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CHAPTER 7

Gastrulation, body folding and coelom formation


Introduction:

 Blastulation results in the formation of several cell lineages, including the epiblast, hypoblast,
trophectoderm, and ICM.

 Epiblast gives rise to three germ layers: ectoderm, mesoderm, and endoderm.

 Gastrulation is the process by which these germ layers form, and it also establishes the
primordial germ cells.

 This chapter focuses on the initial establishment of the germ line.

Development of the Amnion:

 During gastrulation, the trophectoderm is covered by extra-embryonic mesoderm, forming the


chorion.

 Chorioamniotic folds gradually enclose the embryonic disc, creating a sealed amniotic cavity.

 The amnion consists of an inner epithelium derived from the trophectoderm and an outer layer
of extra-embryonic mesoderm.

 In some species, the mesamnion disappears, leaving no connection between the amnion and
chorion, while in others, it persists, causing the amnion to tear during birth.

Early Phases of Gastrulation:

 Gastrulation begins with the appearance of the primitive streak, an elongated accumulation of
cells at the caudal pole of the future embryo.

 The primitive streak is formed by epiblast cells accumulating posteriorly.

 Ingression of epiblast cells between the epiblast and hypoblast marks the initiation of
gastrulation.

 Cells in the primitive streak involute from the epiblast through its basement membrane to
become mes-endodermal precursors capable of forming mesoderm or endoderm.

 This marks the first instance of epithelio-mesenchymal transition, where cells transition from an
epithelial state to a more loosely organized, migratory state.

 The cessation of gastrulation is marked by a suppression of this process.

Formation of Mesoderm and Endoderm:

 Mes-endodermal precursor cells give rise to both endoderm and mesoderm during gastrulation.

 Endoderm-forming cells displace the hypoblast beneath the epiblast, forming the endoderm.

 Some cells remain positioned under the epiblast and trophectoderm, forming intra- and extra-
embryonic mesoderm, respectively.
CHAPTER 7
Gastrulation, body folding and coelom formation
 Endoderm enlarges to create the upper lining of the primitive yolk sac beneath the epiblast and
connects to the hypoderm at the margin of the embryonic disc.

 Mesodermal cells extend beyond the embryonic disc as extra-embryonic mesoderm.

 Both intra- and extra-embryonic mesoderm split into two sheets: somatic (parietal mesoderm)
associating with the epiblast and trophectoderm, and visceral (splanchnic mesoderm)
associating with the endoderm and hypoblast.

 The trophectoderm and extra-embryonic somatic mesoderm together form the chorion, the
outer layers of the embryonic part of the placenta.

 The cavity forming between somatic and visceral mesoderm is referred to as the coelom.

 Initially, the coelom is outside the embryonic disc (extra-embryonic coelom or exocoelom).

 Later, the split between somatic and visceral mesoderm involves intra-embryonic mesoderm,
establishing an intra-embryonic coelom.

 The intra-embryonic coelom, through embryonic folding, will later give rise to body cavities.

 Somatic mesoderm contributes to parietal portions of peritoneum and pleura.

 Visceral mesoderm contributes to the visceral portions of these serous membranes.

Establishment of Intra-embryonic Mesoderm:

 Intra-embryonic mesoderm forms in a posterior-to-anterior direction.

 It progresses parallel with the growing primitive streak.

 After involution, cells spread laterally to form extra-embryonic mesoderm and cranially to form
intra-embryonic mesoderm.

 The entire embryonic disc's growth rate surpasses that of the primitive streak.

 The embryonic disc transforms from an initially oval shape to later become pear-shaped.

 As the disc elongates, the primitive streak gradually shifts to a more posterior location due to
this differential growth.

Development of the Notochord:

 The notochord is a midline structure that plays a pivotal role in establishing the anterior-
posterior embryonic axis.

 It is formed by epiblast cells ingressing through the primitive node, which is located at the
anterior end of the primitive streak.

