DEVELOPMENT OF

FROG
DEVELOPMENT OF
FROG
Kingdom: Animalia
Phylum: Chordata
Sub-phylum: Vertebrata
Order: Anura

Frog is the representative vertebrate having both aquatic and terrestrial mode of
life.
Embryology of frog typically illustrates the sequence of developmental events of
all higher vertebrates.
The green pond frog, Indian Bull frog are the most common species found in India.
In frog sexes are separate and sexual dimorphism is well marked during breeding
season.
Males posses copulatory or nuptial pad and possess vocal sacs which produce
croaking sound during breeding season to attract females for copulation.
GONADS
The male gonads are the testis which are paired, white and ovoid bodies located in the coelomic
cavity near the upper end of the kidneys and are suspended from the dorsal body wall by a
sheath of peritoneal mesentery called mesorchium.
Female gonads are the paired ovaries, attached to the dorsal body wall by a peritoneal sheath,
the mesovarium. During breeding season they assume large size due to storage of mature eggs
and after ovulation they become small sized.
SPAWNING
Act of expelling eggs from the uteri of female frog.
During copulation the male firmly clasps the body of
the female by it’s forelegs and enlarged thumb pads,
this sexual embrace is called amplexus.
Jelly coat of liberated eggs swells up and forms a
froathy, irregular mass-Spawn and its formation is
spawning
In such position, the male is always capable of discharging sperm over the ova as they are
extruded through the cloaca of female.
SPERM
It has a head, middle piece and a tail.
Head is elongated containing nucleus and the acrosome at the anterior end.
Middle piece is short.
Tail is long and filamentous.
Size: 0.03mm in length.
OVUM
Spherical in shape, mesolecithal and moderately telolecithal.
1.75-2mm in diameter.
Egg surrounded by a thin transparent vitelline membrane just over the plasma
membrane, and a jelly coat, formed of 3 concentric layers of albuminous substance.
Jelly coat protects the eggs from mechanical injury and serves to make the eggs adhere
to each other and to submerged objects. Thus they are not readily washed out.
Donot allow bacteria and fungi to creep in
It is distasteful and prevents the eggs from eaten by aquatic organisms.(TYPE STUDY)
FERTILIZATION
It is external and takes place in water.
Egg is laid in secondary oocyte stage( second meiotic division metaphase stage).
Spermic fluid is shed almost simultaneously with ovulation.
Gametic fusion occurs at once. Usually monospermy occurs. When egg is activated
by sperm second polar body is released.
The entry of sperm usually occurs in the animal hemisphere.
The point of contact of sperm head with cytoplasm of the egg bulges out in the
form of cone called fertilization cone through which the sperm enters. It gradually
engulfs the sperm and then begins to retract and carry the sperm.
Within 1-2 min the whole spermatozoan enter inside ovum and the tail
disintegrates.
When sperm enters cortical granules break and lifts up the vitelline membrane and
forms fertilization membrane.
Between fertilization membrane and plasma membrane is the perivitelline space
Because of yolk granules in the vegetal pole the egg rotates slowly within the
perivitelline space that results in the animal hemisphere above and vegetal
hemisphere below. Such a rotation is called rotation of orientation.
Point of sperm entry will mark the ventral side of the embryo, while the site
180 degrees opposite the point of sperm entry will make the dorsal side.
Nieuwkoop centre is a signalling center in the dorsal side of early amphibian
embryo.
It forms in vegetal region as a result of cortical rotation initiate by the sperm
entry. It induces Spemann’s organizer.
The activation of the egg by the entry of the sperm induce the second
maturation division of egg nucleus which was suspended in metaphase 2.
It gives off second polar body at the animal pole and the nucleus becomes
vesicular to become egg pronucleus.
PENETRATION AND COPULATION PATH
Usually the whole sperm enters the ovum, the tail disintegrates and the head
and middle piece travel towards the female pronucleus along a cone shaped
path. This path corresponds to the radius of the egg and is referred to as the
penetration path.
In some cases the sperm does not move straight and changes its course after
penetration. This changed path is copulation path.
Since the amphibian eggs are heavily pigmented, the sperm takes some of
the pigment granules of the surface layer to the interior of the egg. Therefore
the course of the sperm head may be marked by the pigment granules
trailing along its path.
