Gastrulation

of the
Gastrulation is a
dynamic process during whichathe various
blastula become rearranged and reorganised so as to form organ f
stage called gastrula. The
major events that characterise two or theg are orming are,
Up to lation aare thefhree
gastrulation layete
gastrulation the course of development in most
maternally derived instructions that animals was fol owing
is the originated
developmental processes up to gastrulation
in the egg before directe b
fertilizai
genome. Towards the end of are fertilizatior
largely controlled by mhas
activated and the
blastulation, genetic material of
the
the blastomers
synthesis of substantial amounts oI new RNA
During gatrulation the transition and proteins oe
from control by the
embryonic genome begins. maternal genome
to the
A t the
beginning of
This occurs by the gastrulation the rate of cleavage divisions slows
addition of two gap considerably
cycle. Recall that phases (G, and G,
early cleavage cycle is biphasic phases) into the cleavage
In most
embryos there is little growth or involving only the S and M phases,
Gastrulation is one of the most
change in mass as
gastrulation begins.
by active, well-ordered critical stages in development, which is
movements bring about are-arrangements of the cells in characterised
the embryo, These cell
definite form and
name
morphogenetic structure for the
movements (formatiye
movements occur in the embryo embryo and hence the
during gastrulation,movements). Such
morphogenetic
One of the major tubulation and
consequences of
previously may have been 1ar apart, these cell
movements
organogenesis.
is that
inductive interactions resulung in are brought close together. groups of cells, which
the These cells then undergo
The morphogenetic establishment of the major
blastula to a stage
vements during organ systems.
establishment of germcharacterised by thegastrulation result in the conversion
nversion ofot aà
the external ectoderm, thelayers is the presence
key event in of three germ layers. 1nE
middle mesador layers. The
Another consequence of gastrulation.
mesoderm and8astrulation.
the internal
ernal
The germ
endoderm. layers are
rm layers
germ aie

archenteron. The
archenteron." gastrulation is the endoderm.
formation of primititerm gastrulation formation of a primitive gut or
gut or (Greek: gaster primitive gut the
archenteron. =

stomach) literally means ne

means
 Gastrulation
Morphogenetic movements
The term morphogenetic movements encompasses variety of specific behavioural
a
natterna of embryonic cells(The major morphogenetic
result inreartangemnt of ihe embryo from the blastulamovements during gastrulation
the nresence of three germ layersl However, the other to a gastrula characterised by
intolater stages. These cellular morphogenetic movements continue
rearrangements
also enable inductive interactions between
various groups of cells that culminate in the formation of the
organs and organ systems
Morphogenetic movements.are often classified into epibolic movements (epiboly)
and embolic movements (emboly Epibolic movements are confined to the prospective
ectodermal blastomeres {the embolic movements refer tö thë inward migration of
nrespective clfbfdamesodermal and endodermal blastósheres from the external surface
of the blastula to the interior. The embolic movements include the processes such as
invagination, involution, delamination, infiltration, convergence, divergence and
ingression.

EmbolicMovements
Invagination
This is one of the most common types of morphogenetic movements. This involves
an infolding or inpushing of a layer of cells into the bla_tocoel. This process
results in
the formation of a new cavity called archenteron. This cavity opens to outside by
a

the
wide opening called the blastopore (Fig. 4.1). Invagination typically occurs in
development of microlecithal eggs such as that of Branchiostoma.)Invagination mainly
helps in the formation ofendoderm.

Dorsal lip of Chorda cells Blastocoel

Ectoderm
blastopore.
Ectoderm
Invaginating Dorsal lip
Endoderm Blastopore
Archenteron

Ventral lip Endodermm

Vental lip Mesodermal crescent

Blastocoel
Fig. 4.1 Typical invagination process

volution
 DevelopmentalBioloy-Experimental Embryology

Primitive
Primite
rid
Ectode Aree opea

Mesoderm
Eniodem
Y olk
through primitie .
Fig.4.2 Diagrammatic representation ofinvolution reak
example) The inturnede
the blastoporal lips is d
an

blastoporenvolinvoluutiteod
and mesodermalcels from inner of the blastoporeol
along the margin
cells are deposited inside the embryo
streak of birds mammals. and
isalso noticed in the primitive

