Localised growth with constriction

3.Splitting
4.Folding-may be ingrowth(invagination or inpocketing) or outgrowth(evagination)
5.Bending
6.Degeneration- to remove excess parts.

Gametogenesis: is development of gamet or sex cells. Gamets are develops in the gonads
which may be ovary(female for ovum) or testis(male for sperms)
So it is development of sperm or ovum.

Development of sperm from primordial germ cell is spermatiogenisis and Development of

ovum/ova from primordial germ cell is Oogenesis.

Fusion of gamets give rise to zygote by fertilization.

2 types of chromosomes in gamets. They are 1. Autosomes 2. Sex chromosomes(always

paired)
Eg. Cow-60 chromosome→Autosome-58, Sex chromosome-2 ie. 29+x, 29+y
Man-46(23 pairs)→A-44, S-2,ie 22+x, 22+y.

Development/Origin of Gamet:
Gamets develops from primordial germ cells (which is precursor of gamet develop from
germinal epithelium). Primordial germ cells arise from yolk sac endoderm.

Embryo has 4 types of extraembryonic membranes ie yolk sac-which is connected to gut

tube.which is large in birds, and reptails and it is covered by inside endoderm and outside
mesoderm.

Yolksac endodermal cell becomes larger in size. They can be differentiated from other
endodermal cells by alkaline phosphatase and PAS positive reactions.ie very active cells
called primordial germ cells And these cells migrate from yolksac to developing gonad in
genital ridge. This migration is by two methods.

a. By vascular chanel – in birds

b. By Ameboid movement- in mammals.
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Attachement of yolksac to gonad is yolk stalk. Through yolk sac these primordial cell deposit
in gonad. With in gonad, these cells called as primordial germ cells or precursor which is
indifferent stage which may develop into male or female.

SPERMATOGENESIS:It is development/formation of sperm/spermatozoa in male from

primordial germ cells.

It takes place in 3 phases:

a. Spermatocytogenesis
b. Meiosis
c. Spermeogenesis

Section of Testis: shows Seminiferous tubules which has 2 types of cells-a. cells of
spermatogonic series b. sertoli cells(large pyramidal shape)

Primordial cell→spermatogonia cell which are found attached to basement membrane and
there are 2 types of spermatogonia cell

a. A type: oval in shape, obsl/spherical nucleus, light staining little chromatin.

b. B type: spherical shape, spherical nucleus, darker stained chromatin.

A type of spermatogonia cells divide into 6 time A1, A2 , A3 , A4 , A5 and A6 and cannot
become directly into primary spermatocyte( become B type) but B type cells divided into
primary spermatocyte.

A type
˄
A A1
˄
A A2
˄
A A3
˄
A A4
˄
A A5
˄
A A6 →B type
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Spermatogenesis:
Primordial germ cell→ Diploid
˄
A B → Transformation of gonia cell to
˄ Mitosis ↓spermatocytogenesis
2n Primary spermatocyte(diploid)
˄ ↓1st meiotic division
n n Secondary spermatocyte(haploid)
˄ ↓2nd meiotic division
n n Spermatids
↓ spermeogenesis
Spermatozoa/sperm
Ist Meiotic division:
1.Prophase: is very prolonged stage has 5 subphases.
a.Leptotene: thin strand chromosomes
b.Zygotene: pairing of homologus chromosomes
c.Pachytene: homologus chromosomes become short and thick, darkly stained.
d.Diplotene: initial crossing over of chromosomes into star and transformation of characters.
e.Diakinesis: crossing over is completed. Chromosomes starts separating to respective pole
then undergo
2.Metaphase
3.Anaphase
4.Telophase: giving rise to secondary spermatocyte. Here reduction in chromosome number
to haploid hence Reduction division.(2n→n). it takes number of days.

IInd Meiotic division: is from seconadary spermatocyte to spermatids. The process is same as
mitosis (i.e. no prolonged prophase stage).here no further reduction in chromosome number
hence it is Equational division. Its duration is few hours.

Spermatid: is small, spherical cell with nucleus and cytoplasm and which found towards
seminiferous tubules. This nonmotile cell transformed into motile cell sperm. This change is
called as Spermeogenesis. It takes place in 4 phases.
1. Golgi phase: Golgi granules are found above the nucleus in spermatids are called as
Proacrosomal granules. These granules joined to form single acrosomal granule.
2. Cap phase: these acrosomal granules start covering nucleus like cap
3. Acrosomal phase:nucleus start becoming elongated and cap become longer and covers
more than half of the nucleus.
4. Maturation phase: nucleus become very much elongated covered by acrosomal cap
and cytoplasm surrounding nucleus occupies position behind the nucleus to form
tail.sphericle nonmotile spermatid transferred into very long motile spermatozoa.
Tail divided into 3 parts:
a.Principal piece: has centriole with mitochondria on either side.
b.Middle piece: no mitochondria, only thick fibrous sheath.
c.End piece: only centriole.

Portion of residual body remain which is the remnant of cytoplasm and not capable of
fertilization.
These takes place in epididymis and collection of sperm from tail of epididymis.
Spermatogenesis takes place in only matured animals which attains maturity.
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OOGENESIS: It is the process of development of ovum from primordial germ cell.

Primordial germ cell migrates to the developing ovary to become oogonia in foetal stage.
Oogonium divides into primary oocyte(Oocytogenesis). Each primary oocyte undergoes 1st
meiotic division.

Primardial germ cell

↓
Oogonium(foetal stage)
↓ Mitosis
Primary oocyte (at birth)
↓ 1st meiotic division
Diplotene stage(stops)
↓ Ovulation
Secondary oocyte. +1st Polar body(contain only nucleus, no cytoplasm)
↓ 2nd Meiotic division
Ootid + 2nd Polar body
↓
Ovum

1st meiotic division: Prophase-leptotene, zygotene, pachytene, diplotene, diakinesis . At birth

primary oocyte comes upto diplotene stage of 1st meiotic division called Dictyate
stage/resting stage. This will last till the animal reaches puberty. Then metaphase, anapjhase
and telophase phases occurs.

At birth, ovary has cortex and medulla. Cortex has follicles and medulla contains blood
vessels and nerves.
Follicles in different stages of development:1.Primary follicle: primary oocyte covered by
single layer of flat cells.
2.Secondary follicle: primary oocyte covered by more than 2 layers of follicle cells.
3.Tertiary follicle:pri. Oocyte surrounded by follicle cell and theres is antrum
4.Matured follicle: graffian follicle found at one pole to come out.
All these follicles contain primary oocyte. Primary oocyte surrounded by single layer of
squamous epithelial cell is Primordial follicle.→Primary→Secondary→Tertiary→Matured
follicle.
Dictyate stage remains dormant in diplotene stage of 1st meiotic division and further
changes occur after ovulation. After ovulation, it pass through meta,ana and telophase. And
1st meiotic division is complete. 2nd meiotic division is completed only after fertilization.
In all animals except in dog and horse, both 1st and 2nd meiotic division is completed only at
or after fertilization.

OVULATION: is the process of release of ovum from the follicle (graffian). Ovulation can
takes place anywhere on the surface of ovary. The point at which ovulation takes place is
calles as Stigma. In mare sigma is confined to a depression as Ovulation fossa.
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Ovulation is of 2 types:
a. Spontaneous ovulation: is one where it doesnot require any exogenous agent to
ovulation to takes place. Eg. All most all domestic animals including human.(ie. It
occur due to harmone like estrogen, leutinizing hormone)
b. Induced ovulation: which require exogenous (extraneous) agent like
coitus/copulation to bring about ovulation. Eg. Rabbit, Ferret, Mink, Cat, Fox, Camel.

Factors which bring about ovulation:

1. Pressure due to large amount of antrum fluid (liquour folliculi), which make wall of
follicle thin and cause ovulation.
2. Weakening of wall of follicle due to some lytic agent like prostaglandin which brings
about ischemia, weakening of wall of follicle and rupture of follicle and release of
ovum.
3. Contractile activity of smooth muscle in the ovarian stroma results in ovulation.
[In sheep, not too much of liquor folliculi, but follicle is flabby so 1st theory is true. In ant
eater, no antrum in follicle but still ovulation takes place may be due to hormones.]

Hormonal control of ovulation:

a. Pituitary (anterior) releases the hormone FSH (Follicle Stimulating Hormone-
gonadotropic H.) responsible for development of the follicle.
b. Once follicle become mature follicle, Oestrogen is secreted which act as feed back on
pituitary so FSH secretion stops.
c. Pituitary start secreting LH (Leutinizing Hormone) which bring about ovulation.
Once ovum is released, rest of the follicle become Corpus luteum.
d. Corpus luteum secretes Progesteron which is necessary for maintenance of
pregnancy if fertilization occurs. If no fertilization, CL get regress, then next cycles
starts.

In Spontaneous ovulation:
Oestrogen

↑
Pituitary gland→FSH→Follicular development→Mature follicle
↓ ↓
LH → Ovulation → Corpus luteum
Progesteron if pregnancy
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In Induced Ovulation:
Oestrogen

↑
Pituitary gland→FSH→Follicular development→Mature follicle
↓ ↓
Coitus(oestrus/heat) → LH → Ovulation → Corpus luteum
Progesteron if pregnancy

Same as spontaneous ovulation but during release of LH, it requires copulation/coitus

during which there is stimulation of vaginal mucosa for nerve impulse is spinal nerves. Then
information goes from spinalcord to brain to hypothalamus(pituitary) to release LH which is
necessary for ovulation.

Note: there are 2 million oocyte in women is found in ovary during life time. Out of these
only 400 reach maturity. Rest of follicle get regressed i.e. Atretic follicles. Ie at birth
3,70,000 ovum, puberty-1,90,000 ovum and maturity-400.

FERTILIZATION:
It is defined as the union of male(spermatozoa) and female(ovum) gamets to produce
Zygote(single cell).
Fertilization takes place by penetration of spermatozoa into ovum. Fertilization takes
place at 1/3rd of oviduct. Spermatozoa are motile cell move by lashing movement of tail.
Sperms are deposited in female genital tract is called as Insemination. It may be natural or
artificial. Release of sperm from male genital tract is called Spermiation. Sperm speed in
female genital tract is 5mm/sec in bitch, cow 5mm/min. semen is deposited in vagina where
pH is 4.3 i.e. acidic. Acidity in the vaginal fluid is counteracted by secretion of vesicular
gland, prostate and ampulla i.e. seminal fluid. It brings to neutral which act as buffering
agent.
Before sperm get capacity (by biochemical and physiological modification), sperm
undergo changes in female genital tract called capacitation. Capacity is the ability of sperm to
fertilize the ovum under the influence of fluid in female genital tract.
These fluids brings about changes in sperm-
a. Breaking of acrosomal cap (which contains outer and inner acrosomal membrane)-
first breakdown of outer acrosomal membrane and release of hyaluronidase enzyme
(which is b/w outer and inner acrosomal membrane). Hyaluronidase enzyme is also
called as spreading factor.
b. Certain chemical substance present in ovum which attract sperm is called as Fertilizin.
Fertilizin is released by ovum by mature eggs which causes attraction of sperm
towards ovum. It is glycoprotein (33lakh mol.wt.)
c. Antifertilizin is secreted by Spermatozoa, 10,000 mol.wt. these two substance join
and form a reaction similar to antigen-antibody reaction and cause agglutination
which takes place only when they are of same species. And it also prevents
polyspermy.
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Mechanism of Fertilization: ovum is covered by 3 types of structure

a.Plasma membrane or vitelline membrane
b.Zona pellucid which is highly refractile layer
c.Layer of columnar cell ie.Corona radiata.

Fertilization takes place in 3 phases:

1. Penetration of Corona radiata: Corona radiata cell attach to one another by
cementing substance called Hyaluronic acid which is a mucopolysaccharide like
chondratin SO4 of cartilage. Cementing substance prevent entry of sperm. So it has to
dissolve, which is facilitated by hyaluronidase enzyme of sperm and release disperse
corona radiate cell and make enter of sperm.
2. Penetration of Zona pellucid: ZP second barrier which protect female gamete which
is penetrated by sperm with enzyme called Zona lysine or Acrosin (release from
inner membrane of acrosome). Once spem touches ZP, it becomes firmly attached and
penetrate rapidly by protein enzyme like zonalysin or acrosin. Thus penetration of
zona pellucida is called as Zona reaction./Acrosomal reaction. Specific receptor site
present in ovum lost for spermatozoa once head of spermatozoa enter vitaline
membrane. By zona reaction, further entry of other sperms is prevented i.e.
polyspermy is prevented after entry of one sperm.
3. Fusion of oocyte and sperm cell membranes: sperm touches the vitalline or plasma
membrane of ovum. So fusion of cell membrane. Under E/M, in some animals, they
arises cone shape structure called Fertilization cone as soon as sperm touches
membrane. Then cone is withdrawn, during which it drags sperm into cytoplasm of
ovum. Sometime, plasma membrane get completely surrounded and then draws head
of sperm into ovum.

In many animals, below plasma membrane, there are cortical granules, when sperm
head enter cytoplasm of ovum, there is development of cortical granules which block the
ruptured entry point and prevent entry of other sperms so polyspermy is prevented.- Cortical
reaction.ie. Vitelline block. (And there is some change in zona pellucida after its structure
and composition through removal of specific receptor site of spermatozoa is zonar reaction).

Once head of sperm enters ovum(contained male pronucleus and tail is cast off), there
is 2nd meiotic division of Oogenesis and is completed. There is fusion of both male and
female pronuclei to form zygote..

Results of Fertilization:
1. Chromosomal number is restored.- each gamet will have haploid number and by
fertilization it remain
2. Sex of embryo is established. xx-female and xy-male but in birds xy female and xx
male.
3. Initiation of 2nd meiotic division and completion of it.
4. Metabolic activation of zygote to undergo cleavage division.

Parthinogenesis: by activation of egg, cleavage division is induced even without a sperm for
fertilization by artificial means is parthinogenesis. i.e. development of embryo from ovum
that activated by means other than sperm. Eg. Hot fine needle on ovum cause cleavage, sharp
pointed needle dipped in blood in amphibian, fishes. In parthinogenesis upto 7 stage, but half
chromosome number it occur naturally in insects, lizards and lower animals upto frog. In
mammals, parthnogenetic individuals are all genetically females.
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CLEAVAGE: called as Segmentation.

It is the process of division of zygote found after fertilization. Zygote is 80-120μm, largest
mammalian cell. Zygote is single cell which is divided into multicellular organism. Segment
because daughter cell become smaller. Cleavage division is always mitotic. Cleavage involve
both nucleus and cytoplasm division. Division of nucleus is Karyokinesis and division of
cytoplasm is Cytokinesis.

In vertebrates division depends on amount and distribution of yolk material present in egg.
1.Isolecithal-have small amount of yolk distributed uniformly and nucleus is in centre.
eg. Mammals, amphioxus, primitive fishes.
2. Microlecithal-small amount of yolk
3.Megalecithal
4.Telolecithal
In 3 and 4th – cell contains large amount of yolk and pushing nucleus to one side/pole.
In Telolecithal eggs, 2 types-
a. Highly telolecithal egg: large amount of yolk pushing nucleus to one side eg. Birds,
reptails
b. Moderately telolecithal- comparatively smaller amount of yolk. eg. Amphibian.
c. Here at yolk no division occurs and it occurs only at nucleus.

In isolecithal, egg (zygote) divide into equal parts called Holoblastic cleavage. Where as in
Telolecithal, divide equal or unequal cleavage and is Meroblastic cleavage(division only at
animal pole and resulting cells are called as Blastomeres.).eg. meroblastic equal cleavage-
amphioxus.
So in mammals- isolecithal holoblastic equal
In birds- highly telolecithal meroblastic unequal
Amphibia- moderately telolecithal meroblastic equal.

In eggs, where the nucleus is present is called Animal pole and where yolk and cytoplasm
present called as Vegetal pole.

Cleavage division in mammals(isolecithal):

First division is always vertical resulting in 2 daughter cells.
Second division is also vertical resulting in 4 daughter cells.
Third cleavage is horizontal resulting in 8 daughter cells.
Fourth cleavage results in 16 cell in dog after 8 cell results in 12 cells.
In 16th cells stage, daughter cells are arranged in the form of mulberry fruit and
surrounded by zona pellucid called as Morula stage. This occurs in pig by 4 days, horse 4-5
days and cow 4-6 days.
The morula stage egg start migrating from the place of fertilization(upper 1/3 rd of
oviduct) to the horn of uterus.
After 16 cell stage is 32 cell stage. Here cells are rearranged insuch a way that a layer
of cells to periphery(tropoblast) and at one pole there is clump of cells called Inner cell mass
or Embryoblast which give rise to embryo. The cavity formed is called as Blastocele. Whole
structure is surrounded by zona pellucid and called as Blastula.(and process is called
Blastocyst).
Tropoblast cell lining periphery give rise to extra embryonic membrane.

