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Development: Trilaminar Germ Disc


- The process of formation of germ layers ( endoderm, mesoderm and ectoderm) in the embryo - It occurs during the third week of gestation - Gastrulation begins with the formation of the primitive streak on the surface of epibalst - In a 15-16 day embryo primitive streak visible as narrow groove with slightly bulging regions on either side. - The cephalic end of the streak, the primitive node a slightly elevated area surrounding the small primitive pit

A - Dorsal view of a 16 days embryo ( primitive node and streak visible) B- Dorsal view of a 18 days embryo

Cells of the epiblast migrate toward the primitive streak Reaching near the primitive streak these cells become flask shaped, detach from the epiblast, and slip beneath it. (invagination) Some of these cells displace the hypoblast, forming embryonic endoderm Cells which come to lie between the epiblast and newly formed endoderm form mesoderm. Cells remaining in the epiblast form ectoderm ( The epiblast is the source of all of the germ layers)

Formation of Notochord
Prenotochordal cells invaginating in the primitive pit move forward cephalad to reach the prechordal plate These prenotochordal cells become intercalated in the hypoblast at this stage embryo consists of two cell layers that form the notochordal plate Cells of the notochordal plate proliferate and detach from the endoderm to form a solid cord of cells, the definitive notochord. It serves as the basis of the axial skeleton. The notochord and prenotochordal cells extend cranialy to the prechordal plate (an area just caudal to the buccopharyngeal membrane) and caudally to the primitive pit.

At the point where the pit forms an indentation in the epiblast, the neurantic canal temporarily connects the amniotic and yolk sac cavities

The cloacal membrane is formed at the caudal end of the embryonic disc. It consists of tightly adherent ectoderm and endoderm At the same time the posterior wall of the yolk sac forms a small diverticulum that extends into the connecting stalk. This diverticulum is called the allantoentric diverticulum or allantois. This appears at about 16th day of development.

Establishment of the body axes

Establishment of the body axes-anteroposterior,dorsoventral and left right, takes place prior to and during the period of gastrulation The anteroposterior axis is signaled by cells of the posterior margin of the embryonic disc. This Posterior marginal zone (PMZ) secretes an activin like molecule that induces primitive streak formation. This establishes the cranio to caudal axis in the embryo.

Fate Map established during Gastrulation

Regions of the epiblast that migrate and ingress through the primitive streak have been maped and their ultimate fates determined. Cells that ingress through the cranial region of the node become notochord Those migrating at the lateral edges of the node and from the cranial end of the streak become paraxial mesoderm Cells migrating through the mid streak region become intermediate mesoderm Those migrating through the caudal part of the streak form lateral plate mesoderm

TS showing formation of mesoderm A- 17 day B 19 day

Growth of the Embryonic Disc

The embryonic disc, initially flat and almost round, gradually becomes elongated, with broad cephalic and a narrow caudal end Expansion of the embryonic disc occurs mainly in the cephalic region; the region of the primitive streak remains more or less the same size Growth and elongation of the cephalic part of the disc are caused by a continuous migration of cells from the primitive streak in the cephalic direction. This continues until the end of the fourth week At the end of the fourth week the primitive streak shows regressive changes, rapidly shrinks and disappears In the cephalic part , germ layers begin their specific differentiation by the middle of the third week In the caudal part, differentiation begins by the end of the fourth week

Derivatives of the germ layers

The embryonic period or period of organogenesis ( 3rd to 8th week of development)- During this period the three germ layers, ectoderm, mesoderm and endoderm, gives rise to a number of specific tissue and organs.

- At the third week of development the ectoderm is disc shaped, broader in the cephalic than the caudal region - Appearance of the notochord and prechordal mesoderm induces the overlying ectoderm to thicken and form the neural plate. Cells of the plate make up neuroectoderm, initial event in the process of neurilation.