Delamination
Theword delamination means separation of groups of cells from other.
groups/The separation of endodermal, mesodermal and notochordal cells from
othegBn'teleastfishes is a good example for delamination) Acording to a widely ar each
view, the endodermformation in birds takes placebydelamination (Fig. 4.3). pted

Prospective
hypobiest tpibias Subgerminal space Large volk-laden cells Subgeminal space

Posterior Yolk Anterior

Prospective hypoblast Subgerminal space

Postenior Anterior B
Blastocoel
Hvpoblast Epiblast Posterior Anterior

Epiblast Blastocoel
C

Posterior Subgerminal space Anterior

Hypoblast
Subgerminal space
Fig. 4.3 Delamination process Fig. 4.4
in birds Diagrammatic representation ot
infiltration
Infiltration (Polyinvagination)
This
involves the
surface of the detachment
blastula and of jndividual cells
or
arrange themselves asa their íalling into
single layer (Fig. 4.4).the blastocoel.groups
In the
of cells from the
52 blastocoel they
 Fig. 4.5 A. Convergence (Bold lines) Fig. 4.5 B. Divergence (Narrow lines)

Divergence (Spreading)
It refers to the migration of involuted cells from the blastopore or primitive
The involuted
streak. In divergence, the cells move in different directions from a single point.
and primitive
cells of notochord and mesoderm migrate and diverge from the blastopore
streak to their future positions within the developing embryo (Fig. 4.5B).

Ingression
Ingression involves movement of individual or groups of cells from the external
cells leave an
layer of blastula into the blastocoel. During ingression, the migrating
epithelial sheet by transforming from epithelial type of cells into freely migrating
mesenchyme cells. To do so, they must alter their cellular architecture, program ot
motility, and adhesive relationship to the surrounding cells. It is categorised intotwNO
types unipolar andmultipolar. In unipolar ingression, the groups of cells move fromn
a particular region. Segregation of germinal layers in the flagellated blaatula .af Porifera
and Coelenterata is an example for unipolar ingression. Among vertebrates, neural
crest cell are an example ofa mesenchymal cell type that emigrates out of an epithelium.
Migration of groups of cells simultaneously from several points into blastocoel is called
multipolar ingression (Fig. 4.6). Multipolar ingression is observed in the formation of
hypoblast from the blastoderm in the prototherian mammal, Echidna.
Epibolic Movements (Ectodermal Expansion)
Epiboly involves the movements of epidermal and neural areas
rocess. The generalmigration-of these two areas is in the during gastrulative
he embryo. As the ectoderm expands in this direction, it anteroposterior direction of
gradually engulfs and surround=
 Developmental Biology-ERJe

ibuill

A. Unipolar ingression B. Unipolar ingression C. Multipolar ingressi

in sea urchin in hypothetical blastula in hypothetical blast
Fig. 4.6 Diagrammatic representation of ingression

the inwardly moving presumptive notochordal, mesodermal and endoder

(Fig. 4.7).piboly occursdue to the rapid division of the ectodermal celsa
cells
ermal
nr are
and not
to the
stretching of the cells. d

A site of
blastopore

yolk plug
Fig. 4.7 Diagrammatic representation of epiboly
Physiology of Gastrulation
Gastrulation is essentially a
of movement of
morphogenetic movements wouldprocess cause an parts of the embryo. These
consequentdy
of the
an increased increased
oxidation. Thus, it has been expenditure of energy and
frog embryo revealed that oxygen
similar sharp increaseprogressively increases from consumption
885. Both catabolic and
in total
oxygen consumption is cleavage to late
gastrula stage.
functions include the anabolic also observed in the sea urehin
activities occur
for
gastrulation breakdown of
glycogen during gastrulation. The
egg. The energy catabou
is obtained
been discovered by the present in the
that the breaking down of
glycogen present in the eE8.requireu
cells of the
dorsal lip of amount of glycogen becomes
lowestblastopore.
IL
animal pole and The rate diminished in
of the
synthesis and nucleic
acid
at the
vegetal pole. Theoxidation oí
glycogen is
invaginaus
protein turnover and synthesis. During
protein t anabolic activities highest au
protein synthesis is particularly in include
gastrulation, there is a sharp pioase in
new
mRNA. mainly the yolk. protein synthesis. incred
period upto theThe far inactive
so There is a The source of
materials
materials for the
stage. blastula stage, genes, which sudden increase 0
in the production
become active andhadstartnot been
54 producingproducing mRNA the
mRNA in the gastru
 Gastrulation