Fate of Zona pellucid: in mammals, it begins to degenerate in stage of blastula and resulting
blastula to environment of uterus and facilitate implantation to occur. In horse, zona pellucid
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is replaced by a capsule of CHO which is produced by tropoblast cells. Function is could be

protective and capsule remains for 21 days of gestation. So in equine implantation is always
late because of capsule development.
The cleavage division utilizes lot of energy till about 16 cell stage from yolk which is not
sufficient further so implantation occur.
Blastula is in uterus, get nutrition from Uterine milk which is formed by secretion of
uterine gland and some sloughing of cells of epitherlium. This type of nutrition t o blastula is
called as Histotropic nutrition.which is also for few days so for permanent nutrition
implantation takes place ie blastula implanted to uterine mucosa.

Implantation is defined as the embryo getting attached or burrowing for its nutritional
requirement to its maternal tissue. In domestic animals implantation is superficial without
burrowing deep called as Nidation. True implantation is only in primates and rodents where
implantation upto endometrium of uterus.
Implantation always occurs towards the inner cell mass.

3 types of implantation:
a. Superficial-in ungulates and carnivores, blastula gets attached to epithelial cells
superficially without causing damage to uterine mucosa.
b. Interstitial – uterus has number of folds. Blastula gets burrowed in one of these uterus
folds eg. Man, g.pig, primates.
c. Deep or eccentric- blastula get implanted deeply into mucosa giving eccentric
position causing much damage to uterine mucosa eg. Rodents, rat, rabbit.
In b and c,have a layer of uterine tissue will be covering the blastula eroded deeply by
causing injury to uterus and this type of reaction is called as Decidual reaction.which type
gives more bleeding during parturition.

Implantation may be towards or away from mesometrium. If towards(towards broad lig. Of

uterus) –mesometrial implantation and if away-antimesometrial implantation.
During implantation, layer of epithelial cells of uterus covering blastula(towards upper part)
is called Decidua capsularis. And opposite side(towards inner cell mass) is Decidua basalis
and where implantation is abscent is Decidua paritalis.

Cleavage in Birds: In Telolecithal eggs: meroblastic and discoidal

Here egg has large amount of yolk at the bottom doesnot undergo division. Cleavage takes
place only in animal pole where nucleus present.
Division takes place by formation of Furrow-cleavage furrow.
1st division is vertical resulting in 2 blastomeres.
2nd division takes place right angle to 1st or perperndicular to 1st reluting in 4 blastomeres.
3rd-is b/w 2nd and takes place paralled to 1st on either side resulting in 8 blastomeres and
furrows with clumping of cells at centre.
4th- large number of cells at top with cleavage furrow. It appears like a disc of cell with yolk
underlying and is called as Blastodisc. And cavity called Blastocele.
Birds laid egg at this blastodisc stage.

GASTRULATION: Germ layer formation: it is defined as process of formation of primitive

germ layers like mesoderm, ectoderm and endoderm. Embryo with three germ layer is called
as Gastrula stage.-which contain cavity called blastocele, inner cell mass and tropoblast.
Here single layered blastula converts into trilaminar stage.
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Gastrulation steps:
1. Delamination of cells from inner cell mass into blastocele cavity. Single layer of cells
arrive inside the tropoblast is called Hypoblast cells. These cells later become
primitive endoderm.
2. When hypoblast are being formed, the tropoblast cells immediately below ICM
become tall columnar cells called as Epiblast cells. These cells gives disc like
appearance called as Embryonic disc or embryonic shield. These epiblast cells give
rise to Primitive or primary ectoderm.
3. B/w epiblast and hypoblast an extracellular basement membrane appears. Tropoblast
and epiblast are separated each other to form fluid filled cavity called as Primitive
amniotic cavity.
4. So b/w primary yolk sac and primary/primitive amniotic cavity a bilaminar plate is
formed which give development of embryo ie. “Bilaminar germ disc”.

Before stage of germ cells are established, there is rapid elongation of the embryo soon
after the zona pellucida is lost. Rate of elongation is about 1cm/hr in pig. Tropoblast cells
become long and tubular and reaches the length of about 35cm in ruminants and 1mt in pig. It
takes place from 9-16 days in pig and 12-14 days in bovine and in horse, not much of
elongation.

5. Development of Primitive streak:

By the time the hypoblast cells lining the blastocoele cavity to form the endoderm, the
epiblast cells in the embryonic disc region, migrate towards a position along the diameter of
the disc ie. towards the centre. This migration and proliferation of epiblast cells results in
formation of two parallel ridges with a groove in the middle. These are called as Primitive
ridges and primitive groove. These structure is called as Primitive streak. Development of
primitive streak is important event to form third germ layer Mesoderm.
The development of primitive streak establishes the longitudinal axis of embryo.
Primitive streak always start from tail end and go towards head end. It establish a bilateral
symmetry.
From the epiblast cells, another layer of cells migrate by delamination and occupies a
position b/w hypoblast and tropoblast cells and these give rise to Mesoderm.

Development of Notochord:
it is a rod shaped aggregation of cells that extends entire length of embryo. Primitive
streak does not reach complete length of embryo and it may go upto 2/3rd of embryo length
then it stops. At where it stops there is a depression called primitive pit. Infront of P.pit there
is clump of cells these group of cells are called as Hensen’s node/ Primitive knot(i.e
Cephalic end of primitive streak swollen by proliferation of cell called Primitive
knot/hensen’s node). From this node cells start migrating towards head end b/w epiblast and
hypoblast. So it looks like a plate of cells and this plate of cells are called as Notochordal
plate which is a broad sheath like structure. Later notochordal plate starts folding and gives
rise to a row of cells. This chord of cells from hensen’s node towards head end is called
Notochord.
Importance: in adult, notochord doesnot gives any embryonic structure but it has a great
developmental significance.
1. It established Cranio-caudal axis of the embryo
 15

2. It serves as an inductor for the development of central nervous system this process is
called as Neurilation.(i.e. if remove portion of notochord, nervous system will not
develop or if you implant notochord to other embryo, new nerve will develops)
3. It is an organizing center for the development of vertebral column and cranial
bones(like occipital, sphenoid, ehamoid ie floor of cranial cavity).

“As the length of notochord increases, primitive streak become shorter” time may come that
primitive streak lost with only notochord.

Note: B/w epiblast and hypoblast, a layer of cell give rise to mesoderm is development of
Gastrula in mammals.

Gastrulation in other vertebrates:

Isolecithal-eg. Amphioxus- gastrulation takes place by invagination of one portion of the
Blastula so two layer of cells forming ie. hypoblast and epiblast.
Frog: moderately telolecithal
Birds: highly telolecithal eggs: from tropoblast cells, another layer of cells start developing
which similar to hypoblast. b/w epiblast, tropoblast become hypoblast. Not it is 2 layered
structure. Epiblast cells in the middle shows a clump of cells called Notochord. From
notochord cells, some of cells delaminate and occupy position b/w epiblast and hypoblast.
And give rise to mesoderm. Development of primitive streak and notochord are same as
mammals.

Development of extra embryonic membranes:

These are membranes which surround the developing embryo and doesnot give rise to any
part of the embryo. These EEm are structure which develop mainly for protection and
nutrition of the embryo.
These are 4 in number;
1. Yolk sac
2. Amnion
3. Allontois
4. Chorion
Mesoderm is a single layered cell lining on either side of notochord. Later mesoderm cells are
divided into 3 parts.
a. Paraxial mesoderm: is first layer of cell by the side of notochord
b. Intermediate mesoderm: single layer of cells b/w a and c
c. Lateral mesoderm: lateral most region, has 2 parts
1. Somatic mesoderm: joins with ectoderm to form Somato pleura.
2. Splanchnic mesoderm: it joins with the endoderm to form Splanchnopleura.
(splanchnic mesoderm which is lining GIT, respiratory and excretory organs.)

Between somatic and splanchnic mesoderm, there is a space called Coelom or body
cavity.

Yolk sac:
It is first extra embryonic membrane to develop. Well develop in birds and reptails
where large amount of yolk is present. Eggs with little yolk in mammals, yolk sac develop for
short time and then it regresses. Yolk sac is always continuous with the gut tube. Endoderm
cells give rise to gut tube (endodermal cells folded in the gut tube). Connection between gut
tube and yolk sac is called as Yolk stalk. It is always line by endoderm inside, splanchnic
 16

mesoderm outside. Yolk sac gives rise to primordial germ cells. Yolk in yolk sac provide
nutrition to embryo in birds and reptiles where as placenta doesnot develop. As embryo
develops bigger, size of yolk sac become smaller. (Newly hatched chick has small amount of
yolk which is sufficient for one day. So give only water first day).
In chick, at 48hrs of incubation, we can see the blood islant at proximal part of splanchnic
mesoderm. This vascular area of yolk sac is called as Area Vasculosa. In distal part no blood
vessels called as Area vitallina.
Function of yolk sac: digestive enzyme secreted by endoderm cells into yolk and partially
digested food absorbed into cell.

AMNION: means ‘Vessel’ like. It develops by two methods

a. By Folding: “Plect amnion”- usually seen in mammals and birds except in rodents
and primates. Fold starts developing in embryo involving mainly ectoderm and
somatic mesoderm. Three types of folds develops
1. At head region-Head fold
2. At tail region-Tail fold
3. Lateral fold-on either side.
All these folds meet at a one point. After folding they again starts covering embryo
enclosing a large cavity called as Amniotic cavity. Membrane enclosing amnion
cavity is called as Amnion which is made up of ectoderm inside and somatic
mesoderm outside. Amniotic cavity contain amniotic fluid –watery type, which acts
as a bactericidal, act as private aquarium for developing embryo. It prevents injury to
embryo and external shock. Usually called as Water bag. During parturition, amnion
comes out first and helps in dilatation of cervix.
Amount of amniotic fluid is 3.5lit may increase upto 6.5lit in different animals.
Amniotic fluid contain Na, Pot.chloride and fructose. Too much of amniotic fluid
which press on developing embryo is called as Hydramnios.
By amniocentesis, sex of the animal can be detected by looking for x or y
chromosome.

b. By Cavitation: “Schiz amnion”(humans) – tropoblast above inner cell mass or

epiblast cell become differentiated into 2 zones:
1. Cytotropoblast: cell boundaries are distinct
2. Syncitio tropoblast: cells are not clearly demarked, only nucleus is seen. Small cavity
start appearing in syncitio tropoblast and these cells become amnioblast. Cavities fuse
and become larger and start surrounding embryo completely.

ALLONTOIS: “Sausage shape”

During amnion development by folding, it also involves endoderm cells which forms
a tube like structure with head, tail and lateral fold called as Gut tube which later give rise to
digestive tube. It has 3 parts, namely
 Fore gut
 Mid gut-continuous with yolk sac
 Hind gut-from here a diverticulum develop involving endoderm and splanchnic
mesoderm and which become larger and completely surrounds the embryo and called
as Allontois. It also surrounded by splanchnic pleura. It develop by 25 days in the
cow. Membrane lining allontois secete Allontoic fluid which acts as an excretory
organ of embryo (and later become urinary bladder). Secretory activity of allontois
maintain Osmotic pressure of foetal plasma. 4-12 lit in cow and 70cc in sheep. Blood
vessels called allontoic vessels develop later.
 17

Note: some animal contain large bodies ie. faecal matter excreted by embryo called
Hippomanes which accumulate in allontois. Commonly seen in Horse and in pig and
ruminants. It also called as aphrodesiae. Rich in gonadotropic harmones.

CHORION: during development of amnion, head fold and tail fold join so there are 2 layeres
formed.
a. inner layer which give rise to amnion and
b. outer layer which give rise to chorion which completely surrounds the embryo including
amnion, allontois and yolk sac. Chorion is lined by somato pleura (ie. ectoderm outside and
somatic pleura (mesoderm) inside and amnion – somatic mesoderm outside and ectoderm
inside).

All these process are taking place in uterus. Chorion come in contact with lining of uterus.
Villi start developing from chorion called as Chorionic villi. These villi burrow deeply into
endometrium of uterus which give rise to Placenta.

3 types of villi:
1. Primary villi: villi having only ectodermal layer.
2. Secondary villi: ectoderm with connective tissue core of somatic mesoderm
3. Tertiary villi: ectoderm + somatic mesoderm core and blood vessels.
Villi may be separate or joined with one another called as Labyrinthine ie connection and
Non labyrinthine ie no connection.
Villi in chorion that area is called as Chorion fondosum. And area absence of villi is
Chorion leave.
Usually Allontois+chorion join called as Chorioallontois. Chorion and allontoic
membrane attached to maternal tissue called Chorio allontoic placenta.

PLACENTA AND PLACENTATION:

Placenta is defined as area of intimate contact b/w maternal and foetal tissue for physiological
exchange. Placenta means circular like disc or cake. It is coined by Realdus Columbus.
Placenta is seen in mammals. Mammals classified into 3 types
1. Prototheria: animals which lay eggs. Eg.Platypus, dolphin and whale
2. Metatheria: pouched mammals, they give rise to young ones which are worm like
which is carried in pouches. Eg.kangaroo, bandicoots.
3. Eutheria: true placental mammals give birth to young one. All domestic animals
including man.

Classification of placenta:
3 main principles in classification are
I.a. Vascular origin
b. Foetal-maternal relationship
c.Morphology

a. Vascular origin:
1.Yolksac placenta or omphaloid: it develops in early stage but size of yolk sac
gradually get reduced so act for short time. In rabbits, yolk sac placenta participate in
passive immunization by absorbing immunoglobulins. It is temporary.
2. choriovitelline: yolk sac comes in contact with chorion, it is temporary and it is
replaced by chorioallontoic placenta.
 18

3.Allanto-chorionic/chorio-allontoic: permanent one. Allontoic membrane come in

contact with chorionic membrane.

b. Foetal-maternal relationship: degree to which foetal membranes anchor to

endometrium.
(Decidua: modified form of endometrium just after implantation.)
A.1. Deciduate: foetus burrow deeply in endometrium and causing tissue damage to
endometrium are deciduates. There is more bleeding during parturition. Eg: rodents,
primates.
2.Non deciduate: superficial attachment, not much damage, not much bleeding.

In decidual placenta, decidual cell are found which are enlarge, polyhedral or rounded
derived from fibroblast. Eg. In carnivore placenta.

B. depending upon the type of villi.:

1. Labyrinthine: these are deciduate type. Where villi are interconnected.
2. Non labyrinthine: non deciduate type ie. no connection

c. Morphological classification:
1.Depending on shape: 4 types

a. Diffused type: villi are distributed uniformly throughout chorion without any gaps or
microcotyledinary. Eg. Sow, mare, camel.

b. Cotyledenary type: eg. Ruminants, deer. Villi confined to specialized area called
placentoms or cotyledons also called Multiplex. Ie. cotyledons in foetus, caruncles in
maternal side.
Cow-button shape, sheep-depression in the middle

c. Zonary type: villi are confined to belt like zone ie. transversly oriented ring like. Eg.
Carnivores ie dog, cat.

d.Discoid type: villi placed on large flattened area in the form of a disc. Eg.primates,
rodents, man
monkey- bidiscoid.

2.Internal structure:
a.producing folded(plicae, rugae, eg. Sow)
b.Villous type: mare, ruminants (chorionic villi)
c.Labyrinthine type: intercommunication of allanto-chorion eg. Carnivores, rodents.

3.Histological classification/ number of layers of interhemal membrane:

Foetal and maternal blood is separated by endothelium, CT, epithelium v/s epithelium,
CT and endothelium.
a.Epitheliochorial: all 6 layers are intact. Ie. all 3 layers of maternal side persist.
Eg.diffused type, in mare, sow, camel.
 19

b.Syndesmochorial: uterine epithelium is reduced and other 2 layers are intact. Ie

epithelium is lost and CT come in contact with chorion. Eg. Ruminants, deer.

c.Endotheliochorion: endothelium of capillaries of maternal uterus in contact with

chorion here uterine epithelium and CT are absent. Eg. Carnivores.

d.Haemochorion: there is damage to endothelium of blood capillary. So blood is in direct

contact with chorion. Ie. all 3 layers of maternal side is absent so tropoblast freely
exposed to maternal blood. Eg. Primates, rodents.
This classification is given by Gosser, 1902.
Mossman, 1932, In Logomorph(rabbits): he says Haemoendothelial type: where maternal
blood come in contact with endothelium of chorion.
Owers, 1960, acc to him, Indian musk shrew has Endothelioendothelial type.

Sow-chorio allontoic, non decidual, non labyrinthine, diffuse, epitheliochorion

Dog- chorioallontoic, non decidual, labyrinthine, zonary, endotheliochorion.

Functions of Placenta:
a. Provides nutrition: haemotropic nutrition ie b/w blood of foetus and maternal
(embryotropic-a.histotropic-uterine gland secretion b. hemotropic-maternal blood)
b. Exchange of gasses like CO2, O2, and electrolytes by simple diffusion. 20-30cc of
o2/min exchange occurs b/w foetal and maternal blood.
c. Exchange of nutrients, aminoacid, CHO, FFA, Vitamine etc.
d. Hormone production: like progesterone(for pregnancy maintenance), oestrogen(for
parturition) and some gonadotropic hormone like FSH,LH and in mare eCG or
PMSG
e. Also produce Relaxin at the time of parturition.
f. In transmission of maternal antibody by pinocytosis or phagocytosis. Immunoglobins
provide passive immunity against virus and bacterial diseases.
g. Secretes lactogen which is seen in human and other animals.