AMPHIMIXIS
The sperm nucleus becomes vesicular in the egg cytoplasm and forms male
pronucleus. It moves towards the egg pronucleus resulting in amphimixis,
which produces a diploid zygotic nucleus.
• ESTABLISHMENT OF BILATERAL SYMMETRY AND FORMATION OF GRAY
CRESCENT.
Before fertilization the frog egg is radially symmetrical but after fertilization it
becomes bilaterally symmetrical. This is due to temporary shifting of pigments
resulting from a raising and concentration of cortical layer towards the animal
pole. This is accompanied by corresponding thinning of the egg cortex opposite
of the egg.
At future ventral side of the egg the cortical cytoplasm moves downward
towards the vegetal pole while at the opposite, future dorsal side of the egg the
cortical cytoplasm moves upward toward the animal pole.
When the edge of the darkly pigmented cortical cytoplasm rises above the
equator on the dorsal side, the deeper lying lightly coloured marginal
cytoplasm is more exposed, thereby producing a crescent shaped surface
known as the gray crecent.
Gray crescent is a lightly pigmented crescent shaped area of marginal
cytoplasm on dorsal side, exactly opposite the point of sperm entry .
It is formed in the equatorial region just at the junction between the heavily
pigmented animal hemisphere and lighter vegetal hemisphere.
The plane passing through gray crescent and animal pole defines the median
plane of bilateral symmetry and it coincides with the embryonic axis of future
embryo.
Zygote can be divided into 2 similar halves through middle of gray crescent
and the poles.
Usually the first cleavage plane coincides with median axis.
The gray crescent represents the future site of formation of the dorsal lip of
blastopore.
The gray crescentic region will become the posterior side and the opposite
side, the anterior side of the future embryo.
CLEAVAGE
Egg is mesolecithal and moderately telolecithal.
Holoblastic cleavage.
First cleavage occurs about 2.5 hrs after fertilization and completes in 30 min.
First cleavage is in meridional plane, and the furrow from animal pole to the
vegetal pole through the middle of the gray crescent. This makes two equal
sized blastomeres which are identical in quantity and quality of yolk and
pigment granules.
Second cleavage is also meridional but perpendicular to the first one. It
produces blastomeres which are quantitatively different because 2 of them
contains gray crescent material.
Third cleavage is latitudinal and perpendicular to both cleavages. The cleavage
furrow is slightly above the equator towards the animal pole due to less yolk in
that region. It produces macromeres and micromeres.
Fourth cleavage is meridional and double, comprises of 2 cleavge furrows laid
down simultaneously and at right angles to each other and to 3rd furrow. It
results in 16 blastomeres of which 8 are micromeres and 8 are macromeres.
Fifth cleavage is latitudinal and double which divides macromeres and
macromeres. As a result 32 blastomeres.(4 tiers of 2 macromeres and
micromeres)
From this the regularity of cleavage is lost and micromeres divide faster than
macromeres.
An amphibian embryo containing 16 to 64 cells is commonly called a morula.
At 128 cell stage blastocoel becomes apparent and embryo is considered
blastula.
BLASTULATION
Blastomeres are usually assume spherical shape but their surfaces of contact remain
flattened.
After morula stage blastocoel develops. 16-64 cell stage is commonly called morula. In a frog
as early as 8 cell stage a small central fluid filled space called segmentation cavity or
blastocoel appears.
As the multiplication of micromeres is rapid, the blastocoel shifts more towards the animal
pole and is an eccentric position above the level of equator.
The blastocoel becomes infiltered by water and albuminous fluid.
The roof of the blastocoel is covered by a relatively thin 2 or more layers of pigmented black
micromeres and its floor is more or less flat and formed of non pigmented large sized
macromeres.
As cleavage proceeds, the adhesion of blastomeres to each other increases and they arrange
themselves into blastoderm. It is many cell thick in frog.
This is blastula and is coeloblastula.
The micromeres of animal pole tend to spread towards the vegetal pole and consequently
there appears an equatorial thickening called marginal zone or germ ring.
 MBT- mid blastula transition
• Prior to gastrulation the zygotic genomes have to be activated.
• Nuclear genomes are not activated until 12th cell cycle. At that time mid
blastula transition occurs.
• Different genes begin to be transcribed in different cell, the cell cycle acquires
gap phases, blastomeres acquire capacity to become motile.