The concept of germ

Concept of Germ Layers
layers dates back to the last half of the
when Caspar Friedrich Wolff, observed
that in the early eighteenth century,
was
mbryo development of chick, the
tormed eat-like layers. Based on this observation,
in
embryos develop through a process he he postulated
that
termed "epigenesis" replace preformation
theory that was followed by early to
scientists
Pander (whose name has been immortalised until then. In 1817, (Heinrich Christian
which the blastodisc rest as 'nucleus of by naming the small mass of white yolk on
layerswhile studying chick embryos. Between Pander') discovered three
1850 and 1855, Robert primordial germ
refined germ
the cell layer concept. The terms Remak had further
introduced respectively by Huxley in 1871 and "mesoderm" and "ectoderm" were
Lankester in 1873.
Later, this trilayer concept was
proved true for many types of embryos
conceptbecame an accepted embryological principle. Towards the end of
and this
the terms ectoderm, endoderm and mesoderm were 19th century,
inner and middle layers of the introduced to refer to the outer,
embryo respectively. The adult organs do not arise
directly from the cells derived by the cleavages
of the zygote. The
at first arranged into layers
called germ layers from which the embryonic
various
cells are
formed. This concept is known as germ layer organs are
theory.
Among animals, sponges show the simplest
layer. Although they have differentiated cells organization, having a single germ
(e.g.
coordination. Diploblastic animals, Cnidaria and collar cells), they lack true tissue
complexity, having two germ layers, the endoderm Ctenophora, show an increase in
and
diploblastic animals is organised into recognisable tissues. Sinceectaderm. The body of
are the only germ
layers present in Diploblastica (Radiata), theseectoderm and endoderm
are termed primary
germ layers. All higher animals (from flatworms
to humans) are
possessing a mesoderm in addition to the germ triploblastic,
mesoderm layers found in Diploblastica. The
usually originates from the cells emigrating into the
ectoderm and endoderm. The space between the
body of triploblastic animals
The term
secondary germ layers have been applied to refer develop recognisable organs.
to the three
endoderm and mesoderm. The
formation of three germ layers is a commonlayers ectoderm,
-

in the early denominator

development of all vertebrates.
Fate of Germ
Layers
The following chart
during development of typical(Table 4-1) shows the derivatives of the three
vertebrate. germ layers
Ectoderm
The ectoderm
gives rise to the following derivatives.
Somatic Ectoderm:
skin glands Epidermis; skin derivatives (scale, nail,
ear vesicles
including the mammary
glands, etc.); olfactory feather, claws, hair,
(labyrinth);
and proctodaeum. cutaneous sense organs; anterior organs; lens of eye; inner
pituitary;lining of stomodaeum
Developmental Biology-ExperimentalEmbryology

Table 4-1 Derivatives

of the Primary Germ Lava
Mesoderm
Ectoderm

All nervous tissue

Skeletal smooth, and Endodermn
Epithelium of digesti
cardiacmuscle tract(except
Epidermis of skin and Cartilage, bone and other and anal thatofofsto
cavities) ive
epidermal derivatives

(hairs, hairfollicles,
sebacecus and sweat
glands, nails)
Cornea and lens
of eye
Epitheliumoforaland
nasalcavities, of paranasal Serosae of ventral body
sinuses, and of anal canal
Tooth enamel
Epithelium ofpineal and
pituitary glands and
adrenal medulla
Melanocytes
Some cranial bones and
branchial cartilages
(derived from neural
crest)
connective tissues
Glandular derivativeses of
Blood, bone marrow, and digestive tract (liver
lymphoid tissues pancreas)
Endothelium of blood Epithelium of respirato
vessels, and lymphatics tract, auditory tube,ator
tonsils an
cavity Thyroid, parathyroid and
thymus glands
Fibrous and vascular tunicsynus glands
of eyes Epithelium of reproductie
ducts and glands
Synovialmembranes of
joints Epithelium of urethra and
bladder
Organs of urogenital
system(ureters, kidneys,
gonads, and reproductive
ducts)