 The initial cells that ingress through the primitive node form the prechordal plate, a mesodermal
structure located just anterior to the tip of the notochord.
CHAPTER 7
Gastrulation, body folding and coelom formation
 Different organizer regions within the primitive streak contribute to the involution of cells
forming the mesoderm and endoderm.

 In the mouse, three organizer regions are identified: early-gastrula, mid-gastrula, and late-
gastrula organizers.

 The late organizer, the primitive node, gives rise to the prechordal plate and notochord.

 The notochord is delimited anteriorly by the prechordal plate and posteriorly by a structure
called the cloacal membrane, which temporarily seals common openings of the gut, urinary
organs, and reproductive tract.

Early Development of the Ectoderm and Its Derivatives:

 The notochord secretes signaling molecules, including Sonic hedgehog (Shh), which induce the
overlying epiblast to differentiate into neuroectoderm.

 Both the primitive streak (posteriorly) and the nascent neural ectoderm (anteriorly) are visible
on the surface of the embryonic disc.

 In the region anterior to the primitive node, epiblast cells differentiate into neuroectoderm,
leading to the formation of the neural plate.

 The neural plate's lateral edges become elevated, forming neural folds that enclose a midline
depression known as the neural groove.

 The neural folds gradually fuse over the neural groove, completing the formation of the neural
tube.

 Fusion initiates in the future embryonic cervical region and proceeds anteriorly and posteriorly,
resembling the closure of a double zipper.

 Initially, the neural tube opens into the amniotic cavity through the anterior and posterior
neuropores, which eventually close.

 This process establishes the central nervous system as the first embryonic organ system

Development of the Neural Crest:

 Neural crest cells are detached from the lateral border or crest of the neural folds during the
elevation and fusion of the neural folds.

 Neural crest cells do not participate in forming the neural tube but instead migrate widely to
contribute to various tissues, including melanocytes, parts of the nervous system (neurons for
the central, sympathetic, and enteric nervous system), and craniofacial mesenchymal
derivatives.

Surface Ectoderm:

 After allocating cells for endoderm, mesoderm, germ line, and neuroectoderm, the remaining
lateral epiblast differentiates into surface ectoderm.
CHAPTER 7
Gastrulation, body folding and coelom formation
 Bilateral thickenings of the surface ectoderm form the otic placode (develops into the inner ear)
and the lens placode (forms the lens of the eye).

 The remaining surface ectoderm contributes to the epidermis, skin-associated glands,


epithelium of oral and nasal cavities, and the caudal portion of the anal canal.

 The epithelium covering the oral cavity also gives rise to the enamel of the teeth and a portion
of the pituitary gland (adenohypophysis).

Mesoderm and Early Derivatives:

 Formation of the notochord establishes an embryonic midline axis.

 Initially, mesoderm forms a thin, loosely woven mesenchyme on either side of the notochord.

 In the occipital region, paraxial mesoderm proliferates and forms somitomeres (later somites),
which give rise to connective tissue, bone, cartilage, dermis, skeletal muscle, and vertebrae.

 Lateral plate mesoderm remains thin and becomes the somatopleura (somatic mesoderm) and
splanchnopleura (visceral mesoderm) after division into somatic and visceral mesoderm by the
extra-embryonic coelom.

 The intermediate mesoderm differentiates into structures of the urinary and gonadal system.

 Nephrogenic cord forms a temporary kidney, the mesonephros, and the gonads receive
primordial germ cells.

 The embryonic body undergoes anterior-posterior and lateral folding, defining intra- and extra-
embryonic cavities.

 The somatopleura forms the lateral and ventral body wall, while the splanchnopleura forms the
wall of the primitive gut and its derivatives.

 The intra-embryonic coelom divides into peritoneal, pleural, and pericardial cavities, with serous
membranes lining these cavities derived from somatic mesoderm (parietal surfaces) and visceral
mesoderm (visceral surfaces covering organs).

Blood and Blood Vessel Formation:

 Both blood and blood vessels originate from common mesoderm precursor cells known as
haemangioblasts.