 GASTRULATION
Formation of blastula into gastrula.
Blastomeres move to definite positions and form germ layers by morphogenetic
movements. Ultimately it forms triploblastic embryo.
Little bit complicated gastrulation occurs.
Several processes occur simultaneously during gastrulation. It involves mainly 3
processes:
1. Invagination of endoderm
2. Involution of pharyngeal endoderm and chordamesoderm
3. Epiboly of ectoderm
INVAGINATION OF ENDODERM AND FORMATION OF BLASTOPORE
 Gastrulation begins by the formation of groove on the dorsal side below the equator
and exactly below middorsal point of the gray crescent. This groove is the blastopore.
 Blastopore is produced by the invagination of endodermal material.
 The cells of this region assume the elongate shape of a bottle and move toward
the interior of the embryo toward the blastocoel cavity.
 Their elongating neck cells remain to surface of blastula, same time bulky cell
bodies move inward.
 As a result a pull is exerted along their thin necks which creates an indentation at
the surface.
 When the bottle cells multiplies the invagination deepens.
 The area above blastopore is the dorsal lip of blastopore which is derived from
gray crescent.
 on the dorsal side of the embryo, a cavity is formed which extends from the
blastopore on the surface into interior of the embryo. Lined on all sides by the
invaginated cells, this cavity is called archenteron or gastrocoel. It is first narrow
slit like cavity, later as more cells invaginate, it expands at its anterior end and
becomes crescentic, then horseshoe shaped and finally circular.
 Materials bounding the blastopore on sides represents its lateral lips and the
material on the ventral side, the ventral lip.
INVOLUTION OF PHARYNGEAL ENDODERM AND CHORDAMESODERM
 After blastopore formation convergence and involution occurs.
 Convergence: it is the flow of surface materials towards the blastoporal lips
 Involution :rolling up of that materials which have arrived at the dorsal lip of
blastopore.
 The endoderm cells adjoining the dorsal lip of blastopore are first to migrate
inward- they form the prospective pharyngeal endoderm.
 These cells are followed by prechordal plate, notochord and tail mesoderm.
 As these materials flow into the interior around dorsal lip, they become
considerably narrowed and in the process of inward flow, greatly elongated.
 The prospective pharyngeal region later forms the most anterior part of
advancing archenteron.
 The oral and pharyngeal endoderm expands so as to form the spacious foregut,
whose lateral, ventral and anterior walls consists thin layer of endoderm.
 Inside the gastrula, prechordal plate becomes a part of the archenteron roof in
front of the notochordal material.
 The notochord becomes stretched along the dorsal side of the archenteron
and forms a middorsal strip of the archenteron roof. Prospective notochord
elongates in the longitudinal direction and contraction in transverse direction.
 The tail mesoderm remains near the blastopore and is the posterior end of the
embryo.
 The trunk somites and ventrolateral mesoderm into the interior by rolling over
the ventral lips of blastopore.
 The entire mesoderm moves in anterior end from blastoporal end and is called
chordamesodermal mantle between ectoderm and endoderm except where
mouth will be formed.
EPIBOLY OF ECTODERM
 It is the movement of ectodermal cells over the surface of the embryo.
 When the prospective endoderm and chordamesoderm move inside from the
surface, their places are taken by ectoderm.
 Neural and epidermal ectoderm increase their surface areas at the expense of
thinning out of epithelial layer by active migration of ectodermal cells towards
blastopore margin.
 The prospective epidermis expands in all directions but prospective nervous
system expands longitudinally.
 Dorsal lips forms lateral lips by the cells rolling in increases laterally.
 Laterals meet ventrally to form ventral lip and blastopore assumes a shape of
ring. The space enclosed by the rim of blastopore is seen to be filled by yolk
laden macromeres of vegetal region. These cells are called the yolkplug.
 These cells are seen protruding from blastopore as they are too large to be co-
operated in the floor. The embryo is in yolk plug stage.
CLOSING THE BLASTOPORE
 As a result of combines action of epiboly and convergence, the blastopore
reduces in size and yolk plug is drawn inside to archenteron and blastopore closes
ultimately.`
 Archenteron deepens and blastocoel is eliminates.
 All this shifts the gravitational axis of blastula, shifting it about 90 degree away
from blastopore.
 Vegetal region is drawn in to the interior and at the same time to rotate and at
the end of gastrulation the vegetal region comes to lie below archenteron.