Abnormalitis in placenta:
1. Hydroamnion
2. Hydroallontois
3. Retention of placenta: placenta retain without excretion after parturition

Twins and Twinning:

Twins is more than one birth at a time in uniparous animals like cow, horse, man.
There are 2 types of twins:
1.Monozygotic twin: one ovum is released and fertilized by single sperm give single zygote
which divided into two and develop independently. It also called Identical twins. They are
phenotypically and genotypically similar. Same sex ie xx(female) or xy(male).

2.Dizygotic twin: are called Fraternal twins. Here two ovum released and fertilize by 2
separate sperm and 2 zygote formed and develop independently. Sex may be same or
different. Phenotypically and genotypically different.

Freemartin: it is a female co-twin of a male. It is a dizygotic twin where there is male and
female. Here female is sterile and have hypoplastic genital tract. And male is normal and this
female twin is called as freemartin. There is chromal defect which is xx/xy chimarism.
 20

Thought to be brought about by early mixture of blood of two sexes. There is inhibition of
development of female genitalia. Evident that HY antigen from invading xy cell influences
cells in developing ovary to produce Androgen which inhibiting female development or
Antimullerian harmone may be allowed to function. All these influence ovary to behave
like a male gonad. Freemartin usually seen in cattle, goat, sheep and pig.

Teratology:
Study of monsters or abnormal development is teratology. There are three developmental
stages.
a. Stage of predifferentiation: any agent applied during this stage, it has either one or
two effect. It may be lethal to embryo or no effect at all.
b. Stage of early differentiation: is the stage where it is more susceptible for
malformation.
c. Stage of advanced morphogenesis: here there is increased effect of teratogen, increase
with age of embryo.

Different type of anamolies brings about altered morphogenesis events like:

1. Failure in development or agenesis: particular part or organ will not develop. Eg.
Anaophthalmia-absence of eye
2. Development of excesses-excess development of particular part. Eg. Supernumerary
teat, extra finger or limbs
3. Incomplete development-like aplasia or hyperplasia. Aplasia is retardment in
development. There may be incomplete growth like dwarphism. Sex organs are
undevelop, failure to fuse, like cleft palate, septal defect in heart, defect in diaphragm.
4. Incomplete migration: like ectopic cardis, cryptorchidism(failure of testis to descend)
5. Persistent of embryonic structure like persistent of pupillary membrane, persistent of
anal membrane, paternt ductus arteriosus.
6. Malposition of embryonic structures. Eg. Organs in left side found in right side.

Etiological agent bring about malformation classified under:

1. Genetic factor:
a. Abnormal genes
b. Chromosomal abnormalities: autosomal abnormality leadt to mongolism. Where
as sex chromosomal abnormality leadt to monosomy and trisomy.
Ie. Aneuploidy/monosomy: loss in one of chromosome-Turner’s syndrome-ie.
‘XO’ only one x. other x or y is not there.
Polyploidy/Trisomy-excess of one of chromosome-‘XXY’ ie Klinefelter’s
syndrome. It is reported in dog, cat, boar, bull, ram.
2. Environmental factors: factors like
a. Radiation or physical factors eg. X-rays, √ rays.
b. Chemical agents: carbonic anhydrase, some of drugs which inhibit DNA
metabolism, plants like veretran species.
c. Infectious diseases.
d. Nutritional deficiency
e. Harmones like ICTH.
f. Many agents like nicotin, tetracycline.
 21

SPECIAL EMBRYOLOGY/ORGANOGENESIS
MESODERMAL DERIVATIVES

DEVELOPMENT OF SKELETAL SYSTEM:

Mesoderm divided into 3 parts:

a. Paraxial mesoderm: give rise to veterbral column, skull, somatic muscles, dermis of
skin
b. Intermediate mesoderm: heart, circulatory system, urogenital system
c. Lateral mesoderm: connective tissue, muscles lining the GIT, repiratory tract and
urogenital tract.

Paraxial mesoderm: found on eitherside of notochord. It arrange in the form of blocks.

These blocks are called as Somites.ie. blocks of paraxial mesoderm located on either side of
notochord. Somite consist of 3 parts.
1. Sclerotome: ventromedial position-gives vertebral column
2. Dermatome:dorsolateral position-gives dermis of skin
3. Myotome: dorsomedial position-gives skeletal muscle.
Cavity inside somite is called as Myocoel.

VERTEBRAL COLUMN:

No. of sclerotome= no. of vertebrae.

Ist stage/early stage: sclertotomes are in the form of segmentally arranged. In the prevertebral
stage, each sclerotome is divided into two parts.
a. Cranial loose portion
b. Caudal dense portion
On either side of thses is myotome and dermatome. In between sclerotomal block is
Intersegmental artery. In middle of each is Intersegmental nerve.

IInd stage/late stage: caudal dense portion of one joins with cranial loose portion of next one
ie. rearrangement. When these are joining, myotome attached to two adjacent sclerotome.
And also rearrangement of intersegmental artery to the middle which was b/w sclerotome and
nerve become intersegmental ie b/w the blocks.

So in center notochord, and dense portion form the body of vertebrae, and loose portion
forms arches and process. These encloses Neural canal.

Somite appears first 20hrs of incubation in chick embryo. Every 1hr there is extra somite.
20+2=22hrs→2 somite so 30hrs incubation→10 pairs of somites. So age can be detected.
 22

In pig, 14 days→1st somite. In dog and horse,16 days→ 1st somite. In birds 30-40hrs, we can
access the age, after this not caluculated.
Another type of aging is by Crown Rump Length →used in mammals.

Note: Measurement of foetal body→ Crown Rump Length: CRL: measured as a curved line
in centimeters with a calibrated inelastic thread along the vertebral column between most
cranial part of frontal bone to the rump at ischiatic tuberosity which is designated as Crown
Rump Length.
Approximate age of the fetuses was calculated by using formula which is given by Soliman,
1975.
Y=28.66+4.496X (CRL<20cm)
Y=73.544+2.256X(CRL>20cm)

Where as Y→is the age in days, X→is the CRL in centimeters

Atlas/1st Cervical vertebrae-has no body and contain arch and process ie ring and wing.(only
loose portion) because dense portion joint with 2nd Cervical vertebrae. Ie. odontoid process
joint to C2. And last coccygeal veterbrae only bodies because of dense portion.

In b/w body of vertebrae, the notochordal cells occupying place which give rise to
Intervertebral disc. Each intervertbral disa has two parts namely,
a. Peripheral-annulus fibrosus: derived from notochord
b. Central-nucleus pulposus-derived from mesenchymal cells surrounding notochord.

In cervical vertebrae, body, arech and transverse process. Transverse process is divided into
two parts ie dorsal part and ventral part. In thoracic portion,Ventral part form ribs or costal
process. In rib, head and tubercle and in b/w gap, give rise to foramen transversarium. This
arrangement found in all cervical vertebrae except C7. In lumbar region, costal process gives
attachment to transverse process so it is long and in sacral region, it forms wing of sacrum.

Abnormalities during development of vertebral column:

Chondrodisplasia: there is inhibition of differentiation of annulus fibrosus of intervertebral

disc. This weak annulus fibrosus has increased tendency for rupture and protrusion of nucleus
pulposus.

Block vertebrae: most vertebrae are free but sometime fused. Eg. Thoracic and cervical
vertebrae may be fused.
 23

Hemivertebrae: there is a cuniform hemivertebrae. Ie one part is present and other part is not
there .

DVC: deviation in vertebral column: like

Torticolis-twisted neck.
Scoliosis- lateral deviation.
Kyphosis-dorsoventral deviation.
Lordosis-ventral deviation.

Incomplete formation of arch: here neural canal not completely develop so spinal cord is
exposed to outside, sometime protrude out of skin called Spina bifida.
Protrusion of spinal cord along with meninges. It may be protrude through out skin
and is covered by skin, called Spina occulta.

Development of Ribs: (develop from somites) ribs developed from costal process of
vertebrae. Costal process forms in cervical region as ventral division of transverse process, in
thoracic region they form into ribs. It forms cartilaginous model laid down in the form of
bone later by enchondral ossification. Part of it remains as cartilage called Costal cartilage.

STERNUM: arises in the form of pair of cartilaginous bars called as sterna bars/sterna plate,
appearing on ventrolateral aspect of the body. Later as the body walls get closes ventrally,
Fusion of cartilaginous bars takes place in cranio-caudal direction. So there is single
ossification centre for each sternabrae, last part remains in the form of xiphoid cartilage.(ie
enchondral ossification) To these, costal cartilage gets attached. It is osseocartilagenous
structure.
Anamolis: Bifid sternum/sternal cleft: incomplete fusion b/w sternal plate.

Development of Appendicular skeleton:

It develop first from mesenchyme cartilage.

Four types of bones: short, flat, long and sesamoid bones. Long bone give rise to radius,
humerus, femur. Short bones- carpals, tarsals,. Flat bones-scapula, hip bone.
Long and short bones develop by intra cartilaginous type of ossification. This bone develops
by local mesenchymal tissue and not from somites.
Appendage buds or limb buds- is outgrowth of mesodermal tissue covered by ectoderm.
Forelimb bud and hind limb bud.
Flat bones develop from intramembranous type of ossification and also from local
mesenchymal tissue. Sesamoid bones develop from tendons of muscles.
 24

JOINTS: develop from mesenchyme in between two developing bone may differentiate into
fibrous tissue or cartilage.

SKULL: developmentally skull consists of 2 parts. Namely

1. Neurocranium: has 2 part
a. Chondrocranium
b. Desmocranium/Dermocranium
2. Viscerocranium

Chondrocranium: is portion of skull which forms floor of cranial cavity. These are derived
from enchondral or intracartilagenous type of ossification. Infront of first pair of somite and
notochord, is Occipital somite. Infront of occipital somite, is 3 pairs or cartilages and 3
unpaired cartilages.
3 unpaired cartilages are:
a. Parachordal plate →give rise to basioccipital plate
b. Hypophysial cartilage/plate→give rise to body of sphenoid(basi/postsphenoid)
c. Trabaculie cranii→gives rise to body of presphenoid and perpendicular part of
ethamoid.
3 paired cartilages are:
a. Otic capsule/otic vesicle→give rise to petrous part of temporal. B/w basioccipital and
petrous temporal there is foramen lacerum for IX, X and XI cranial nerves.
b. Ale sphenoid(temporalis)→give rise to temporal wing of basisphenoid. It has foramen
ovale for mandibular nerve(part of trigenminal ie V)
c. Orbito sphenoid→give rise to orbital wing of sphenoid and it has foramen
orbitorotendum for III, IV, VI and Vth cranial nerves. In front of this is optic foramen
for IInd cranial nerve. All these form floor of cranium.

Desmocranium/Dermocranium: bones which form lateral wall and roof of cranium like
frontal, parietal, interparieta., supraoccipital and squamous temporal are develop by
Intramembranous ossification of local mesoderm(all flat bones).
Viscerocranium: develop from bronchial arches/pharyngeal arches/gill arches. Each arch is a
mesoderm.

1st arch: divided into 2 parts.(supplied by V cranial nerve)

a. Maxillary arch→dorsal part. It gives rise to bones of face like maxilla,nasal,
pterygoid, palatine, premaxilla, vomer and malar. All these develop by
intramembranus type.
b. Mandibular arch→ventral part. Mandible first develop as cartilage called as Meckel’s
cartilage (cartilage of ventral part of mandibular arch). Reminents of Meckel’s
cartilage are malleus and incus bone, sphenomandibular ligament. Mandibular arch
later laid down in the form of mandibular bone by intramembranous type.
 25

2nd arch/Hyoid arch→cartilage divelops is called Reicher’s cartilage which give rise to
stepes, bulla tympanica, stylohyoid, tympanohyoid, part of body of hyoid, ceratohyoid and
epihyoid. It supplied by VII cranial nerve.

3rd arch→give rise to lingual process and part of basihyoid and cartilages of larynx like
thyroid, cricoids and arytenoids. Supplied by IX cranial nerve.

4th ,5th, and 6th arch→ give rise to epiglottis and partly other cartilages. And thyrohyoid.
Supplied by X cranial nerve.

Anomalies in skeletal system:

In Limbs:
1.Absence of limbs→Amelia-complete absence of limbs
2.Syndactyla→some of the fingers are fused.
3.Polydactyla→more than 5 digits.

In sternum: cleft sternum-fusion between bars is abscent.

Skull:anamolism of fascial skeleton

Brachygnathia→there is a short upper jaw.
Prognognathism
Cyclopia-fusion of 2 orbits in the same single line.
Hydrocephalus-there is expansion of cranial cavity
Anencephaly-occipital bone fused with atlas.

MUSCULAR SYSTEM

It is necessary for locomotion of body, respiration, swallowing, catching the food and
transport of substances.
Skeletal muscle arises from Myotome of somites, bronchial arches or from local
mesenchyme. Precursors of myotome are myoblasts. These arises from paraxial mesodermal
cells. The process of development of muscle cells from myoblasts is called as
Myogenesis.(i.e. development of muscle).

Each muscle arise from myotome is divided into epaxial and hypaxial group.
Epaxial muscle(Epimere) are those which are supplied by dorsal branch of spinal nerves
and muscles above the transverse process. Their main function is extensors of the vertebrae.
 26

Hypaxial muscle(hypomere): are those supplied by ventral branch of spinal nerves and are
ventral to transverse process in position. Their main function is flexors of spine/vertebrae.

These muscles are segmentally arranged:

Axial group:
Epaxial muscles:
Long muscle group:
1. Ilio costalis-thoracic, lumbar part.
2. Longisimus-lumbar, thoracic and cervical
3. Spinalis et semispinalis- lumbar, thoracic and cervical
Short muscle group:
1. Interspinalis-inb/w spines. Develop in thoracic region
2. Intertransversaris-b/w transverse process.
3. Multifidi-b/w articular and transverse process.

Hypaxial muscles: is divided into 3 groups.

a. Dorsomedial group: called as Prevertebral group. In lumbar region- sublumbar
muscles like psoas major, psoas minor, iliacus. In cervical and thoracic region-longus
colli.
b. Lateral group: at lumbar/abdominal region- abdominis externus, abdominis internus
and transverse abdominis. In thoracic region- intercostalis externus, intercostalis
internus and transvers thoracis. In cervical region-trapezius, Rhomboideus,
sternocephalicus.
c. Ventral group: lumbar/abdomen-rectus abdominis, thoracic- rectus thoracis, cervical-
infra hyoid.

Muscles of Appendicular skeleton(Limbs):

Two theories:
1. Limbs develop from local mesoderm ie. from limb buds.
2. They develop from migration of myotomal blast(myoblast) of hypoxial group
muscles.
More evidence to say that they develop from myotome because of nerve supply by ventral
spinal nerves like brachial plexus and sacral plexus.
Muscles of limb are divided into dorsal group ie extensors and ventral group ie flexors.

Muscles of Pelvic region:

It is develop from local mesoderm. It has a dorsal division, lower sacral and coccygeal
region. Ie somites transformed into myotome and these myotome attached from sacral to
coccygeal to form pelvic diaphragm ie coccygeus, retractor ani and muscles of perineum ie
bulbocavernosus, ischiocavernosus, retractor penii etc.
 27

Diaphragm: innvervation is by 5th, 6th and 7th cervical spinal nerves. It is located in between
thoracic and abdomen but nerve supply is by cervical region. Ie. earlier it fromed b/w
thoracic and cervical region and later it migrate to last part of thoracic region but with same
nerve supply.

Muscles of eyeball: muscles arise from preotic somites. Infront of first pair of somite ie
occipital somite is otic then preotic somite. Preotic somites are 3 in number. They are
a. Premandibular: give rise to dorsal rectus, ventral rectus, medial rectus and inferior
obliques which are innervated by IIIrd cranial nerve.
b. Mandibular somite: which give rise to muscle superior obliques which is innervated
by IVth cranial nerve.
c. Hyoid somite: give rise to lateral rectus and retractor bulbi muscles which are
innervated by VIth cranial nerve.

Muscles of Head:
These are mainly derived from Bronchial arches.
1st arch:Mandibular arch: innervated by 5th cranial nerve and muscles derived are muscles of
mastication like massator, temporalis, medial and lateral ptergoideus, digastricus and
myelohyoideus.
2nd arch:hyoid arch: innervated by 7th cranial nerve and muscles derived are muscles of lip,
nostril, cheek, eyelid and ear. They are nasolabialis, caninus, buccinators, orbicularis oris,
orbicularis oculi, ear muscles. Ie muscles of facial expression.
3rd arch: innervated by 9th cranial nerve. Muscles of pharynx like cranial constrictor of
pharynx ie pterygopharyngeus, stylopharyngeus.
4th,5th and 6th arches: innervated by 10th cranial nerve. They innervate caudal constrictor of
pharynx like crico, thyro, hyopharyngeus and laryngeal muscles like cricothyroideus.

