Gastrulation in Birds

…telolecithal eggs & discoidal meroblastic cleavage.

• By the time a hen has laid an egg, the blastoderm
contains some 20,000 cells.
• Most of the cells of the area pellucida remain at
the surface, forming the epiblast,
• Other area pellucida cells delaminate & migrate
individually into the subgerminal cavity to form
the polyinvagination islands (primary
hypoblast).
 Gastrulation in Birds
• Shortly thereafter, a sheet of cells from the
posterior margin of the blastoderm migrates
anteriorly to join the polyinvagination
islands, thereby forming the secondary
hypoblast
 Gastrulation in Birds
• The two-layered blastoderm (epiblast and
hypoblast) is joined together at the margin of the
area opaca, and the space between the layers forms
a blastocoel

• The avian embryo comes entirely from the

epiblast. The hypoblast does not contribute any
cells to the developing embryo
• Rather, the hypoblast cells form portions of the
external membranes, especially the yolk sac
 Gastrulation in Birds
• All three germ layers of the embryo proper
(plus a considerable amount of
extraembryonic membrane) are formed
from the epiblastic cells
 Gastrulation in Birds
The primitive streak
• It is the major structural characteristic of avian,
reptilian, & mammalian gastrulation
• This streak is first visible as a thickening of the
epiblast at the posterior region of the embryo, just
anterior to Koller's sickle
• This thickening is caused by the ingression of
endodermal precursors from the epiblast into the
blastocoel & by the migration of cells from the
lateral region of the posterior epiblast toward the
center
 Gastrulation in Birds
• As these cells enter the primitive streak, the
streak elongates toward the future head
region.

• At the same time, the secondary hypoblast

cells continue to migrate anteriorly from the
posterior margin of the blastoderm
 Gastrulation in Birds
• The primitive streak defines the axes of the
embryo.
• It extends from posterior to anterior;
migrating cells enter through its dorsal side
and move to its ventral side; and it
separates the left portion of the embryo
from the right
A

C
B
 Gastrulation in Birds
• As cells converge to form the primitive
streak, a depression forms within the streak.
This depression is called the primitive
groove, and it serves as an opening through
which migrating cells pass into the
blastocoel. (Fig.A)
• The primitive groove is analogous to the
amphibian blastopore.
 Gastrulation in Birds
• At the anterior end of the primitive streak is a
regional thickening of cells called the primitive
knot or Hensen's node (Fig. B)
• The center of this node contains a funnel-shaped
depression (a.k.a primitive pit) through which
cells can pass into the blastocoel.
• Hensen's node is the functional equivalent of the
dorsal lip of the amphibian blastopore
 Gastrulation in Birds

• The embryonic germ layers are formed by

the migration of cells in the epiblast toward
the primitive streak, and their ingression to
form the middle and lower germ layers (the
embryonic mesoderm and endoderm).
 Gastrulation in Birds
• The anterior portion of the primitive streak
and the node serve as a passageway for
cells.
• The first cells to pass through the area are
the cells that are fated to become the
endoderm.
• These cells migrate to the bottom of the
cavity & replace the hypoblast cells
 Gastrulation in Birds
• The prospective mesoderm cells are the
next to enter the primitive streak and leave
the epiblast.
• However, they do not move down to join
the hypoblast but instead make a U-turn and
migrate outward btn the epiblast and
hypoblast to form a separate layer btn the
two.
During gastrulation in birds and mammals, epiblast cells converge at the midline and ingress at the
mitivestreak. Ingression of these cells results in formation of the mesoderm and replacement of some of the
hypoblast cells to produce the definitive endoderm.

                                          

      
• The epiblast & the epiblast cells remain to
become ectodermal cells

• The ectoderm undergoes epiboly to

surround the yolk mass.
                                            
  
A cross-section through the embryo allows
us to observe the three germ layers that form
during gastrulation: ectoderm, mesoderm,
and endoderm.