 Finally gastrula contain 3 germ layers.outer layer- neuroectodermal portion
middorsally and epidermal ectoderm around the embryo.
 Archenteron- lumen of the gut. At this stage the walls and floor of archenteron
are composed of endoderm, roof is of chordamesoderm.
 Gradually the free edges of the endoderm approach each other in mid line and
fuse, thus closing mid gut.
NEURULATION
 It is the formation of neural tube from neural ectoderm.
 It occurs after gastrulation.
 After the closure of blastopore the neurectoderm situated along the middorsal
position of fully formed gastrula is called neural plate or medullary plate. This extends
from dorsal lip of blastopore to anterior end. The cells of neural plate become thick
and elongated and arranged in columnar epithelium.
 The lateral margins of neural plate become thickened and elevated as neural folds or
medullary folds. Anterior end of neural plate also folds transversely.
 A depression or groove is formed between folds in the centre of neural plate and is
called neural groove.
 Gradually the neural folds grow inward towards one another and fuse in the middle
line to form a tubular structure called neural tube.
 The cavity of neural tube is neurocoel.
 Closing of neural tube begins just in front of mid region and progresses back and forth.
 Neural fold remain unfused in the anterior end and is called neuropore. But it
closes ultimately.
 Posterior end of neural fold merge into the sides of blastopore and their union
encloses blastopore.
 Now neurocoel communicates with the archenteron through the blastopore.
The blastopore acts as a temporary passage or channel between the
neurocoel and archenteron called neurenteric canal.
 As the neural folds fuse, the neural tube becomes constricted off from the
dorsal ectoderm. Now the surface of ectoderm becomes a continuous sheet of
cells above neural tube.
 Neurocoel is broadest at the anterior part and it develops as the brain and
posterior part is narrower.
 The formation and closure of neural plate is known as neurulation and the
embryo at the time of neural tube formation is called neurula stage.
 Later neurenteric canal is interrupted and the blastopore fuse. After the
closure of blastopore a new opening appears slightly below the position of the
original blastopore and it becomes anus in future.
NEURAL CREST
 A group of cells on the sides of the neural plate are not incorporated in the
neural tube or epidermis.
 When neural tube separates from the epidermal ectoderm these cells occupy
the space between the neural tube and overlying epidermis as a semicircular
peripheral band around the neural tube called neural crest.
 It is seen as two patches on either sides of neural tube.
 This gives rise to branchial skeleton, dorsal root ganglia, sensory nerves,
sympathetic nervous system and cells of adrenal medulla.
CHORDAGENESIS
• Notochord develops from the chorda mesoderm.
• The presumptive notochordal cells along with mesoderm involute and stretch out along the
longitudinal axis as a narrow sheet called chorda mesodermal mantle in the mid dorsal
region of the roof of archenteron.
• By the time the blastopore closes, the chordamesoderm mantle separates from the
adjoining endoderm and comes to lie between endoderm and ectoderm in its definitive
position.
• Later chorda- mesodermal mantle delaminates into the prechordal plate in the middle and
somatic mesoderm on the sides.
• Pre chordal plate cells very quickly expand and arrange themselves into a cylindrical rod.
• Later these assume vacuolated appearance and an elastic and fibrous sheath is developed
around these cells called notochordal sheath.
• As a result the rod becomes turgid and represents the notochord.
• Its lengthening occurs by the swelling of notochordal cells with in the space provided by the
notochordal sheath, which allows them to slide and interdigitate in long axis of sheath but
not in other planes.
MESODERM AND COELOM FORMATION
 The embryo elongates during coelom formation.
 The presumptive mesoderm is located on either side of the notochord, they grow
laterally and then downwards between the ectoderm and endoderm until they
meet ventrally below the gut.
 The mesoderm becomes subdivided into a series of segments called somites
called somites.
 Each somite can be divided into 3 parts:
EPIMERE or DORSAL MESODERM
• Lie on either side of notochord and each mesodermal band becomes transversely
constricted into a series of segments called somites.
• Outer layer of somite is thinner and is dermatome, it develops into the derivatives
of the dermis.
• Inner layer is thicker called myotome, it develops in skeletal muscles of back.
•
INTERMEDIATE MESODERM or NEPHROTOME
• It is the mesoderm lying on either side of the somites also called nephrotome.