Muscles of tongue: develop from occipital somites. And are innervated by 12th cranial nerve.
Muscles are styloglossus, hyoglossus and genioglossus and intrinsic muscles of tongue.

Muscle innervated by 11th cranial nerve are sternocephalicus and trapezius. Spinal accessory
is considered as modified spinal nerve. In fishes and amphibian only 10 cranial nerves and
11th is modifies spinal nerves. These muscles derived from somites. Some evidence says they
derive from 5th and 6th bronchial arches.

Smooth muscles: derived from splanchnic mesoderm. Splanchnic mesoderm lining gut tube
ie digestive and respiratory tract are made of smooth muscle. Smooth muscle lining
urogenital system and blood vessels are derived from local mesoderm may be splanchnic or
somatic mesoderm.

Cardiac muscle: derived from splanchnic mesoderm that lined with primitive heart tube.
 28

CARDIOVASCULAR SYSTEM

It includes heart and blood vessels.

Vascular system: blood vessels appear by about 3rd week and provide nutritional requirement
of the embryo. It arises from splanchnic mesoderm. Cells of splanchnic mesoderm form
clusters of cells. These are called as Angiogenetic cell clusters. They lie on the lateral and
cranial aspect of the embryo forming a horse shoe shape plexus of blood vessels. The
cephalic portion of this clusters is called as Cardiogenic area. Surrounding intraembryonic
coelomic cavity form pericardium.
Other clusters of angiogenic cell appear bilaterally parallel and close to the midline.
These clusters of cell acquire a lumen. These cell become endothelial cell and it forms a pair
of longitudinal vessels. These are called as dorsal aortae. Later they get collected to
angiogenic cell clusters.

Sagital section of embryo: at 2 point endoderm of gut tube and ectoderm are in contact. They
are
a. Buccopharyngeal membrane (in front):is also called prochordal plate.
b. cloecal membrane.
Angiogenic cell clusters found infront of prochordal plate. This happens by 4-5 somite
stage. Later when head fold is formed, clusters found below the gut tube because of folding ie
cardiogenic area get shifted to ventral to gut tube. This cardiogenic area surrounded by part of
coelom called Pericardial cavity. This primitive heart tube becomes enlarged and bulges into
pericardial cavity.

Formation of Heart:
First heart form in the form of two tubes in splanchnic mesoderm ie two endocardial
tubes. Lateral thicker wall(outside) form epicardium, medial thinner wall called endocardium
and middle myocardium. Both these covered by septum of mesoderm called dorsal
mesocardium. Mesodermal cell adjacent to lateral endocardial tube become epimyocardium.
Meanwhile blood vessel develop in the yolk sac called as Vitelline vessels(these are directly
open into heart tube) and the allantoic vessels( one umbilical vein from placenta) with in
embryo. There are two veins, precardianl vein, postcardinal vein, both join to form common
cardinal veins also called as Duct of cuveir. These two veins join to form Sinus venosus
which later get connected to paired endocardial tubes.
Each endocardial tube has lateral wall ie epimyocardial and medial ie endocardial
tube. b/w these two form myocardium. Two endocardial tubes are separated by septum of
endocardium, which later disappears. Then it is in the form of single tube like structure. It has
4 parts. They are,
a. Sinus venosus: where two vein joins
b. Ventricle
c. Bubus cordis(bulbus)
d. Truncus arteriosus(truncus): continued by dorsal and ventral aortae.
 29

Straight tube first change into loop called bulboventricular loop. This loop is formed in
endothelial tube which forms bulbus and primitive ventricle. Where as sinus not involved in
loop. Close end of loop is directed to right and somewhat ventrally. Truncus and sinus
venosus lie in the midline. So cranial portion is bubus cordis and ventral portion is ventricles.
After first loop is formed it start bending such that ventricle portion lies ventrally on to right
and sinus portion lies caudally. Truncus and bulbus lies on the dorsal aspect.

First change in sinus portion: it forms two bulging on eitherside to form left and right atrium.
Atrium is placed where 2 cardinal veins open. Now it is in the form of single tube without
any partition. First change occurs is by

Development of septum in the ventricle: septum is formed by formation of endocardial

cushion. This endocardial cushion has muscular part and a membranous part. This give rise to
interventricular septum(b/w 2 portion of ventricle). Meanwhile, three endocardial cushion
between atria ie right, left and middle. Membranous part of endocardial cushion of ventricle
join middle cushion of atria. So ventricle completely divided into right and left. Space
between right and left endocardial cushion give rise to atrio-ventricular foramen ie
tricuspid(right) and bicuspid(left).

Septum formation in atrium: extending from dorsal aspect of atrium, develop a membranous
septum called septum primum. There is opening between septum primum and middle
endocardial cushion called ostium primum/primary internal foramen. Later another
septum develop towards right side of septum primum called septum secondum. It extend
whole length of atrium and touches middle endocardial cushion. When it touches, a foramen
develop at dorsal aspect of septum secondum called osteum secondum. So blood flow from
right atrium→ostium secondum→b/w septum primum and secondum→left atrium. At left
atrium, pulmonary veins opens. Blood goes to pulmonary veins. Lungs are not functional in
embryo. So blood goesto left ventricle→Aorta. Circulated blood is deoxygenated.
Oxygenated blood circulate only in umbilical vein. Immediately after birth, the lungs start
functioning. So blood flows from lung to left atrium and more blood goes to ventricle, so
septa at atrium come together and completely closed and gap between primary and secondary
septum give rise to foramen ovale. Two septa join to form Fossa ovalis.

Truncus: is a single tube from ventricle. A septum start developing in truncus region called
as Aortico pulmonary septum or spiral septum. Because of this truncus(bulbus) portion is
divided into aortal and pulmonary trunk. Aorta is in contact with left ventricle and pulmonary
trunk with right ventricle.

Anamolies in development of heart:

Ectopic cardis: ie heart is situated outside the thoracic cavity. Usually located in ventral part
of neck or thoracic inlet. It formed by failure of descent of heart from gut tube to pericardial
cavity or thorax.
 30

Atrial septal defect(ASD): when primary and secondary septum not communicate, then
there is gap called ASD ( it takes 1-11/2month to completely develop to 2 atria. In newborn
septum is not complete).

Persistent ventricular septal defect(VSD): interventricular septum is not completely

develop ie membranous part. Here communication between right and left ventricle.

Persistent truncus aorticus: failure of aortic and bulbar crest to form bulbar septa.

Tetrology of fallot: involves multiple defect anamolies in heart. It consists of pulmonary

stenosus, malformation of aortic pulmonary septum, right ventricular hypertrophy, overriding
aorta, VSD. It is common in dashound dogs.

ARTERIAL SYSTEM
Here we study the modification of aortic arch, ventral aorta, dorsal aorta and its branches.
There is paired dorsal aortae.
There are 6 aortic arches corresponding to bronchial arches. Found on either side of the gut
tube. Ventral aorta is continuation of truncus portion of the heart. (connection between dorsal
and ventral aorta-aortic arches/paired vessels form).

Fate of aortic arches:

Ist and IInd aortic arches completely disappears. Small remnant of Ist aortic arch is called as
Maxillary artery. Remnant of IInd is Stepedial and hyoid arteries.

IIIrd aortic arch gives rise to form common carotid artery and internal carotid artery. External
carotid artery arises as a branch of internal carotid artery which supply to brain.

IVth aortic arch: on left side give rise to Arch of aorta. And right side give rise to right
subclavian artery. Connection on left side is present forming arch of aorta and right side
connection with dorsal and ventral aorta disappears.

Vth aortic arch completely disappears.

VIth/pulmonary arch: form pulmonary trunk.

Connection between VIth aortic arch and dorsal aorta forms Ductus arteriosus which later
form ligamentum arteriosum. This pattern is in humans.
 31

Left subclavian artery arises as a direct branch from aorta.

Aorta has 3 parts:

1. Ascending aorta/ventral aorta
2. Arch of aorta
3. Descending aorta/dorsal aorta.
Ventral aorta between IV and V arch gives rise to ascending aorta. Arch of aorta is the IV
aortic arch proper. Descending aorta is continuation of dorsal aorta b/w IV to VI.

In domestic animals: on right side, connection between III and IV portion of ventral aorta
gives rise to brachiocephalic artery and from brachiocephalic artery right and left common
carotid artery arises.

In adult carnivores and pigs: arch of aorta on left side gives rise to brachiocephalic trunk
which give rise to right subclavian and 2 commom carotid artery. Left subclavian arise as
direct branch from aorta.

In ox and horse: Zipped up portion of 2 common carotid artery is brachiocephalic trunk as a

single trunk.

In vertebrates upto amphibian: two aortic arches on both sides.

In birds, aortic arch is on right side.

III aortic arch gives internal carotid artery continue to cranial cavity and give rise to e
cerebral artery at ventral aspect of neural tube forming arterial circle and continue on pons
and medulla oblongata as basilar artery.

Branches of aorta:

Dorsal aorta: 3 types of vessels distributed in cervical, thoracic and lumbar regions. They are
paired parietal-dorsal branch, paired visceral-lateral and paired visceral-ventral in embryonic
stage.
paired parietal-dorsal(DPP) branch:
Cervical region →7 parietal branch. Later connection b/w 7 disappear, ie. they joined and
establish connection with subclavian artery. This forms Vertebral artery(arise as a branch of
subclavian and enter cranium and supplies as ventral spinal artery)
Thoracic region→ dorsal inter costal areter(13 or depending on no. or ribs)
Lumbar region→ lumbar arteries(6-7)
 32

paired visceral-lateral(LPV): no branches in cervical and thoracic region. In lumbar and

abdominar region→ renal artery for kidneys, testicular artery in male/ovarian artery in
female.

paired visceral-ventral(VPV): vitelline arteries: no vessels at cervical region. These

supplies the gut tube. Paired visceral become single vessel (becoz gut tube become single due
to lateral fold).

In thoracic region→ bronchial a., oesophageal a.

Lumbar region→ coeliac a. for foregut(stomach, 1st and 2nd part of duodenum), cranial
mesenteric a. for midgut(duodenum, jejunum, ileum, part of caecum and colon), caudal
mesenteric artery for hind gut(colon and rectum).

Dorsal aorta continuous and at 5th lumbar level gives middle sacral and coccygeal
artery. Last branch is called as umbilical artery which is paired ie left and right. It passes
through placenta and form connection b/w mother and fetus for purification of blood.

Internal iliac artery arises as a branch of umbilical artery. Original connection

between umbilical artery and aorta is lost when internal iliac artery arises. So now umbilical
artery become 1st branch of internal iliac artery. Int.iliac a. supplies substance in pelvic
cavity. External iliac artery arise as a branch of internal iliac artery.

In head region→IIIrd aortic arch gives rise to internal carotid artery supplies to brain. External
carotid artery arise as a branch of internal carotid artery.

VENOUS SYSTEM

By 5th week, 3 pairs of veins developed with in the embryo. These are,
1. Vitelline vein(Omphalomessenteric vein)→ carries blood from yolk sac to sinus
venosus
2. Umbilical vein→ originate from chorionic villi. Carry oxygenated blood from
embryo.
3. Cardinal veins→ found within the embryo. These are 3 in no.
a. Precardinal vein(anterior)
b. Postcardinal vein(posterior)
c. Common cardinal(duct of cuvier)
These cardinal veins drain blood from embryo proper.
All these veins joins sinus venosus.
 33

VITELLINE VEIN: before entry into sinus venosus, they form plexus of veins around
duodenum. This passes through Septum transversum ( which divides the coelomic cavity
into pleural sac, peritoneal sac and pericardial sac. This septum give rise to diaphragm later).
At the level of septum transversum from duodenum as a diverticulum liver develops forming
hepatic cords. b/w hepatic cords, these vitelline vein give rise to hepatic sinusoids. This
plexus of vein on left side gets reduced. And blood from left side of the liver rechanneled
towards the right side resulting in enlargement of right vitelline veins and this results in
formation of right hepatocardiac chanel. (is called post hepatic portion of the venacava.
Later give rise to 1st portion of caudal vena cava). Proximal part of left vitelline vein
disappears. Then there is network of blood vessels around duodenum give single vessel. It
has 3 parts namely, proximal anastamoses, middle and distal anastamoses. From distal
anastamoses, vein arise from duodenum gives single vessel which has 3 parts namely, cranial
dorsal on dorsal aspect of duodenum, middle ventral on ventral aspect of duodenum and
caudal dorsal. Right part of cranial, left part of middle disappear and remains as S-shape vein
which become Portal vein. It has connection with the umbilical vein on both sides. Blood
from midgut empties into liver by superior mesenteric vein which drain intestinal loops.

UMBILICAL VEIN: initially located on either side of liver and become hepatic connection
to sinusoids. Proximal part of left umbilical vein and reminder of right umbilical vein
disappears. So only distal part remains. So there is connection from right proximal portion to
left distal part, give rise to new vessel called Ductus venosus. This brings venous blood from
placenta to the liver. It is not connected to sinusoids but bypasses the sinusoids of liver.
Totally 3 veins forms:
1. Portal vein:from vitelline vein
2. Sinusoids of liver: from vitelline and part of umbilical vein
3. Ductus venosus: from right and left umbilical vein.
CARDINAL VEINS: right and left precardinal veins and right and left post cardinal veins
join to form common cardinal vein which open into sinus venosus. Cardinal vein are main
vein drain venous blood from embryo. Additional vein called subcardinal vein which drains
from kidney and supra cardinal vein which drains from body wall b way of intercostals
veins.
Development of Anterior or Cranial vena cava: new anastamosing vessel from left
precarinal to right precardinal arises is called Brachiocephalic vessel. Rest and left
precardinal completely disappear. Now right precardinal divides into external jugular vein
and internal jugular vein. These join to form common jugular vein. From this right and left
subclavian are given off. And then these two become right and left brachiocephalic
(forelimb→brachium, from head and neck→cephalic). Two right and left brachiocephalic
vein join to give Cranial venacava which opens into sinus venosus on right side. Small
reminant of left precardinal opening into sinus venosus give rise to Coronary sinus.

Caudal venacava: has 5 sources:

1. Hepatocardiac chanel: due to enlargement of right vitelline vein form 1st part of
caudal vena cava.
2. 2nd part derived from anastamoses between two subcardinal vein(right and left). It
happens with in mesonephron or kidney called Renal anastamose. Portion infront of
this is prerenal portion. After anastamos is postrenal. Hepato cardiac chanel
 34

anastamoses with right prerenal portion of right subcardinal vein and it form 2nd part
of caudal venacava.
3. 3rd part derived from Renal portion i.e. right renal anastamos of right subcardinal vein.
4. 4th is connection between postrenal and right supracardinal.
5. 5th part is Post cardinal veins join at the level of sacral part gives rise to common iliac
called iliac anastamose. Right common iliac gives external and internal iliac vein and
left part disappear. i.e. iliac anstamose gives 5th part.
Most of the left post cardinal and sub cardinal vein completely disappear. Rest of supra
cardinal veins gives rise to hemiazygus vein(after gives of 4th part).
Hemiazygus vein→ from left supracardinal eg.pig, ox
Vena azygus vein→form if right supracardinal vein persist. Eg.horse. they drain coronary
veins to sinus venosus.

FOETAL CIRCULATION

In c/s of umbilicaus, extension of allontois, 2 umbilical artery and 1 umbilical vein. Whole
thing is surrounded by connective tissue called Mucoid connective tissue(Whatsons jelly).
Aorta terminate by 2 umbilical arteries carry impure blood from foetus to placenta for
purification. Purified blood brought to foetus through umbilical vein which opens into hilus
of liver. At hilus of liver, there is connection b/w umbilical vein and hepatic vein from
sinusoids of liver called Ductus venosus. Hepatic vein opens into caudal venacava. Caudal
vena cava bring venous blood from thorax, abdomen and hindlimb of embryo. It opens into
ventral portion of right atrium. Another portal vein bring digested food from intestine and
join liver to sinusoids of liver. Venous blood from head, neck, forelimb come through cranial
venacava. This opens into right atrium. From right atrium blood goes to left atrium through
Foramen ovale. And also passes through right atrio ventricular opening to right ventricle
then to pulmonary artery(mainly from head and neck) then goes to lung(but not functional).
So connection b/w pulmonary artery and (dorsal) aorta form Ductus arteriosus. From dorsal
aorta blood distributed to all parts of body through branch of aorta. Then ultimately aorta
divides into umbilical artery which goes to placenta.
Umbilical vein(oxygenated)→ductus venosus→caudal venacava(both oxygenated and
impure blood)→foramen ovale→left atrium.
More oxygenated blood in ductus venosus→right atrium.
Left atrium→left ventricle→aorta→common carotid artery→head, neck(more oxygenated
blood)
Venous blood from cranial venacave→right atrium→right ventricle→pulmonary
artery→thorax, abdomen, hindlimb(less oxygenated blood)
Two short cuts:
1. Formen ovale
2. Ductus arteriosus
3.
After birth: formen ovale closes forming Fossa ovalis. So blood from right atrium goes to
right ventricle → pulmonary artery→lungs→left atrium→left ventricle→aorta.
 35

Ductus arteriosus become Ligamentum arteriosum. Ductus venosus of liver also closed by
forming Round ligament of liver. Connecting body wall to visceral surface of liver.
Umbilical artery become Round ligament of bladder. So four foetal reminants.
(ductus venosus→ligamentum venosum, notochord degenerates but helps in neurilation).