 Haemangioblasts differentiate into two types of precursor cells: haematopoietic stem cells
(which form blood cells) and angioblasts (which form endothelial cells).

 These endothelial cells come together to create the foundation for blood vessel formation.

Endoderm and Its Early Derivatives:

 The endoderm is responsible for forming the inner epithelial lining of the gastrointestinal tract
and its associated structures.
CHAPTER 7
Gastrulation, body folding and coelom formation
 Initially, the endoderm forms the epithelium of the roof of the primitive yolk sac, and with
embryonic folding, it becomes the lining of the primitive gut.

 The primitive gut consists of the foregut, midgut, and hindgut, each contributing to different
parts of the gastrointestinal system.

 The foregut gives rise to the pharynx, middle ear, thyroid and parathyroid glands, liver,
pancreas, and parts of the esophagus, stomach, and duodenum.

 The midgut contributes to most of the small and large intestines.

 The hindgut forms the transverse and descending colon, rectum, and part of the anal canal.

 At the caudal end of the hindgut, the cloaca is formed and later separates into the
gastrointestinal and urogenital systems.

 The allantois develops as an outgrowth from the hindgut into the extra-embryonic coelom and
plays a role in forming the allantochorionic placenta in domestic animals.

Primordial Germ Cells:

 Primordial germ cells are a population of epiblast cells set aside during mesoderm and
endoderm formation for the future germ line.

 They become recognizable during gastrulation, often in the posterior rim of the embryonic disc.

 These cells are displaced from the embryonic disc to the wall of the definitive yolk sac and, to
some extent, the allantois.

 Their relocation pathway may involve passive carriage with the endoderm.

 Ultimately, primordial germ cells find their way to the genital ridges, where they continue to
multiply until they either initiate meiosis (in female embryos) or enter mitotic arrest until
puberty (in male embryos).

Certainly, let's summarize the key points in a clear and organized manner:

Gastrulation and Germ Layers:

 During gastrulation, the epiblast forms the primitive streak, leading to the formation of
endoderm and mesoderm, including primordial germ cells.

 The remaining epiblast becomes the ectoderm, one of the three primary germ layers.

Development of the Central Nervous System:

 The central nervous system, derived from the ectoderm, is the first organ to form.

 Neurulation occurs, resulting in the establishment of the neural tube, which differentiates into
brain vesicles and the spinal cord.
CHAPTER 7
Gastrulation, body folding and coelom formation
 Neural crest cells, a specialized cell population, detach before neural tube closure and
contribute to various tissues in the body, including parts of the nervous system.

 Surface ectoderm, derived from the ectodermal germ layer, forms the epidermis, which includes
structures like hair, horn, claws, hooves, and subcutaneous glands.

Endoderm and Its Derivatives:

 The endoderm gives rise to several vital structures, including the gastrointestinal tract,
respiratory tract, parts of the urinary system, parts of the middle ear, the thyroid and
parathyroid glands, the liver, the pancreas, and the stroma of the tonsils and thymus.

Mesoderm and Its Subdivisions:

 The mesoderm is divided into paraxial, intermediate, and lateral plate mesoderm.

 In the cranial region, paraxial mesoderm forms somitomeres, which later contribute to head
formation.

 More caudally, somitomeres give rise to somites, each of which has three components:
sclerotome (forms the vertebral column), myotome (gives rise to musculature), and dermatome
(forms skin layers).

 The urogenital system is derived from the intermediate mesoderm, and the development of the
genital system involves primordial germ cells.

 The lateral plate mesoderm splits into somatic and visceral sheets, creating the intra-embryonic
coelom.

 The somatopleura associates with the trophectoderm to form the chorion and with surface
ectoderm to create the somatopleura.

 The splanchnopleura associates with the endoderm, forming the serous membranes covering
internal organs.

 Blood and blood vessels, the cortical portion of the adrenal glands, the spleen, and other
structures are also derived from the mesodermal germ layer.

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