• The kidney develops from this.
VENTRAL AND LATERAL PLATE MESODERM or HYPOMERE
• Lying lateral to the nephrotome.
• It is separated into 2 layers by a split in the middle to form an outer layer of
somatic mesoderm or parietal layer which lines the ectoderm of the body and
an inner layer of splanchnic mesoderm or visceral layer that lies along the
outer surface of endoderm.
• The somatic mesoderm and the ectoderm together constitute the
somatopleure and the splanchnic mesoderm with endoderm forms the
splanchnopleure.
• The somatic and splanchnic layers are separated from each other except at
their free ends, by a cavity which forms the coelom of the organism.
• When the 2 lateral mesodermal plates fuse in the midventral line, the lateral
coelomic cavities also unite to form a single continuous coelom, also called
splanchnocoel.
• In later stages of development this cavity expands and becomes the body cavity
of adult animal.
• The pericardial cavity is developed from splanchnocoel.
 ORGANOGENESIS
BRAIN
• It is an ectodermal derivative.
FORMATION OF NEURAL TUBE FROM NEURECTODERM
DIFFERENTIATION OF BRAIN
• The anterior part of neural tube is distinguished as encephalon which develops into
regions of brain.
• As the embryo increases in length the neural tube assumes a slightly concave
orientation.
• Encephalon is divided into 3 primary brain vesicles: procenphalon( forebrain),
mesencephalon( mid brain), rhombencephalon( hind brain)
• In the head region the floor of the embryonic brain vesicle almost the posterior end
of prosencephalon bends ventrally around the anterior end of notochord. The
process of bending is cranial flexure.
• The floor of the anterior end of the brain is called tuberculum posterius, it forms the
posterior limit of fore brain.
• Opposite to tuberculum posterius on the roof is dorsal thickening . Lies just behind
closed neuropore.
FOREBRAIN
Anterior brain is again divided into anterior telencephalon and posterior diencephalon.
Posterior limit is tuberculum posterius and anterior limit is lamina terminalis.
Two antero lateral pouches develops from the telencephalon known as telencephalic
vesicles which later develop into cerebral hemispheres. They have a cavity called
telocoel or lateral ventricle.
The lateral ventricles communicate with diencephalon cavity (diocoel) by a foramen
called foramen of monro.
The extensions of lateral ventricles into olfactory lobes are known as olfactory
ventricles or rhinocoels.
The roof and dorso- lateral walls of cerebral lobes thicken to form the cortex or
pallium, while floor and ventro- lateral sides become thickened into corpes striata.
DIENCEPHALON
It is behind telencephalon, the cavity is called third ventricle or diocoel.
The roof of diencephalon becomes non- nervous, which together with vascular
piamater covering it, forms a finger like projection that extend into the diocoel as well
as lateral ventricles, called anterior choroid plexus.
Behind anterior choroid plexus a long finger like outgrowth known as the epiphysis is
seen. Epiphysis grows into brownspot.
Between anterior choroid plexus and epiphysis is habenular ganglion and
commissure.
From ventrolateral wall of the diencephalon a pair of evaginations develop called
optic vesicles.
The floor of the diencephalon gives out a funnel like out pushing called infundibulam.
An invagination from roof grows toward infundibulam to form Rathke’s pocket which
detaches from it and joins infundibulam and forms pituitary gland.
In the floor anterior to infundibulam there are 2 thickenings called optic recess. Called
optic chisama and torus transversus.
MESENCEPHALON
The dorso lateral walls bulge out as a pair of optic lobes or corpora bigemina.
The floor and sides of the mid brain is thickened into tracts of grey matter called crura
cerebri, which is a pathway between fore and hind brain.
The cavity of mesencephalon is mesocoel it is narrowed due to thickening of lateral
walls and roof.
This forms the iter or aqueduct of sylvius.
It connects 3rd and 4th ventricles of the brain.
RHOMBENCEPHALON
The hind brain is demarcated from the mid brain by a transverse constriction.
A transverse thickening in the roof of rhombencephalon develops into cerebellum.
Behind the cerebellum is the posterior choroid plexus.
The ventral and ventro lateral walls of rhombencephalon give rise medulla.
Rhobocoel is the 4th ventricle. It communicates anteriorly to 3rd ventricle and
posteriorly to spinal cord.