UROGENITAL SYSTEM

1.Urinary system
2.Genital system
Developmentally urinary and genital system are very closely related arise from
intermediate mesoderm.
Urinary system develops from intermediate mesoderm in the form of Urinary ridge
along caudal wall of the abdominal cavity. It opens into cloeca.
In vertebrates, urinary system is divided into
1. Pronephros - early kidney, function in fishes.
2. Mesonephros – middle kidney, function in Amphibia and reptails.
3. Metanephros – definitive kidney, in birds and mammals.
[By beginning of 4th week, the intermediate mesoderm in the cervical region looses
contact with the somite and forms segmentally arranged cell clusters. These are called
Nephrotomes. These nephrotomes grow in lateral direction and contain a lumen and they are
tubural in shape. This type of tubules are called Nephrogenic tubules/nephric tubules, open
into the coelom. Externally such openings are called External glomerulli. Suppose they
don’t open into coelom, tubules are open internally called as Internal glomerulli. It consists
of two parts: Nephric tubule and external glomerulus which constitute Excretory unit. All the
tubules joined to form a common duct. It may be pronephric duct, metanephric duct and
mesonephric duct.]

In birds and mammals all the 3 are seen but only metanephros is functional.
Pronephros: is seen in the chick embryo in the form of tubules by 36hrs. It appear in the
form of tubules with extension or invagination of somites called pronephric tubules called
Nephrotome opens into coelomic cavity. Nephrotome are paired and segmentally arranged
solid cell groups in cervical region. Nephrotome consists of pronephric tubules, open into
pronephric duct.
External glomeruli are the tubules in connection with coelomic cavity. Tubule get
connection from dorsal aorta is called Internal glomeruli.
Later pronephric tubule loose connection with somite and form independent tubule
opening into coelomic cavity called Nephrostomes. Unsegmentally arranged tubules in
thoracic and lumbar region called as Nephrogenic cord. Segmented portion is Nephrotome (in
cervical region). Pronephric tubule develop in the from of 10 somite cell group in cervical
region (7-14 somites in birds, 36hrs).by 4th week in the mammals. Later these tubules
completely disappear.
 36

Pronephric tubule have external glomeruli only.

Mesonephros: is unsegmental portion which develops by 9-26 somites in the mammals. 29-
30 somite of 55hrs in birds and usually they are unsegmented in lumbar and sacral portion.
They also in the form of tubules called mesonephric tubule, these joins to mesonephric duct
called Wolffian duct opens into cloeca.
Mesonephric tubule differ from pronephros in that they have internal glomeruli and
are S-shaped tubules. All these tubules joins to form Mesonephric duct. Mesonephros become
vestigial later and important in development of genital system.
Metanephros: appear by 5th week, permanent kidney. Medial to mesonephros, is
Metanephric blastema. An outgrowth from mesonephric duct arise at where mesonephric
duct join cloeca called as Ureteric bud. Ureteric bud is a single tube arise as diverticulum.
Ureteric bud becomes longer and widens into funnel shape structure called Renal pelvis.
Then calyx major and calyx minor. This is surrounded by metanephic blastema where we get
different portion tubules of nephrons i.e. upto collecting duct.
Urorectal sinus/cloeca: it is common opening for mesonephric duct and ureteric bud. It has
connection with Allontois from where bladder develops. Septum of mesoderm arise and start
dividing cloecal common opening into 2 parts called Urorectal septum. 2 parts are:
a. Dorsal rectal sinus – where rectum opens
b. Ventral urogenital sinus – where mesonephric duct opens. Urogenital sinus is
continuous with allontois which later becomes Urinary bladder.

At junction between bladder and urethra there is single opening by mesonephirc

duct(MND) and ureteric bud. Later MND and ureter become separate with separate opening.
Allontois is lined inside by endoderm and outside by splanchnic mesoderm. UG sinus
also lined by endoderm inside. (MND and ureter derived from intermediate mesoderm). 1st
MND, then uteter opens. Later MND form ductus deferens in male opens at neck of bladder
and ureter opens first because ureteric bud start moving on dorsal aspect of bladder which is
lined by mesoderm.
Urinary bladder between two ureter and neck of bladder is called Trigonum visicae. All
portion of ureter lined by endoderm and having folds except trigonum(mesoderm lining)
vesicae and no folds ie smooth.
UG sinus later become pelvic portion of urethra.
Anamolies associated with Urinary System:
1. Hydronephrosis: kidney tissue is replaced by fluid filled cyst. Because of blockage of
ureter.
2.Renal cortical hypoplasia: there is arrest in development of metanephric mesoderm.
Common in dog and cat.
3. Ectopic ureter: ureter opens into urethra instead of bladder when there is no migration of
ureter.
4. Double ureter: two ureter: one opens into bladder and another to urethra.
5. Pelvic kidney: first kidney develops in pelvic cavity and later it shifted to abdominal
cavity. If no migration, it remains in pelvic cavity so the name.
6. Horse shoe kidney: sometime both the kidney join and look like horse shoe.
 37

GENITAL SYSTEM

It involves the development of gonads, duct system and external genitalia.

Development of Gonads:
Two stages:
a. Indifferent stage: where sex is not established.
b. Definitive stage
Urogenital ridge from intermediate mesoderm divided into 2 parts, medial genital ridge and
lateral urinary ridge. Medial to mesonephros, smaller genital ridge develop(ie genital
blastema). It is lined by mesodermal coelomic epithelium. Yolk sac endoderm migrate by
ameboid movement to dorsal mesentery and get lodge with in the genital ridge called
primordial germ cell in mammals and by vascular system in birds.
Development of gonads in both sexes takes place by 4 steps:
1. Arrival of germ cells at gonadal ridge.
2. Formation and proliferation of blastema from different gonad
3. Formation of cords of cells which surround the germ cell.
4. Differentiation and development into testis in male or ovary in female(definitive
stage)
Upto 1-3 steps is common for both the sexes.

Development of Testis(male):
First set of u-shape of cords of cells arise from coelomic epithelium arise from
periphery towards the centre. These are called as Primary sex cords or Medullary cords.
These cords of cells migrate to interior of the gonad and peripheral cords of cell disappear
and forms a network of tubule and give rise to Rete testis.
Second set of cords arise from coelomic epithelium are called Secondary cord or
cortical cord. These secondary cortical cords differentiated into seminiferous tubules. It
contain sertoli cell and cells of spermtogonia series. Sertoli cell and basement derived from
coelomic epithelial cell (secondary cortical cords). In b/w sertoli cell, cells of spermatogonia
series ie spermatogonia cell derived from primordial germ cell. These germ cell begin to
undergo meiosis. However incorporation of germ cell into seminiferous tubule and
interaction b/w cell types stops meiosis and subsequent spermatogenesis. Meiosis takes place
only after sexual maturity in the male.
In b/w seminiferous tubules, Interstitial cells or Leydig cells are present. Sperm
cells(spermatogonia series) are endodermal in origin. Sertoli and Interstitial cells are
mesodermal in origin. Two population of interstitial cells are found. One appears in the
embryonic testis and produces androgen which is necessary for development of male genital
system.
Other population forms postnataly at the onset of sexual maturity and becomes the
interstitial cells of adult testis.
 38

Development of OVARY (female):

From coelomic epithelium cords of cell arise called Cortical cords called as
Secondary sex cords. Primary cords remains same. These secondary sex cords become
differentiate into follicular cells.(primary, secondary and tertiary follicles). Follicle cells are
homologus to sertoli cells. Cords of cell towards the medulla completely disappears →
medullary cords. Remnant of these gives rise to Rete ovary.
With in follicles, is Oogonia. Granulosa cell surround oogonia derived from
primordial germ cell. These oogonia get differentiated into primary oocyte. These primary
oocyte undergoes 1st meiotic division till late prophase (diplotene stage). And primary oocyte
→ secondary oocyte and stops at diplotene stage called Dictiate stage. At this stage it is
arrested until sexual maturity.
Oocyte in foetal ovary is not surrounded by follicular cell complete meiosis
prematurely but degenerate. This occurs to high degree in ovary and as many as 90% oocyte
degenerate. Among domestic mammals, highest germ cell survival found in pig in which
50% cell towards medulla give rise to Rete ovary which completely or partially degenerate.
Degenerated medullary part contain blood vessel and nerves and go to form the
medulla of the ovary. Those which donot degenerate, form rete ovary. Formation of rete
ovary appears to influence onset of meiosis in female at the time of oestrus.

DUCT SYSTEM:
Two sets of ducts appear.
1. Mesonephric duct(MND) also called Wolfian duct.
2. Paramesonephric duct(PMND) also called Mullerian duct.
Parallel and ventral to MND is PMND. Indifferent gonad is present and MND and PMND
opens into urogenital sinus.
Indifferent gonad get differentiated into testis or ovary due to XX or XY
chromosome. XY is because of Hy antigen in male is secreted by Y chromosome in male. So
differentiate into testis. Testis start secreting Mullerian inhibiting factor(MIF). This is
responsible for arrest in development of PMND but influence development of MND.

In male, Gonad→ development of seminiferous tubule and rete tesis (network of tubule).
PMND completely disappear and MND remains.
In MND→ number of mesonephric tubule join to form mesonephric duct. Mesonephric
tubule in the region of testis give rise to Efferent ducts and they establish connection with rete
testis. Part of efferent duct gives head of epididymis. MND give rise to other part of
epididymes (body and tail) and ductus deferens and these open into cloeca or urogenital
sinus. Tubules infront of testis completely degenerate and remnant of this become
paradidymes. Remnant of tubules behind testis become Appendix of testis.

In female: because of XX chromosome, indifferentiate gonad get differentiate into ovary.

Ovary does not secrets MIF. So MND degenerate and PMND remains.
Cords of follicles, rete ovary, cortex and medulla in ovary because of absence of MIF, there
is persistence of PMND or ovarian duct and degeneration of MND.
 39

PMND does not establish connection with ovary. So it forms oviduct and uterus (horn, body
and cervix). Sometime find reminant of mesonephric tubule and form paraoophorn and
enoophorn. Cranial most portion form oviduct, unfused portion form horns of uterus. Fused
portion form body and cervix. Last portion where two fused PMND, opens into UG sinus
form cranial portion of Vaginal. Rest of vagina is derived from UG sinus.

Various uterine shape in different species:

In some long horn, short body→ carnivores, pig
Short horn, long body→ cow, mare these depend on extent of fusion of PMND.
In primates only body and cervix, no horn.

Two PMND fused and give rise to body and cervix of uterus. Junction where PMND
joins to UG sinus is Vaginal plate. Fused portion is called Mullerian tubercle is a solid
structure. Mullerian tubercle give rise to cranial part of vagina. UG sinus form caudal part of
vagina. Cervix open into vagina → Postro vaginalis of cervix. This part of vagina has
columnar epithelium. Rest part of vagina lined by stratifies squamous epithelium. UG sinus
establish connection with allontoise(urinary bladder) form External urethral orifice. Portion
infront of this orifice is vagina and behind orifice is Vestibule. So vagina derived partly from
Mullerian tubule (PMND, mesoderm), UG sinus(endoderm).

Accessory glands in Male:

MND opens into UG sinus. Vesicular gland derived from MND as a diverticulum
where it joins the UG sinus (2 outgrowths). It is mesodermal in origin. From urogenital sinus
two diverticulum arises called as Prostate gland. Bulbo urethral gland also derived from UG
sinus as a diverticulum at the level of Ischial arch.
In female: accessory glands are major and minor vestibular glands found at the floor of
vagina. These are homologus to prostate and bulbo urethral gland.

Urorectal septum;(mesoderm) divide cloeca into dorsorectal sinus and ventro urogenital
sinus.
Cloecal plate: where ectoderm of body wall and endoderm of hindgut or UG sinus meet.
Later it divided into 2 parts by urorectal septum.
From behind at the level of cloecal plate,
a. Anal opening
b. Urethral opening
Surrounding anal opening is a fold of mesoderm called anal fold. Surrounding urethral
opening another mesodermal fold called urethral fold. On either side of urethral fold two
more folds called Genital fold or Scrotal fold. Junction between anal fold and urethral fold
become elongated giving rise to perineum. This is in indifferent stage.

Development of External Genitalia: in indifferent stage urethral fold become elongated to

give rise to Genital tubercle or Phallus in a cranial direction.
 40

In male: Genital tubercle give rise to body and glans penis. From MND to ischial arch form
pelvic urethral. Portion of urethra incorporated into genital tubercle and become Penile
urethra.
Scrotal folds forms the Scrotum.
In female: urethral folds give rise to Pelvic urethra or Vestibule.
Scrotal folds form the Labia vulvae.
Genital tubercle give rise to body, and glans of clitoris.
Ligaments of Genital organs:
Female: mesonephros completely degenerats and only MND is formed. In place of
mesonephros, mesenteric fold of tissue is present. In medial edge of this fold are two cord
like ligaments. Ligament cranial to gonad give rise to Suspensory ligament of ovary.
Ligament caudal to gonad form, Proper ligament of ovary. Mesenteric fold persist and
suspens the structure form from PMND give rise to mesosalphinx and broad ligament of
uterus. Mesosalphinx may completely enveloping forming Ovarian bursa.
In male: mesonephros form suspensory ligament of testis in a transitory structure. Ligament
caudal to testis form proper ligament of testis or mesorchium. Another ligament suspending
tail of epididymis is Ligament of tail of epididymis.
Proper ligament of testis + ligament of tail of epididymis form Gubernaculum testis which
plays major role in descent of testis.

Descent of Gonads:
Both ovary and testis move to a position caudal to their original position(lumbar region)
→descent to their position in male and female.
In female: there is a intra abdominal descent which is greater in the ungulates like ox and
horse. In pig and carnivores retained in their original position ie. Sublumbar region.
In male: testis descents from intra abdominal level to the scrotum
1. Intra abdominal passage
2. Inguinal passage
3. Scrotal passage
All the passage are by shortening of gubernaculums which push testis into scrotum. So
gubernaculums helps in
1. Transfer of testis to scrotum
2. Accessory function in descending process brings about formation of inguinal canal
3. Dilate inguinal canal sufficient to testis to pass through.
4. Ground substance of CT. of the gubernaculums secretes steroid harmone which helps
in descent of testis to occur.
Male Indifferent stage Female
Gonad Spermatogonia Primordial germ cell Oogonia
Sertoli cell Mesonephric Follicular and
cell(mesodermal cell) granulose cell
Rete tesis Medullary cord Rete ovarii
 41

Interstitial Interstitial blastema Interstitial cells of

cells(Leydig cells) ovary
Mesonephric Efferent duct Mesonephric tubule
tubules
Paradidymis, Cranial tubules Enoophron,paraoophor
Appendix of testis on (vestigeal)
Epididymis, Ductus Mesonephric duct Gartner’s duct, ureter
deferns, Ureter,
Vesicular gland
Vestigial giving Paramesonephric duct Oviduct, Uterus(horn,
rise to uterus body), cervix, part of
masculineus(horse) vagina(cranial portion)
Prostate gland, Urogenital sinus Vagina, vestibule
bublourethral
gland,pelvic urethra
External urethra Urethral folds urethra
genitalia
Scrotum, phallus Genital(scrotal) fold Labia vulvae
Body and glans Genital tubercle Body and glans of
penis clitoris

Anamolies in Genital system:

Hermophrodite: where gonadal tissue of both sexes is present in an animal. It may contain
ovotestis (ovary or testis)
Pseudohermophrodite: gonads of one sex and genital organs of other sex. Male
pseudohermophrodite is common than female.
Cryptorchidism: failure of testis to descent into the scrotum. It may be unilateral or bilateral.
Hypsospadia: in male, where urethral folds meet creating an opening at ventral aspect of
urethra.
Epispadia: where two urethral fold donot meet at dorsal aspect of urethra.
Gonadal dysgenesis: due to chromosomal abnormalities.