Neural Crest: Branchial skeleton;

ganglia and sensory nerves, adrenal
pigment cells like chromatophores; Some cartilage elements of head and neck.medull,
Neural Tube: Brain, spinal cord, cranial
and spinal motor nerves, retina and
optic nerve, posterior pituitary.
Endoderm
The endoderm gives rise to: Lining of
respiratory system; bladder, certain endocrine digestive tract; liver and pancreas,
lining of vagina, urethra and associated glands (parathyroid, thyroid,
ear cavity in
mammals, etc. reproductive glands; auditory tube andthymusj
midale
Mesoderm
The mesoderm
gives rise to the
Notochord: It is retained in following
reduced or obliterated. prochordates; inderivatives.
the
Epimere: Dermatome- remaining chordates,
chordates, itit DEbecomes
Dermis of skin;
Skeletal muscle and appendicular
Mesomere: Excretory skeleton. Sclerotome Vertebral column;.
lumn; Myotome
Hypomere: Somatic organs and
peritoneum, mesenternes, layer reproductive tracts.
Parietal
heart, blood
56 peritoneum; Splanchnic ayer Visceral
cells, blood
vessels, eonnd layer Vi
 EARLY DEVELOPMENT OF
AMPHIQxus
(Branchiostoma
Amphio (Branchi
nh-phylum
derthe.sub-phylum C lanceolatum) is a
Cephalochordata
mal and dev
animal of the primitive marine chordate
amorphose into thedevelopment
aviparosed adult
is Phylum
indirect with Chordata. Amphioxusanimal
adult
ncamo:larity with some organism. The a
free-living
exhibit similarity with some invertebrates, early developmental ciliated larva that
is an

time
same tim
representative of vertebrate such the Echinodermata,
are represent as stages of amphioxus
The gonads
development in its simple form.
and the at

sexes are separate, but no

cin male)
well-marked sexual
female) dimorphism.
the
testes (in the and ovaries In
the adult
the (in the
on both sidesanimal.
the
een
body between tthe body wall, bordering the atrium and are located
of the
There are about 28 pairs on each side. the muscle
groups (myotomes).
Upon sexual aturity, the sperms and First polar body
ova are discharged into the atrium by the
rupture of the body wall and they escape
through the atriopore into the
SurTounding seawater.

The ovumn
When the ovum is fully grown it is
approximately from 0.01 mm to 0.12 mm
in diameter and is of the microlecithal
ype as it contains a very small amount Sperm
of yolk. It is also called isolecithal
because the yolk granules are almost Egg of Amphioxus
uniformly distributed in the egg
eyloplasm. The egg nucleus or germinal
CE lies towards one side of the egg, the animal pole. The opposite side is called
remain bounded by a plasma membrane.
getal pole. The ooplasmic contents called
membrane of mucopolysaccharides
d e the plasma membrane is a thin
the vitelline membrane.
 94 DevelopmentalBiology

The spermn structure of flagellate

the typical
It is very small, has

middle piece, and tail.

The Head is spherical
the middl
matatozisoz0avery hashmving
Fertilization
in seawater. Females sno.
It is external and takes place pawn throughout the
toward evening, while the animolo
and summer and always aremals
female. ne of these sperms
shed sperms into the
vitelline membrane and
water near the
sWimming
enters the egg usually
near Acrosome
-Head
perae
the vegetal pole of the Fertilization membrane
Middle piece
egg. As soon as the sperm
Anphioxus First polar body-
reaches the egg the sperm
vitelline membrane iIs
lifted off from the surface
of the egg to form a
fertilization membrane.
The space between the
plasma membrane and
fertilization membrane is
Tail
known as perivitelline
space. It gets filled with a
viscous fluid called
perivitelline fluid. The Fertilized egg
entry of the sperm into the
cgg also stimulates the
second maturation
division and the second
animal pole in the polar body is released. It remains
perivitelline attached to the egg at-
to form the
male and female space. Meanwhile, the sperm and egg nuclei
of the
egg and fuse to form a pronuclei enlarg
respectively. They migrate towards the equa
place slightly above the zygote nucleus. The fusion
of the two
equatorialplane of the egg pronuclei laks
Cleavage
The fertilized
furrow egg
undergoes
and occurs holoblastic almost equal avage. The first cleavag
is
meridional
the
zygote into two equal from 60 to 90 cleavage. Tis
animal pole to the blastomeres minutes after fertilization. It divide
vegetal pole. in a
meridional plane fertilization.
lane which
which ex extendsfrom
 95
Early developmentofamphioxus
second akes place
cleavage
onal but at
at right angles to the within
first cle 45 minutes after the tist. It
the first. is also
errahona/
Theal
The