 DEVELOPMENT OF EYE
It is the photoreceptor.
Tissues of eye are derived from 3 sources:
• The wall of diencephalon forms retina
• The epidermal ectoderm forms lens
• Mesenchyme of head give rise to fibrous and vascular coats.
During tail bud stage a pair of ventro- lateral evaginations develop from wall of
the diencephalon called optic vesicle and its cavity is optocoel.
Basal part of optic vesicle is connected to brain by optic stalk.
Optic vesicle continues to push outward until it comes in contact with the
inner surface of lateral head epidermis and induce thickening called lens
placode
External wall of optic vesicle that has a contact with epidermis flattens out and
invaginates to form a double walled cup called optic cup.
The invagination of optic vesicle is extended to ventral side of optic stalk and groove
is formed called choroid fissure. Blood vessels enter optic cup through this. At its
beginning choroid knot can be seen where cells of iris arises.
Development of optic cup obliterates the optocoel and the inner and outer walls are
therefore brought into close proximity.
The two layers are continuous with one another at the rim of optic cup which forms
the edges of the pupil, later edges surrounding pupil thin and form iris.
Pigment granules develop in the external layer of optic cup and forms the pigmented
layer of retina.
Inner layer forms the thicker nervous layer. These nerves come out of the optic cup
as optic nerve.
When optic cup forms, the lens placode invaginate to develop a cup like structure,
the lens cup which gradually pinches off called lens vesicle.
Later it is places within the ring of optic cup supported by suspensory ligaments.
The inner cells of lens vesicle form lens fibres and it divided enormously, thus
the inner cavity pf lens vesicle is reduced and it becomes solid lens.
When lens vesicle pinches off from ectoderm the free edges of ectoderm
extends over the lens and in conjunction with head mesenchyme forms cornea.
Mesenchyme cells form choroid coat. Outside of it forms sclera.
3 chambers in the eye.
• The space between the iris and cornea is the anterior chamber
• The space between lens and the iris is the posterior chamber.
• Aqueous humour fills these chamber. Posterior chamber produces them and it
is drained in the anterior chamber.
• The largest chamber is the vitreous chamber which is filled with vitreous
humour derived from retina and lens. This maintains the shape of the eye.
• Eyelids and nictitating membrane develop at later stage.
• Inner lining of eyelids forms conjunctiva which is fused with cornea.
DEVELOPMENT OF HEART
Heart is a derivative of mesoderm and it develops about at tail bud stage.
It develops due to the induction of archenteric floor.
The splanchnic mesenchymal cells grow ventrally toward the midline below the
pharynx and forms heart.
Splanchnic mesoderm at the region of pharynx proliferates and the cells aggregate
to form 2 small masses called endocardial cells.
They become organised into 2 tubes known as endocardial tubes which fuse
together to form a single tube which forms the rudiment of heart.
Bilateral symmetry is evident in the heart development.
Mesoderm converges on either side of endocardial tube and encloses them.
The growing ends of mesoderm meet ventrally at first and fuse below endocardium.
There appears ventral mesocardium temporarily and it disappears.
Mesodermal sheets meet above endocardium and below pharynx.
 when fusion of 2 mesoderms are complete, the heart is suspended from
above by a dorsal mesocardium.
Endocardium is now surrounded by inner myocardium and outer pericardium.
The space between these are pericardial cavity.
Myocardium give rise to striated, syncytial and involuntary muscles of heart.
The rudimentary heart gets divided into 3 chambers: 2 thin walled auricles
and 1 thick walled ventricle.
Posteriorly it receives 2 vitelline veins and anteriorly it leads into truncus
anteriosus or ventral aorta.
Development of heart occurs within the limited space.
The developing heart tube undergoes flexion and torsion within the
pericardium so that the heart becomes a reversed S shaped coiled tube.
Posterior part of the original tube becomes folded dorsally and anteriorly.
Anterior part of heart is folded ventrally and posteriorly.
 the ventricular part of heart is brought downward and auricular part is
upward
The posterior limit is represented by sinus venosus and it the point of fusion
of vitelline veins.
A sheet of endocardium from the roof of atrial chamber and divides the
auricle into left and right chambers.
Ventricle remains undivided.
The truncus arteriosus becomes partially subdivided by vertical and
longitudinal septum and give rise to the ventral paired aortae which are
connected with various aortic arches.