ENDODERMAL DERIVATIVES
DIGESTIVE SYSTEM
Development of Pharyngeal region.
There are 6 pharyngeal arches. I, II, III, IV, V and VI arches. Arches are mesodermal. From
each arch has got a aortic arch from which nerve, muscle, and skeletal element(bone or
cartilage) are derived. Each arch is separated from endoderm inside is called bronchial
pouches/pharyngeal pouches. (endodermal pouch). Outside lined by ectoderm called as
Pharyngeal cleft(I, II, III, IV, V and VI)
 42

Development of Tongue:
At the floor of the pharynx by about 3rd week, from the 1st arch, a pair of enlargements takes
place. This is called as Distal tongue bud or Lingual swellings.
From 1st and 2nd arch in the midline there is a medial swelling called Medial tongue bud or
Tuberculum impar.
At junction of 2nd and 3rd arch, a pair of swelling called as Proximal tongue bud or Cupola
or Hypobronchial eminence.
From 4th arch,behind cupola another swelling called as Laryngeal swelling/epiglottic
swelling give rise to Epiglottis.
Distal lingual swelling become elongated and united one another and give rise to tip and body
of tongue. Medial swelling give rise to rest of body of tongue. Proximal swelling or cupola
give rise to Root of tongue. Where two distal lingual swelling joins give rise to Sulcus
linguae. Junction between 1st and 2nd arch, especially in ruminants gives rise to Torus linguae.
Musculature of tongue and other CT of tongue derived form respective arches and only
epithelial lining derived from particular pouch.
Epithelial lining tip of tongue→ 1st pouch
Epithelial lining body of tongue→ 2nd pouch
Epithelial lining root of tongue→ 3rd pouch.
Nerve supply to tongue is by 3 types:
1. General sensory nerve(sense of touch, pain is supplied by lingual branch of
mandibular nerve which supplies cranial 2/3rd of tongue and caudal 1/3rd is supplied
by lingual branch of IX) -derived from 5th and 9th cranial nerves.
2. Gustatory fibre/taste fibers- derived from 7th and 9th cranial nerves.
3. Motar nerve- from 12th cranial nerve.
Nerves of 1st arch is V, 2nd arch is VII, and 3rd arch is IX. but do not have general sensory
nerve from VII(becoz, eventhough body of tongue by 2nd arch, epithelial lining of root of
tongue overlies body of tongue(3rd pouch) so no GSN from 7th only 5th and 9th).
Muscles of tongue arise from occipital somite so supplied by 12th nerve.
Anomalies during development of tongue:
Microglossia: short lingual swelling.
Macroglossia: elongation is very much long.
Forked tongue: two lingual swelling remain separate without union.
Tongue tie: during elongation of lingual swelling there is attachment with floor of mouth
called Franulum linguae. If attachment is too much, no movement of tongue called as tongue
tie.

Derivatives of Pharyngeal pouches/bronchial pouches: 6 pouches

1st pouch: divides into dorsal and a ventral part. Dorsal part has broad(distal) and
narrow(proximal) part. Broader distal part become Tympanic cavity(middle ear cavity).
Narrow portion give rise to Isthasian tube/pharyngotympanic tube(connection b/w pharynx
and middle ear). Also called as Tubotympanic recess. Ventral part disappears.
 43

2nd pouch: also has dorsal and ventral part. Ventral part disappears. Form dorsal part, mucus
membrane of pharyngeal tonsil develop. Aggregation of lymphatic tissue around tonsil
derived from mesoderm of the arch. This mucus membrane is smooth or having crypt.
3rd pouch: dorsal and ventral part. Dorsal portion divides into cords of cells which give rise
to External parathyroid or parathyroid crypt. This get migrated and found at bifurcation of
external carotid and maxillary artery. Ventral part give rise to epithelial reticular cells of
thymus(cervical portion). In between thymocytes similar to lymphocytes which derived from
mesoderm.
4th pouch: dorsal and ventral part. Dorsal part gives internal parathyroid always found
associated with thyroid gland. Ventral part give rise to Thoracic part of thymus.
5th and 6th pouch: give rise to Ultimobranchial body. This is a group of epithelial cell
which is found as a separate structure in vertebrates upto birds but in mammal, get
incorporated with thyroid cell called parafollicular cells or ‘c’ cells which concern with
calcitonin secretion but in birds it is a separate structure located at the level of thoracic inlet.

THYROID GLAND:
It is derived from endoderm at the floor of the pharynx. It appears 1st as endodermal
diverticulum called Thyroid diverticulum at the level of 2nd branchial pouch. It is closely
associated with root of tongue and the connection is called Thyroglossal duct. Proliferation
of cells of thyroid gland, duct get obliterated and lumen become solid cord of cells.
Diverticulum become enlarges and gives number of follicular cells and connectiong with
pharynx is lost. And it moves more caudal direction so it passes more towards area of 4th
pharyngeal pouches (ie larynx and trachea). These cords of cells give rise to follicles lined by
cuboidal epithelium. Ultimobronchial body get incorporated in thyroid cells and looses its
identity and cells become parafollicular cells or C cells. Later it divides into two lobes
connected by isthmus.

FOREGUT
Foregut is divided into pharyngeal region gives tongue, thyroid gland etc. behind this is
oesohageal region.

Development of Oesophagus:
It develop from portion of foregut behind pharynx. It maintain its original tubular shape.
Lining epithelium is derived from endoderm of foregut. While surrounding mesoderm gives
rise to muscle and other CT elements. There is increase in length of oesophagus to keep up
the growth of neck and thorax.
Anamolies:
Megaoesophagus: there will be increase length of oesophagus.
Oesophagus acalasia: there will be constriction of lumen of oesophagus.
 44

Development of Stomach:
It appears as a dilatation of foregut behind the oesophagus. This dilatation takes place
both dorsally and ventrally. Enlargement is more on dorsal aspect compared to ventral aspect.
After enlargement, it undergoes rotation to 900, then what was dorsal become left and ventral
become right. There is further enlargement of the left side. This enlargement of left side
becomes greater curvature. While right side becomes lesser curvature. Liver which is on right
side of abdominal cavity rapidly enlarges. So this rapid enlargement pushes cranial end of
stomach further to left so that long axis is roughly at right angle to the long axis of the
embryo. Cranial part of the dorsal enlargement grows more than caudal part and form fundic
portion. Narrow part(caudal) ie pyloric part continuous with duodenum. Boundary of dorsal
enlargement form greater curvature and original ventral part form the lesser curvature. So the
stomach of adult has greater curvature towards left and lesser curvature towards right. This is
in monogastric animals(dog, horse, pig)
In ruminant stomach: first it is in the form of tube. All rumen, reticulum, omasum and
abomasums are of same size. Left side shows two enlargements ie rumen and reticulum.
Right side is omasum and abomasums. Enlargement of rumen forming dorsal and ventral sac.
Remaining part of foregut after stomach form 1st part of duodenum.

Development of Liver:
Parenchyma of liver is derived from endodermal cells of foregut immediately after
stomach called as Hepatic diverticulum or liver bud. These diverticulum goes on dividing
into number of branches to form cords of cells called Hepatic cords. In between hepatic
cords, sinosoids derived from umbilical vein and vitelline vein. Where hepatic diverticulum
joins foregut(duodenum) is form Hepatic duct or bile duct. At this point where hepatic
diverticulum joins foregut another diverticulum arises which form the Gall bladder. From gall
bladder, cystic duct arises. Cystic duct + hepatic duct join to form common bile duct or
ductus choledochus.
Liver cell is covered by CT capsule. This capsule is derived from the mesoderm
surrounding. Hepatic cords are endodermal origin. Capsule is called Septum transversum
which later become part of diaphragm.

Development of Pancreas:
Derived from endoderm of foregut. It arises as two epithelial diverticulae of foregut. Dorsal
pancreatic(DPB) and ventral pancreatic bud(VPB). VPB comes out from common bile duct
as a diverticulum ie as a branch of hepatic diverticulum. Where it joins common bile duct.
DPB arises opposite to this ie it is from main part of foregut. Then there is differential growth
and torsion of initial part of duodenum brought about by rotation of stomach. Because of this
rotation, dorsal and ventral pancreatic bud become close to one another and becomes single
mass with left lobe (DPB) and right lobe (VPB) and body (centre). Each one of these bud
have independent duct.
Duct of VPB is called Principal or main pancreatic duct or duct of wirsung.
Duct of DPB is Accessory duct or duct of Santorini.

Different type of duct system in different animals:

 45

In some ventral remains, dorsal disappears. Ie MSc PhD

In horse and dog→ two ducts, both are remain. Main duct open along with common bile duct
which open on a major duodenal papillae. Another duct, accessory duct is further apart open
separately.
In man, sheep→ only a single duct which derived from ventral pancreatic bud so principal
pancreatic duct. Here two duct join to form a single duct.
In cow, pig→ main duct is Accessory duct to which duct from VPB come and join.
Rest of foregut giverise to first part of duodenum, 2nd and 3rd part is by midgut.
So foregut→ oesophagus, stomach, liver, pancreas and 1st part of duodenum develops.

MIDGUT
It is the portion of gut tube b/w foregut and hindgut. It is in connection with yolk sac.
Midgut is in the form of a straight tube. Later enlargement of midgut forming a U-shape
structure. So there is Cranial limb and Caudal limb. In the axis(centre) there is cranial
mesenteric artery. There is rotation of midgut at 1800, so what was cranial limb become
caudal and what was caudal become cranial limb and it is 1st rotation.
From original cranial limb, we get part of jejunum, 2nd and 3rd part of duodenum, and part of
ileum. From original caudal limb part of ileum, caecum and ascending colon. This is in dog
where colon is made of ascending and descending colon.ie only one rotation.
In cow and pig, there is 2nd rotation of first part of colon→ ascending colon is very much
enlarged to give rise to 3 parts of colon namely, proximal loop, spiral loop and distal loop in
cow. And in horse, ascending colon has right ventral, left ventral, right dorsal and left dorsal
parts.
Yolk sac is attached b/w cranial and caudal limb and when yolk sac is reduced, it is in the
form of cicatrix scar tissue called as Meckel’s diverticulum in fowl.

HINDGUT
It is last part of the gut tube behind the midgut. Usually in the form of straight tube. From
hindgut we get transverse colon and descending colon and rectum. This is retained in all
animals. Rectum opens into cloeca or urogenital sinus.
Anus: is derived from 2 sources. Cranial part is formed by partitioning of cloeca into a dorsal
rectal sinus and ventral urogenital sinus by urogenital septum(mesoderm). Endoderm form
caudal part of anus.
All these are endoderm and is in contact with ectoderm called Cloecal plate/cloecal
membrane. That part of cloecal membrane covering anal canal is Anal membrane.
Anamolies:
Megaoesophagus: lack of oesophageal peristalysis. Embryological basis is not known, but
physiological disruption of swallowing reflex or neuromuscular maldevelopment.
Intestinal atresia: involving atresia coli, atresia jujuni and atresia ileai. It is due to failure of
embryonic gut to recanalise.
 46

Atresia ani: rectum and anus come in contact with ectoderm at cloecal membrane. If cloecal
membrane is not rupture ie persistant anal membrane.
Urorectal fistula: urorectal septum not completely develop. So rectal particle goes to the
urogenital sinus. Communication between rectum and urinary tract.
Persistant Meckel’s diverticulum: midgut (cr. And cd.) unite and communicate with yolk sac.
This connection with yolk sac completely disappear with small scar like tissue called
Meckel’s diverticulum in poultry. Occasionally yolk sac present as a diverticulum to which
loops of intestine descents and there is incomplete development of body wall usually called
as Umbilical hernia. Due to failure of embryological intestinal mass to return to abdominal
cavity.

RESPIRATORY SYSTEM
Development of Larynx:
Larynx-trachea arise from floor of pharynx in the form of a groove or sulcus in the midline at
about the level of IV-V branchial arch. This groove is called as Laryngo-tracheal groove.
Laryngo tracheal groove on either side has Arytenoid swellings. This groove become deeper
into underlying mesenchyme. In growth, from the sides of groove form a pair of ridge and go
towards each other and fused and forms a septum. That changes the groove into blind tubular
structure which opens into pharynx.
Septum is called Tracheo-oesophageal septum which separate dorsally located oesophagus
and ventral larynx and trachea. Blind tube form the trachea and the larynx.
As soon as trachea-oesophageal septum is formed a pair of elevation called arytenoids
swellings at the floor of pharynx along with midline at the level of IV arch and around the
opening of tracheal tube arises. In front of it is covered by Hypobranchial eminence which
give rise to root of the tongue.
Depression between hypobranchial eminence and arytenoid swelling become lumen of the
larynx.
Caudal part of hypobranchial eminence forms the epiglottis. So epiglottic cartilage come
from hypobranchial eminence at the level of III and IVarches.
Arytenoid cartilage derived from IV arch
Thyroid cartilage from III and IV arch
Cricoid cartilage from V arch.
Intrinsic muscles of larynx from V and VI arch. Mucus membrane lining larynx comes from
respective branchial groove.
Mucus membrane lining epiglottis→ III and IV arch
Arytenoid and thyroid→IV and V
Cricoids → V arch.
 47

Development of Trachea:
It arises as a diverticulum from the floor of oesophagus called as Tracheal diverticulum. It
is in the form of tube separated from the oesophagus by septum called trachea oesophageal
septa/groove. Groove become deeper and deeper and finally gets separated from oesophagus.
So oesophagus dorsally and trachea ventrally.
Trachialis is mesodermal in origin arising from oesophagus.

Development of Lungs:
Trachea starts elongation and gives rise to two buds called as Lung buds.(Caudal end of
trachea arises Tracheal primordium which gives two parts called Lung buds). These lung
buds form branchings into primary, secondary, tertiary into underlying mesenchyme which
separates the bronchial tree. Branching is called as Dichotomus branching. These branching
give rise to bronchi and alveoli.
3 stages of development of lung:
1.Pseudoglandular stage: looks like a secretory gland. Almost covered by cuboidal to
columnar type of epithelium.
2.Canalicular phase: 2nd stage give rise to bronchial tree forms 3-4 generation of terminal
branch which expands called canalicular phase. Here it is lined by more squamous type of
epithelium. Here enormous increase in vascularity of lungs.
3.Terminal sacular or Alveolar stage: during this stage, the terminal branches appear as
dilated thin walled sacules resembles alveoli of lungs.
Anomalies:
Oesophago tracheal fistula: when trachea gets separated from oesophagus, it joins cranially
lying larynx. Some time trachea communicate with the larynx, it retained the original
connection with the oesophaus called as oesophago tracheal fistula. Which bring about
bloating of the stomach or food particle may enter into lungs.

Development of Coelomic cavity:

First formation of coelom is indicated by splitting of lateral mesoderm into somatic and
splanchnic mesoderm. Lining gut tube is splanchnic mesoderm and lining body wall is
somatic mesoderm. In between these two is coelom. Coelom has 2 parts,
1. Extraembryonic part: form extraembryonic membrane(amnion, chorion, allontois)
2. Intraembryonic part: part is with in embryo. These are pair of chanels become
continuous with extraembryonic coelom. The development of head region, IE part
become horse shoe shape. it has got a cranial part. Coelomic cavity surrounding the
heart is called as Pericardial coelom. Caudal part is Peritoneal coelom. these two
has a connection on either side. This communication is called as Pericardio
peritoneal canals.from this we get the pleutal cavity and peritoneal cavity. In the
early embryo, all parts of intraembryonic coelom are continus with one another. It
give rise to 3 major cavities like pleural, pericardial and peritoneal cavity.
Separation of pleural cavity from the pericardial cavity: heart is covered by layer of
mesoderm and coleom cavity called pericardial cavity. Pleura pericardial membrane are fold,
appears in mesoderm called pleura pericardial septum/fold/membrane. These fold fused with
 48

mediastinum. As lungs are expanded, so mesoderm lining the lungs forms the pleura.
Mesoderm lining heart forms the pericardium. Pleura pericardial septum become deeper and
giverise to mediastinal pleura. Again expansion of lung and occupies position lateral to
heart. So we get visceral pleura, mesenchymal pleura. So become pericardial cavity and
pleural cavity. Lungs go ventrally around heart form a sac like structure called pericardial
sac. Sac like structure surrounding lung become pleural sac.
Separation of pleural and peritoneal cavity:
First indication of separation of pleural and peritoneal cavity is formation of Septum
transversum from ventral aspect of body wall. This divide partly pleural and peritoneal
cavity. From dorsal wall, another septum arises called pleuroperitoneal membrane. Fusion
of these membrane from dorsal and septum from ventral aspect separate original coelom into
pleural and peritoneal cavity and these membrane and septum get incorporated into
diaphragm.
Development of Diaphragm:
It is a muscle and mesodermal in origin. It divide thorax from abdomen. At first, it arises as a
CT membrane. Later muscle get incorporate into to it by somite. Innervate by 5,6 and 7 th
cervical vertebrae(myotomes). Ventral portion of diaphragm arise from Septum transversum.
Lateral portion are paired arises from mesoderm and pleura peritoneal membrane. Passing
through diaphragm are aorta, oesophagus and venacava. Surrounding oesophagus(gut tube)
and aorta(dorsal) is the mesenchymal tissue. Tissue surrounding is hiatus oesophagus and
hiatus aorticus. Meanwhile myotome from 5,6 and 7th cervical get migrated to form the
diaphragm so supplied by brnches of 5,6 and 7th.
Anamolies during development of Respiratory system:
Oesphageal tracheal fistula: if original connection is not lost, there is communication b/w
oesophagus and trachea forming fistula by incomplete partition of oesophagus and trachea.
Tracheal hypoplasia: results in trachea with a reduced diameter. Common in bracheocephalic
breeds.
Pulmonary hypoplasia: here lungs are smaller than normal with poor alveolar development.
Accessory lung: observed in dog, horse with in the thoracic cavity. Abnormal bud develop
from trachea.