but right angles to the fi

uSt
hlastomeres Cleavage furrow to form four equal-sized
third
cleavage occurs within thirty
blastomereesin
The the latitu minutes after the second and divides the
second and
fiur
Teavagerurrowsto: (horizontal)
TOwsto form eight blastomeres. The plane plane rigt
third cleavage
at
at right angles
furrowtopasse
the first two

ovethe
Ce fuator of the four blastomeres, as age furrow passes slightly
i m a lp o l e
are smaller a
result, the four blastomeres
res near the
micromeres
and the Zygote
d
the vegetal pole
calle

f o u rn e a r

larger calle macro-

are

meres

fourth cleavage is 2-Cell stage

The 4-Cell stage
meridional and is a
Micromeres
gain
i.e., two in
double cleavage Micromeres

B
and the furrows
Number,

from the
animal pole
pass
at right Macromeres
o the vegetal pole
to each other to form Macromeres
angles
blastomeres of which 8-Cell stage 16-Cell stage
16 32-Cell stage
are micromeres and
eight Radial cleavage in
eight are macromeres. amphioxus
The fifth cleavage is latitudinal and double, it divides the eight micromeres and
the eight macromeres Simultaneously into four tiers of eight to form 32 blastomeres.
The sixth set of cleavage planes are approximately meridional and synchronous
producing 64 blastomeres (32 micromeres and 32 macromeres). Gradually cleavage
becomes irregular, micromeres divide more rapidly, the synchronous character of
the division's declines, and the increase in numbers of blastomeres becomes
arithmetical rather than geometrical.
Blastulation
The process of formation Blastocoel
Blastocoel
of the blastula is called
blastulation. As the cleavage
proceeds more and more
Dlastomeres are produced
Blastula
and a semifluid material
Blastula hemisection
accumulates in the centre of
De mass of cells. This serves
 Art whecome ar
that they
h Ph al the astewmes

emhwing a eentral cavity,

the hlaslow d in
blastoeoel, filled nged in a
wlh
Matkrm

ture
his s t h is the blavulaA rully
ith
formed blastul.with Me
thnd el which a e arranged in the form
oe
wIdN O

AFNIN the
a t t N)ells
blastenvwl ln the
Dlatula cells at the vegetal n o l
pole are mna e
somew columnar
animal ke. The blastula of amphiorusis callcalled coeloha
han thw at the

INaNm Nar shapvvd maxs of eells

Gastrulation
tIs the pess by which the single- layered blastui is converted into
avennd sta ture, the gastula. During this process the three germn a.
layers an
wyan fwng anvas pesent on the surtace or the blastula move to ta

NtNN in the embryo, these ovements are called formative or mom

take ther
tNNCnNnts ln anphiovus, gastrulation begins when there are about s e orphogen
cmdyo ie, heween 9h and 10" cleavagCs. The man processes invotvent ellh
axtrulatn of amhionus are () nvaginilion (2)1involution (embolic moveme
nneannt ofoells trvm the surtace into the terior of the embryo) and (3 epihe
nNents. (4) Antenpisterior elongation of the embryo.
1. lavagination
Gastrulation is initiated with
tlattening of the blastoderm at the vegetal pe
a
Thisendaemal plate then gradually invaginates or bends inwards into the blaslecune
hesumptive ertnderm

Blastel

PesunuNae
Presumpave
hesuie cadrienm lnvaginating woxhond
endodem
Pesumpave
esudem
l

Ectodem

Noonthvd
DANal lip of
blastopox*
Vemeral
blasapueAnhenten
byp ot Endodem

Blastoportun D
Gastrulation of amphioxus