                                      
 Gastrulation in Mammals
• …….remember my lecture on mammalian
cleavage
• Mammalian development parallels that of reptiles
and birds
• ….why?
• …..because birds & mammals are both
descendants of reptilian species
• ……but the mammalian embryo obtains nutrients
directly from its mother and does not rely on
stored yolk
….. Gastrulation in Mammals
• The chorion forms the fetal portion of the
placenta.
• It will induce the uterine cells to form the
maternal portion of the placenta called
decidua.
• The decidua becomes rich in the blood
vessels that will provide oxygen and
nutrients to the embryo.
 ….. Gastrulation in Mammals
• Gastrulation in mammals begins with segregation of cells
within the ICM
• This results into formation of the hypoblast (a.k.a
primitive endoderm) layer

• The hypoblast cells delaminate from the ICM to line the

blastocoel cavity, where they give rise to the
extraembryonic endoderm, which forms the yolk sac

• These cells do not produce any part of the newborn

organism as in avian embryos
 ….. Gastrulation in Mammals
• The remaining ICM tissue above the hypoblast is
now referred to as the epiblast
• The embryonic epiblast is separated from the
other epiblast cells, which form the amnionic
cavity (fig 1. b, c)
• The cavity is filled with a secretion called
amnionic (amniotic) fluid, which serves as a
shock absorber for the developing embryo while
preventing its desiccation
 (A & B) blastocyst
immediately prior to
gastrulation

Fig. 1: Tissue formation in the human embryo between

days 7 and 11.
 ….. Gastrulation in Mammals
• The embryonic epiblast is believed to
contain all the cells that will generate the
actual embryo, and it is similar in many
ways to the avian epiblast
 ….. Gastrulation in Mammals
• Gastrulation begins at the posterior end of the
embryo
• Like the chick epiblast cells, the mammalian
mesoderm and endoderm migrate through a
primitive streak.
• Also like their avian counterparts, the migrating
cells of the mammalian epiblast detach from their
neighbors and migrate through the streak as
individual cells
• Those cells migrating through the node give rise
to the notochord.
 The movt of the epiblast cells through the
primitive streak. The ingressing epiblast cells are
thought to replace the hypoblast cells. while at
day 16, the ingressing cells fan out to form the
mesodermal layer.

Fig 2. Amnion structure and cell movements during

human gastrulation.
Fig. 3
 ….. Gastrulation in Mammals
Formation of extraembryonic membranes
• While the embryonic epiblast is undergoing cell
movements, the extraembryonic cells are making the
distinctly mammalian tissues that enable the fetus to
survive within the maternal uterus

• The initial trophoblast cells of mice and humans divide &

give rise to a population of cells wherein nuclear division
occurs in the absence of cytokinesis.

• The original type of trophoblast cells constitute a layer

called the cytotrophoblast, whereas the multinucleated
type of cell forms the syncytiotrophoblast
 ….. Gastrulation in Mammals
• The cytotrophoblast cells contain proteolytic
enzymes that enable them to enter the uterine wall
and remodel the uterine blood vessels so that the
maternal blood bathes fetal blood vessels

• The syncytiotrophoblast tissue is thought to

further the progression of the embryo into the
uterine wall by digesting uterine tissue (Fig. 4)
 Uterine
epithelium

ICM
Splits to
form
Epiblast
and
Hypoblast

Fig. 4
Schematic diagram showing the origins of early mammalian
tissues
 FORMATION OF THE COELOM
• Coelom is a space completely enclosed in mesoderm or body cavity
—the space between the gut and body wall
• In deuterostomes, the mesoderm often forms from the archenteron
wall, and the coelom develops within in it, sometimes as a direct out-
pouching of the archenteron (enterocoely) e.g Ehinodermata and
chordata.
• Protostomes, the coelom forms as a split in mesoderm (schizocoely)
e.g Annelida, Mollusca and Arthropoda
 Identical twins
Identical twins (roughly 0.25% of human births) may occur as separation of early
blastomeres, or even by the inner cell mass into two regions within the same
blastocyst.
» End of Gastrulation Lectures