Peritoneal cavity: peritoneum lining abdominal cavity. There is gut tube. Gut tube is held
dorsally and ventrally by 2 folds of peritoneum called as Dorsal mesogastrium and Ventral
mesogastrium. During rotation of stomach, dorsal mesogastrium comes to left side and
ventral mesogastrium comes to right side.
Dorsal mesogastrium has a cavity called Omental bursa. Dorsal mesogastrium attach to
greater curvature become greater omentum. On lesser curvature become lesser omentum.
Along ventral mesogastrium or lesser omentum, liver is developing so lesser omentum
become connection b/w lesser curvature of stomach to visceral surface of liver. And spleen is
developing along dorsal mesogastrium. This portion form gastrospleenic ligament. Pancreas
develop as diverticulum on dorsal aspect. Partly covered by peritoneum and partly
retroperitoneum. Liver to periphery is falciform ligament.
In midgut and hindgut, loops of interstine covered by dorsal and ventral mesogastrium for
mesocolon, mesojejunum, mesoduodenum.
 49

Anamolies during development of diaphragm:

Diaphragmatic hernia: 3 types
1. Pleuroperitoneal-along peritoneal membrane.
2. Peritoneo pericardial – peritoneo pericardial membrane.
3. Hiatus hernia- hiatus
First one is common where communication b/w pleural and peritoneal cavities.
Hiatus hernia occurs at point where oesophagus passes through diaphragm due to incomplete
fusion of pleura peritoneal membrane and hiatus oesophagus.

ECTODERMAL DERIVATIVES

Development of Nervous System:

Ectoderm infront of Hensen’s node forms a thick plate which is slipper shape called
Neural plate. Lateral edges of neural plate has Neural folds. In b/w neural fold is Neural
pit.Two folds approach one another to form Neural tube.
Fusion of neural folds first takes place in cervical region. Later it extends cranial and
caudal direction. Where fusion takes place in cranial direction, there is opening called
Anterior Neuro pore. And in caudal direction there is no definite neurofold which expand in
the form of rhomboid like structure called Rhomboidal fossa (here fusion is incomplete).
Neural tube give rise to central nervous system consists of brain and spinal cord.
Development of CNS is called Neurilation. Notochord induces development of neural tube.
Note: notochord plays exact role. If remove the notochord, neurilation doesnot develop. So
notochord induces development of neural plate. This is called Induction. So notochord act as
inductor for development of nervous system. Inductor is any structure or substance which
help in development of other structure. Induction also defined as process where by one
population of cells referred to as inductor, acts upon another population of cells to change
behavior of 2nd population in some developmentally significantly like to grow or undergo
morphogenesis.
Properties of inductor:
Inductive effect is lasting and permanent and resulting changes in the induced cell population
are passed on to successive generation even after the inductor has gone. Cells thus
permanently affected are determined or evocator. Here notochord is inductor and neural tube
is evocator or determined. In regard to timing, each system inductors emerge in its strictly
sequential manner ie. primary inductor followed by secondary inductor which follow tertialry
and soon.
Eg. Notochord induces development of neural tube. From neural tube→optic
vesicle→lens→cornea. So notochord is primary inductor. Neural tube is secondary inductor.
Tertiary induction leads to formation of optic vesicle.
Induce cell population must be at a stage of differentiation where by compitation be affected
by inductor. Ie. inductor appears prematurely or long location, no inductions occurs.
Mechanism of induction: exactly not known.
 50

Neural tube is a cylindrical structure. Cranial portion of neural tube give rise to brain
and caudal portion give rise to spinal cord. This neural tube is lined by epithelium called as
Neuroepithelium. some says it is stratified columnar or pseudostratified columnar epithelium.
Neuroepithelium get differentiated into 2 types of cells. Namely
a. Glioblasts:give rise to neurogia cells like astrocyte, oligodendroglia, ependyma.
Except microglia cells which derived from neural crest.
b. Neuroblasts: give rise to neurons.
Development of Neurons: neuroblasts cells are spherical shape without any process, Apolar.
Later two process develop ie Bipolar. In third stage, one of these process withdrawn so
Unipolar. In 4th stage, get multipolar, Where one process remain and other process develop.
Ie one become axon and other process become dendrities.
Unipolar stage seen in embryonic condition.
Pseudounipolar →bipolar cell processes start approaching one another. They join and become
separate again to form pseudounipolar. One process become axon and another dendrite.
At 1st neural tube has only one neuroepithelial layer. Later, neuroepithelium have 3 zones.
1. Outer Marginal layer: form white matter as it contains processes of neuron.
2. Middle Mantal layer: grey matter. Contain cell bodies of neuron
3. Inner Ependymal layer: surrounding central canal or choroid plexus of ventricles of
brain.

Development of Spinal cord:

It develops in the region of caudal part of neural tube behind brain. It is made up of
neural tube with small central canal.
In vertebral column, it has 3 layer, namely ependymal layer, mantle layer and
marginal layer. It is circular in shape. in mantle layer, more of neuroblast added to give rise to
neurons. It becomes Grey matter. Population of neuron is more towards lateral aspect
compared to middle so middle portion become sunken. So we get 2 horns in dorsal aspect
called Alar plate. Ventral aspect called Basal plate. Two horns in alar plate called Dorsal
horn. And in basal plate called Ventral horn. Two are separated by Sulcus limitans. Area
above sulcus limitans is Alar plate, and below is Basal plate. At the level of dorsal horn,white
matter sinks to form Dorsal median septum and ventral, Ventral median fissure.
Between two dorsal horn, portion is called Roof plate. b/w ventral horn is Floor
plate. From mantle layer, get alar plate ie dorsal horn. It contain sensory and association
neurons. Basal plate forms ventral horn which has motor neurons.
Roof plate give rise to Dorsal median septum and Floor plate give rise to Ventral median
fissure.
Marginal layer contain processes of neurons. It forms white matter. It is divisible into dorsal,
lateral and ventral fasiculi.
Cells lining the central canal are Ependymal cells.
Origin of peripheral nerves/spinal nerves: dorsal root contains dorsal root ganglion which
derived from neural crest. From dorsal root ganglion, dorsal root is sensory root. Ventral root
formed by neuron in ventral horn and intermedio grey column, ventro lateral. Neurons of
these give rise to axons. All these axons join to form Ventral root. When dorsal and ventral
 51

root join, they form Spinal nerve. Ie combination of dorsal and ventral root so combination
of neural crest and basal plate.
Nerve process go outside CNS, and end in various parts of the body. These nerve axons are
covered by myelin sheath. This myelin sheath in peripheral nervous system is formed by
Schwann cells(derived from neural crest). Where as axons with in CNS are covered by
myelin fiber, myelin sheath form by oligodendroglia(derive from Glioblasts).

Development of Brain:
Brain develops from the cranial portion of neural tube. This portion undergoes three
enlargements called Brain vesicle. They are,
1. Procencephalon →Forebrain
2. Mesencephalon →Midbrain
3. Rhombencephalon →Hindbrain
Hind brain continuous in the Spinal cord. Flexure develop between spinalcord and brain is
called as Cervical flexure.
Procencephalon divided into Telencephalon and Diencephalon.
Mesencephalon remains as it is.
Rhombencephalon divided into Meterncephalon and Myelencephalon.
Fate of each region:
Myelencephalon: Medulla oblongata and Pons.
At the level of myelencephalon and Metencephalon, neural tube is open(not completely
closed). Ie sulcus limitans. Lateral portion become Alar plate. And Medial portion become
Basal plate. Basal plate contains motor nuclei of cranial nerves(12, 11, 10,9,7,6,5,4,3)
Alar plate give rise to Sensory nuclei from 10th to 5th.
Basal plate →Motor nuclei from 12th to 5th. (becoz 3rd and 4th in midbrain).
Roof plate →Choroid plexus of 4th ventricle and Medullary vela (at roof of 4th ventricle).
Floor plate→Pontine tract and pyramidal tracts (actually migration of white matter).

Metencephalon: give rise to Cerebellum. Deep flexure formed at the region of pons is called
Pontine flexure. Dorsal portion of pontine flexure give rise to two structure called Rhombic
lips. This rhombic lips has Extra and Intra ventricular portion. (These two portion join to
form Isthamus). These goes to form cerebellum. So called as Cerebellar plate. Two intra
ventricular portion join to form Vermis of cerebellum. Two extra ventricular portion form
Lateral hemisphere. Cerebellar plate has 3 regions like marginal, mantle and ependymal
layers. Mantle layer of cerebellum form the cerebellar cortex which is divided into 3 layers
namely,
1. Outer Molecular layer
2. Middle Perkinje cell layer
3. Inner Granular cell layer.
White matter in marginal layer become inside and processes of these cells form the cerebellar
medulla.
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Mesencephalon(Midbrain): originate from original mesencephalon. Only structure resemble

spinal cord.
Alar plate give rise to Corpora quadrigemina ie rostral and caudal colliculus.
Basal plate give rise to Motor nuclei of 3rd and 4th cranial nerve, red nucleus, substantia nigra.
Roof and Floor plate→no structure.
Cerebellar peduncle on ventral aspect of midbrain made up of fiber tract derived from
migration of white matter in ventral direction.
Ventral part(floor plate) give rise to interpeduncular fossa. Portion of ventricle becomes
narrow and give rise to Aqueduct of sylvius.

Diencephalon: another flexure at region of midbrain is called Cranial flexure. infront of

midbrain it involve diencephalon and Telencephalon. Diencephalon has Thalamus,
Epithalamus and Hypothalamus.
Alar plate(dorso lateral) give rise to Thalamus, epithalamus and hypothalamus(some say in
basal plate)
Basal plate(ventro lateral) →no motor area, so no structure. Sometime hypothalamus concern
with ANS.
Roof plate(dorsal plate)→ choroid plexus of 3rd ventricle and pineal.
Floor plate(ventral plate)→no structure.

Telencephalon: develop as two extensive lateral outer growth to form two Cerebral
hemisphere. Cavity between two cerebral hemisphere form Lateral ventricle.
Basal plate→ no structure. Some include Corpus striatum found at floor of lateral ventricle
(caudate and lenticular nuclei)
Alar plate→ cerebral cortex.
Roof plate→ corpus callosum and choroid plexus of lateral ventricles.
Floor plate→no structure.
Cortical area of cerebral cortex has 3 parts.
1. Neopallium → forming gyri and sulci of cortex.
2. Palleopallium → form hippocampus.
3. Archipallium → basal part of brain ie olfactory part, rhinencephalon.

NEURAL CREST:
In Neural plate, 2 neural fold meet in the form of Neural tube. Top most cells of neural fold
get separated from neural tube and occupy dorso-lateral position. These cells are called
Neural crest cells. And these migrate in two direction.
a. Dorso-lateral migration- cells are usually found under the skin(ectoderm) and they
give rise to pigmented cells like Melanoblast which become melanocytes in epidermis
of skin, Xanthoblast in amphibian and others depending on color, pigmented, and
Iridoblast.
 53

b. Ventral migration: cells of ventral migration give rise to number of

structures.namely
1. Sensory ganglia of mixed nerves(5,7,9,10). 5th Trigeminal ganglia/semilunar
ganglia. 7th geniculate ganglia. 9th otic ganglia.
2. Dorsal root ganglia of spinal cord.
3. All sympathetic ganglion – thoracic and lumbar, stellate and cranial cervical
ganglions.
4. Parasympathetic ganglion- ciliary ganglion, myenteric plexus, meissner’s plexus.
5. Some of neuroglia cells- satellite cells surrounding pseudounipolar neurons
(amphicytes).
6. Microglia
7. Schwann cells derived from neural crest in PNS (axon of nerve surrounded by
myelin sheath). In CNS, axon surrounded oligodendroglia are derived from
Neuroblasts.
8. Adrenal medulla: chromaffin cells gives chromaffin reaction when tissue fixed in
chromium salt(zenker’s/helly’s fixative) stain reveals brownish colored pigment.
Eg.adrenal medulla cells. It is called as APUD reaction ie cells of APUD series ie
aminao peptidase uptake and decarboxylation series.
9. Dentine or teeth.

Anamolies during development of Nervous system:

Cerebellar dysplasia and atrophy: during rapid cerebellar growth and differentiation of
external germinal layer brings about hyperplasia and degeneration of perkinje cells.
Dysraphiasis: result from failure of normal closure of neural tube.
a. Cranial bifida:involve cleft in neurocranium. Here meninges along with neural tube
coming out. It could be meningomyocoel along with meninges or meningo
encephalocoele- both brain and meninges.

b. Spinal bifida: part of spinal cord seen outside where arches of two verterbrae not
completely fused. It could be Aculta ie closed by skin, not seen outside but condition
is there. In spina bifida it coming out of skin.

c. Syringomyelia: abnormal cavitation of spinal cord. Extending over several segment. It

is due to incomplete aclusion of central cavity in alar plate region.
d. Anencephaly: part of brain or whole brain may be absent.

Hydrocephalus: too much accumulation of CSF in ventricles. Ventricles become large and
brain tissue become thin called Hydocephalus. It is due to excessive CSF production or
impaired CSF removal from subarachnoid space ie blockage in subarachnoid cisternae.
Hydroencephaly: cerebral hemisphere replaced by two fluid filled sac. Rest of brain is
normal may be due to vascular defect affectiong migratory neuroblasts. Nutritional deficiency
or viral infection like BT in ruminant and panleukopenia in cat cause this.
 54

ENDOCRINE GLANDS
HYPOPHYSIS : Pituitary gland
It derived from two sources namely,
1. Stomodeum ectoderm→ pars anterior
2. Ventral portion of diencephalon→pars nervosa
The ectoderm come in contact with endoderm is bucco pharyngeal membrane. Area infront of
buccopharyngeal membrane is Stomodeum which is lined by ectoderm of body wall. Later
stomodium become mouth cavity. From ectoderm of stomodeum dorsal out growth arises
which is called as Rathke’s pouch/Hypophysial recess. Meanwhile from diencephalon
(portion of neural tube) another ventral outgrowth arises which forms the
Infundibulum/Infundibular recess. Both are ectoderm but develop from two sources ie
ectoderm of diencephalon and stomodeum. It occurs at 3 week old embryo.
By 2nd month, Rathke’s pouch looses connection with the stomodeum and comes and lies
infront of Infundibular recess of diencephalon. Rathke’s pouch has double wall. Cranial part
give rise to Pars anterior or pars distalis. Posterior part form Pars intermedia. Pars
anterior has thicker wall compared to pars intermedia.
Infundibular recess has 3 parts namely,
1. Pars nervosa
2. Infundibular stalk
3. Median eminence.
Portion of pars intermedia surrounded by cells called Pars tuberalis.
Adenohypophysis is gland like. It includes Pars anterior/distalis, pars intermedia and pars
tuberalis.
Neurohypophysis has pars nervosa, infundibulum and median eminence.
Rathke’s pouch arises as a vesicle or pouch. Cranial wall is thicker than intermedia and there
is residual lumen which disappear in some(man, ox) and remains in some(dog, pig, cat,
horse). Before disappearance of rathke’s pouch, portion of Pars intermedia forms cone like
structure calles as Cone of Wulzen. Seen in buffalo, sheep and goat mainly made up of
basophiclic cells. Later this cone disappears.
Pars anterior made up of 2 types of cells like chromophobes which has no affinity for color,
resting stage transferred to acidophil or basophil. Chromophils are acidophils and basophils.
Pars intermedia is made up of cells resembling basophils.
Rathke’s pouch looses connection with stomodial ectoderm but some time connection is
persisting. This condition is called as persistent of craniopharyngeal canal. This is
surrounded by bones of skull (sella turcica) so small canal like present. Sometime it form a
tumour called craniopharyngeoma.
Structure arises just behind Rathke’s pouch as out growth of stomodeum is called as Seessels
pocket in the dorsal direction (in chick and pig). Named after American embryologist Albert
Seessels(1910). It gradually disappear so no importance.
 55

ADRENAL GLAND: is derived from 2 sources. It has cortex and medulla. Cortical tissue
derived from mesoderm (mesonephros) and medulla derived from ectoderm of Neural crest.
Cortex develop at 10mm stage (37 days in human) from columnar mesodermal cell situated
in abdominal wall. These cell proliferate and form cords which separate into large cell mass
on either side of dorsal aorta. First group of cells are primary/foetal cortex. This primary
cortex completely disappears and these cells later replaced second group of cells from the
surrounding mesoderm and to form the structure secondary/adult cortex or Definative
cortical cell, which is invaded by neural crest cells to form the medullary tissue.cells of
neural crest gets differentiate into chromophils and chromophobes.
(they derive from sympathetic nerve cell from neural cortex). It is at 16mm stage (44 days),
these cell shows evidence of catacholamines (10th week of foetal life).
(foetal supra renal are large at 4th larger than kidney. 20 times relative size in the adult. Great
size is due to acidophilic cortical cell. Foetal cortx undergo involution, net loss of weight).
Adult cortex later differentiate into 3 parts namely, Zona glomerulosa, Zona fasiculata and
Zona reticularis.

EPIPHYSIS/Pineal gland: pineal gland appear early in development as invagination from

roof of diencephalon. This outgrowth extend upward and forward and remain as hallow.
Later become solid by proliferation of cells of wall and neuroglia cell ie connected to
diencephalon. Nerve fiber from epithalamus to stalk of pineal. So ectoderm in origin.

SENSE ORGANS
SENSE OF OLFACTION:
Region of head infront of telencephalon, ectoderm shows 2 thickenings called as Olfactory
Placode.(placode-means thickening). Usually ectoderm give rise to sense organs. From
Placode, get a depression called Olfactory Pit. Which come in contact with brain
vesicle(neural tube) ie Telencephalon. Cells of Olfactory pit give rise to two types of cells.
namely, Olfactory cells/neuroepithelial cell. And Supporting cells.
Olfactory cells are bipolar cells. Cerebral portions show Olfactory hairs. Axons of bipolar
cell pass into underline CT and from a bundle of nerve called Fila Olfactoria. These go and
join olfactory lobe of Telencephalon by passing through the cribriform plate of ethamoid.

SENSE OF VISION/EYE:
From diencephalon of brain vesicle two out pocketing develops laterally. This is called as
Optic vesicle (33hrs in chick embryo). Each optic vesicle undergoes a invagination to give
rise to cup like structure called as Optic cup. Optic cup has outer layer(thinner) and inner
layer(thicker). Outer wall which is thinner becomes pigmented layer of retina which forms 1 st
layer. Inner thicker wall gives rise to Nervous layer of retina(other 9 layers). Ganglionic cell
layer develops on mantle layer(by neuroblast) and rods and cone cells. B/w outer and inner
wall there is a space which is called as Intraretinal space which later completely disappears.
Connection between diencephalon and optic cup form Optic stalk which later give rise to
Optic nerve.
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Optic cup has two portion. Cranial and caudal portion. Cranial part is thinner wall compared
to caudal. Thicker portion become Pars Optica Retinae. Cranial thinner portion forms Pars
ciliaris retinae and Pars iridica retinae. Both these combined called as Pars Caeca retinae.
Immediately infront of optic vesicle is ectoderm of body wall which undergoes a
thickening which is called Lens Placode. Lens placode undergoes invagination inside optic
cup which form Lens vesicle. Lens vesicle forms two walls. It loses connection with
ectoderm. It has cranial thinner wall made up of single layer of cuboidal cell forming Lens
epithelium. Caudal thicker wall has fibers arrange in a radiation manner called as Lens
fibers. Space in between lens epithelium and lens fibers disappears completely. Lens vesicle
migrates and gets lodged between iris and ciliary body. Whole thing is surrounded by
mesoderm which gives rise to outer Sclera and middle Choroid. Mesodermal cells migrate
inside between the lens and optic stalk forms Vitreous body. Other portion iris and ciliary
body derived from mesoderm. (only pigmented layer of iris and ciliary body derived from
Neural ectoderm).
Outer epithelial layer of optic cup in addition to pigmented layer of retina in Pars
optica retinae, it give rise to smooth muscle cell in iris. These are dilator and constrictor
muscle of the iris. All muscles of body derived from mesoderm except these two(ectoderm).
Rest of ectodermal body wall, after giving lens, infront of optic vesicle, arises
Corneal vesicle/cornea. Cornea has 5 layers like,
1. Corneal epithelium: derived from ectoderm of body wall.
2. Other 4 CT layer derived from mesoderm between iris and corneal epithelium.
Stroma and inner epithelium derive from surrounding mesenchymal tissue.
From surface ectoderm of body wall, infront of cornea, two outgrowth arises. These
give rise to Eyelids. First two eyelids are join with one another. And later get separated.
Optic cup at ventral aspect has gap/fissure called as Optic fissure which later become
Choroid fissure. Into choroid fissure runs an artery called Hyaloid artery (central artery of
retina). Later this gap should get closed. In not closed, leads to a abnormality. Mesodermal
cell surrounding optic vesicle give rise to muscles of eyeball.
Aqueous humor: present between iris and cornea in anterior and posterior chamber. It is
secreted by posterior chamber by epithelial cells of ciliary body. These get accumulated in
both anterior and posterior chamber.
(at j/n b/w cornea and iris, irido corneal junction, there is mesh work of blood vessels which
drain the excess aqueous humor accumulation in two chambers.)
Attachment between two eyelids is called Ankyloblepharon. This condition is lost just before
birth in ungulates. In carnivores persist for 10-15 days even after birth.
Lacrimal gland is ectodermal origin develop as an out growth in the form of buds from upper
fornix conjunctiva.

Anomalies of Eye:
Microophthalmia: associated with abnormality in development of vitreous body.
Two eyelid should get separated during separation if eyelid protruds out more than normal is
called Ectopion.
If protrusion of eyelid is not sufficient, turn inward called Entropion.
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Persistant choroid fissure leads to Coloboma. It may involve any part of eye. Sometimes
involves iris or lens also. Coloboma iris.
Anophthalmos: absence of eyeball.
Sometime two eye close to give Cyclops condition.
Glaucoma: abnormal development of anterior chamber ie there is increase accumulation of
aqueous humor.

DEVELOPMENT OF EAR
Ear has 3 parts.
1. External ear - ectoderm in origin
2. Middle ear – endoderm
3. Internal ear – ectoderm
Auditory vesicles – endoderm + mesoderm.

INTERNAL EAR:
Lateral aspect of ectoderm of Rhombencephalon shows two lateral thickenings which
is called as Auditory placode. Appears by 3rd week. Auditory placode become invaginated to
Auditory pit and invagination become deeper and giving rise to Auditory vesicle or
Otocyst/otic visicle. This vesicle get separated from ectoderm. It is oval in shape. near
auditory vesicle, there is ganglion found ie Ganglia of 7th and 8th nerve from neural crest.
Medial aspect of Auditory vesicle shows another small outpocketing which give rise to
Endolymphatic duct. Auditory vesicle become elongated with dorsal broader part and
ventral narrow part.
From Dorsal broader part arises 3 Semicircular canals and middle Utricle and Sacule ie
Vestibule. In b/w utricle and sacule there is utriculosacular duct. Medial aspect of this
shows endolymphatic duct.
Ventral narrow part becomes coiled and give rise to Cochlea duct. Connection between
cochlear duct and sacule is called Ductus reunions.
Membranous labyrinth is formed by ectodermal body wall. It is surrounded by osseous
labarynth by mesodermal tissue.
Each semicircular duct has enlargement called Ampulla. Cells lining ampula give rise to
Cristae ampularis which is specialized area for equilibrium for rotatory acceleration.
Utricle and Sacule contain specialized area called as Maculae.
Cristae and maculae lined by neuroepithelial cell terminate in a ganglia of 8th cranial nerve ie
vestibular ganglion.
In Cochlear duct, spiral organ of Corti lined by neuroepithelial cell and supporting cell. Axon
of cell joined to form spiral ganglion.
Vestibular ganglion and spiral ganglion axons joined to form Vestibulocochlear nerve.
Note: membranous portion – surface ectoderm and Osseous portion – mesoderm.
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MIDDLE EAR:
1ST Pharyngeal pouch become enlarged with proximal narrow and distal broader part. Narrow
part form Tubotympanic recess or Eustasian tube (connection b/w middle ear cavity and
pharynx).
Broader distal part gives middle ear cavity/tympanic cavity.
1st Pharyngeal cleft-external auditory canal. At this point ectoderm is in contact with
endoderm. This forms the Tympanic membrane(externally ectoderm, internally endoderm
b/w fibrous tissue from mesoderm).
B/W ectoderm and endoderm, mesodermal tissue is present which give rise to structure of
middle ear like Auditory ossicles (maleus, incus, stapes) and muscles(tensor tympani,
stapedius). Maleus and incus are derived from 1st pharyngeal arch where as Stapes from 2nd
pharyngeal arch. Tensor tympani muscle is attached to the maleus so nerve of 1st arch ie
trigeminal nerve innervate tensor tympani. Stapedium muscle derived from 2nd or hyoid arch
and stapes derived from 2nd arch so innervate by branch of Facial nerve.
EXTERNAL EAR:
Derived from 1st pharyngeal cleft. It give rise to bony part and cartilaginous part of external
ear.
Pinna: from 1st and 2nd arch, arises 3 hillock from each arch so totally 6 hillocks. Later
rearrangement of hillocks to give rise to parts of external ear.
Parts are helix, antihelix, tragus, antitragus.
Helix is derived from 2nd and 3rd hillock
Antihelix – 4th and 5th
Tragus – 1st
Lola – 6th
2nd ,3rd , 4th , 5th cleft disappear completely. If cleft will be persistent after birth, like opening
are called Cervical sinus or cervical cysts.

DEVELOPMENT OF FACE AND PALATE

Portion of head infront of stromodeum shows Olfactory pit. Olfactory pit is
surrounded by two prominent ridges ie medial and lateral nasal process. Inside, 1st arch is
divided into maxillary and mandibular process.
Fronto nasal process is formed by proliferation of mesoderm ventral to brain
vesicles and it form upper boundary of stomodium (stomatodaeum). On either side of fronto
nasal prominence is olfactory pit. (olfactory pit is surrounded by 2 ridges called medial and
lateral nasal process). This takes place by 5th week. Stomatodeum, 1st pharyngeal arch and
fronto nasal process gives rise to face.
During 7th week, maxillary process (dorsal divison of 1st arch, mandicular process is
ventral portion) continuous to increase in size and closed in medial direction. During this
time two medial nasal process becomes closer to one another. So there is a cleft between
medial nasal process and maxillary process. Fusion of two medial nasal process goes to form
the upper lip and medial alae of the nostril.
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Lateral alae of nostril is formed by lateral nasal processes. It does not contribute to
upper lip. As the size of maxillary process or swelling increases and touches the optic vesicle.
During this process, a deep furrow develops between two lateral nasal swelling and maxillary
swelling. This goes to form Naso lacrimal duct. Upper portion form nasolacrimal sac.
Fusion of two medial nasal process which are not complete form Philtrum.
Lower lip and lower jaw is formed by fusion of extended mandibular process. Nasal
septum is formed by proliferation of mesodermal cell at midline of fronto nasal process.

Development of Palate: as the two maxillary process grow medially, two medial nasal
process are not only seen on outside but also develops deeply. These process are called Inter
maxillary segment. It has 3 parts.
1. Labial component – see from outside – upper lip.
2. Upper jaw component/jinjual component – form gums of teeth.
3. Palatine component- roof of mouth cavity or hard palate. This is called as Primary
palate.
Secondary palate is formed by two shelf like out growth from maxillary swellings called
Palatine shelves. Appear by about 6th week. Palatine shelves located downwards on either
side of tongue.
By 7th week, palatine shelves go up/assend and occupies a horizontal portion which goes to
form secondary palate or Definative palate. This secondary palate unites in Nasal septum. At
this point primary and secondary palate fusion occurs. This triangular area is called Foramen
Incisurus. This give rise to Hard palate.
Note: at fusion of secondary palate, tongue like structure called Uvula in man which is
portion of soft palate.
Anamolies:
Unilateral cleft lip: there is a gap between maxillary process and mandibular nasal process on
one side.
Bilateral cleft lip: gap on both side.
Oblique facial cleft: non fusion of the optic vesicle, nose and upper lip.
Cleft palate: two secondary palate do not meet. So wide gap at root of mouth. It usually
associated with cleft lip.

DEVELOPMENT OF MAMMARY GLAND

It is a modified sweat gland.
A pair of external thickenings on the ventral aspect of body wall found which are called
Mammary ridges or lines (milk lines) found bilaterally. It extends from pectoral region ie
base of forelimb bud to medial aspect of hindlimb bud.(ie pectoral to inguinal region). These
ridges grow inside ie invagination and get canalized to form duct system.
This primordial consists of superficial layer of squamous, deeper layer of cuboidal cell.
Thickening penetrate mesenchyme underlying and gives origin to 16-24 secondary branches
which form duct system.
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During 8th and 9th month, initial down growth become canalize forming duct system.
Secondary out growth become lactiferous duct. Tertiary form alveoli and small duct of
gland. So original thickening form Nipple or Papilla or teat.
Duct system is under control of oestrogen and alveoli is under progesterone.
In carnivore and pig→ develops on both side, 5 pairs.
Primates & elephants → only cranial ie pectoral ones and caudal disappears.
ungulates→ only caudal one persists and cranial disappears.
Witches milk: new born baby secreting milk due to hypertrophy of gland and secretory
activity due to production of prolactin by pituitary gland in infants before or after birth in
both sexes due to large quantity of estrogen.
Anamolies:
Additional teat development→ polythelia.
amastia→ absence of development of mammary gland.
polymastia→ extra mammary glands.
Gynaecomastia→ male also show enlargement of mammary gland.

DEVELOPMENT OF SALIVARY GLAND

Salivary gland develop from ectoderm of stomodium. Thickening of stomodial ectoderm
invaginate and give rise to solid cords of cells. Later these cells form(acquired) a lumen and
form the secretory acini and ducts. This is surrounded by mesodermal cell which form
capsule and interlobular connective tissue.
Parotid gland: first appear at angle of mouth in the form of solid cords. These grows away
from labial groove, becomes elongated and separates from the epithelium. Secretory portion
of gland is located at the region of the ear.
Primordial of sublingual and mandibular gland arise at labio-jinjual groove between lower
jaw and tongue. on either side and at floor of buccal cavity mandibular gland migrate and get
separated from mouth. So duct of mandibular and sublingual gland open simultaneously at
floor of mouth.

DEVELOPMENT OF TEETH
Teeth is ectodermal and mesodermal in origin.
Earliest development of teeth is the appearance of dental groove. ectoderm undergoes
invagination forming groove. this groove elongates and assembles a cap like structure which
is called as Dental lamina. Later further invagination of outerlayer forming bell shape which
is called as Enamel organ. (bell shape structure surround by mesoderm tissue).(connection
between enamel organ and ectoderm is called as Dental lamina). Underlying mesodermal
tissue around dental lamina is called Dental papilla.
Enamel organ has 3 layers:
1. Outer enamel epithelium→ lined by squamous cell.
2. Inner enamel epithelium→ lined by cuboidal cell.
3. Stellate reticuleum→ in between 1 and 2nd.
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Cells of inner epithelium become taller and become columnar cells. These are called as
Ameloblast. Meanwhile outer enamel epithelium and stellate reticulum completely
disappear. Peripheral cells of dental papilla arrange in the form of rope of cells which are
called as Odentoblast. Ameloblast are tall columnar in shape start secreting enamel
substance. It arises as a cuticular secretion from ends of ameloblast. Then it laying down
enamel prisms.
(In between prism inter prismatic substance which are secreted by ameloblast). Mean while
odentoblast start secreting organic matrix of dentin ie Pre dentin. Later calcium salt deposited
on predentin forming dentin.
Mesoderm surrounding root of developing tooth become condensed and hightly vascular to
form Dental sac. Elements of dental sac are Cementoblast (which deposit in the cytoplasm,
matrix of cement and get ossified by deposition of calcium salt).
Dental papilla form the Dental pulp occupying root canal of tooth.

DEVELOPMENT OF SKIN
Ectoderm(epidermis) and mesoderm(form dermis) together form the skin.
Epidermis: formed by single layer of ectodermal cells which differentiate into
I. Inner germinative layer →which proliferate and differentiate to form
1. stratum basale,
2. stratum spinosum
3. stratum granulosum
4. stratum lucidum &
5. stratum cornium.
II. Outer layer→ shed off and mix with secretion of sebaceous gland to form pasty
substance called Vernex caseosa.
Neural crest migrate into basal layer of epidermis and differentiate into melanoblast cell at 3rd
month of gestation.
Dermis: mesoderm from dermatome of paraxial mesoderm (somite) gives dermis. From
dermis numerous projections develop called as Dermal papilla. Layer below this dermis is
hypodermis.

DEVELOPMENT OF HAIR:
Germinative layer of epidermis(ectoderm) develop a down growth into dermis and its lower
end expand to form hair bulb. Bulb invaginated by mesoderm in the form of projection
called as papilla. Germinal cells on papilla become spindle shape and get keratinized to form
body of the hair.
Root sheath is formed by cells remains at the parietal layer of downgrowth close to the bulb.
Sebaceous gland: from lateral wall of epidermal outgrowth sebaceous gland develop as a
sacculation and then proliferation and degeneration of germinative cell form gland.

Sweat gland: develop as a downgrowth of ectodermal layer of skin as a cord, then cord
become canalized and extend upto dermis and distal end coiled to form gland which open on
the surface at later stage of intrauterine life.
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Hoof: develop at midterm pregnancy and develops at the tip of digit. It is developed in the
stratum lucidum of epidermis. First epidermis become keratinesed at tip and extend upto 3rd
phalnx. Germinal layer of root of hoof proliferate and form a matrix and matrix transformed
into hoof structure.