The process, can be divided into four stages: (1) Local elongation of the ectoderm cells in a midline zone of the disc and their reorganization into a pseudostratified epithelium, the neural plate. (2) Reshaping of the neural plate. (3) Bending of the plate into a neural groove. (4) Closure of the neural groove into a neural tube from the midportion to its cranial and caudal ends

After induction, the elongated, slipper-shaped neural plate gradually expands toward the primitive streak By the end of the 3rd week, the lateral edges of the neural plate become more elevated to form neural folds,and the depressed middle portion forms the neural groove Gradually the neural folds approach each other in the midline and fuse to form neural tube. This fusion begins in the region of the future neck (5th somite) Until fusion is complete, the cephalic and caudal ends of the neural tube communicte with the amniotic cavity by cranial and caudal neuropores Closure of the cranial neuropore occurs approximately at day 25( 18-20 somite stage) Posterior neuropore closes at day 27 ( 25 somite stage)

Formation of neural groove and fold

Neurilation is then complete,and the central nervous system is represented by a closed tubular structure with a narrow caudal portion, the spinal cord and broader

cephalic portion, the brain vesicle

By the time neural tube is closed, two bilateral ectodermal thickenings the otic placodes and the lens placodes, become visible in the cephalic region. Otic placodes invaginate and form the otic vesicles, which will develop into structures needed for hearing and equilibrium Lens placodes invaginate and form the lenses of the eyes ( During 5th week)

Neural crest
As the neural folds elevate and fuse, cells at the lateral border or crest of the neuroectoderm begin to dissociate, this cells population, the neural crest will under go an epithelial-tomesenchymal transition as it leaves the neuroectoderm by active migration and displacement to enter the underlying mesoderm. Derivatives of the neural crest- crest cells give rise to a heterogeneous array of tissues

Formation and migration of neural crest cells in the spinal cord

Derivatives of the neural crest Connective tissue and bone of the face and skull Cranial nerve ganglia Adrenal medulla Dermis in the face and neck Arachnoid and piamater (leptomeninges) Spinal (dorsal) root ganglia Sympathetic chain and preaortic ganglia Parasympathetic ganglia of the GIT Cells Melanocytes - Glial cells - Schwann cells - Odontoblasts - C cells of the thyroid

The ectoderm gives rise to organs and structures that maintain contact with the outside world

E.G. - The central nervous system - The peripheral nervous system - The sensory epithelium of the ear, nose and eyes - The epidermis including the hair and nails - Subcutaneous glands, the mammary glands, the pituitary gland and enamel of the teeth

Mesodermal Germ layer Derivatives

By about 17th day mesodermal cells close to the midline proliferate and form a thickened plate called paraxial mesoderm More laterally the mesoderm layer remains thin and called as lateral plate mesoderm Intermediate mesoderm connects paraxial and lateral plate With the appearance and coalescence of intercellular cavities in the lateral plate it divides into two layers 1. Somatic or parietal mesoderm layer- a layer continuous with mesoderm covering the amnion 2. Splanchnic or visceral layer- a layer continuous with mesoderm covering the yolk sac ( Together these layers line a newly formed cavity, the intraembryonic cavity. )

Paraxial Mesoderm
By the beginning of third week, paraxial mesoderm is organised into segemnts somitomeres Somitomeres first appear in the cephalic region and then proceeds cephalocaudally Each somitomeres is consists of mesodermal cells arranged in concentric whorl In the head region somitomeres form in association with segmentation of neural tube into neuromeres. ( Which contribute the majority of the head mesenchyme) From the occipital region caudally, somitomeres organize into Somites. The first pair of somites arises in the cervical region of the embryo at the 20th day of development

Formation of somites
Approximately 3 pairs of somites appear per day At the end of the 5th week, 42 to 44 pairs are present There are 4 occipital, 8 cervical, 12 thoracic, 5 lumber, 5 sacral and 8-10 coccygeal pairs The first occipital and the last 5-7 coccygeal somites later disappear. The remaining form the axial skeleton. During this period the age of the embryo is expressed in number of somites

Number of somites correlated to Approximate age in days Approximate age ( Days) 20 21 23 No. of somites 1- 4 4- 7 7- 10

25 26 27 28 29 30

10- 13
13- 17 17- 20 20- 23 23- 26 26- 29 34- 35

Sclerotome- Cells forming the ventral and medial walls of the somite become polymorphous and shift to surround the notochord. These cells collectively known as scleretome. They will surround the spinal cord and notochord to form the vertebral column - Cells at the dorsolateral portion of the somite migrate as precursors of the limb and body wall musculature. - Cells at the dorsomedial portion of the somite proliferate and migrate down the ventral side of the somite to form Myotome. - The remaining dorsal epithelium forms the Dermatome and together these layers constitute the dermotomyotome

Each segmentally arranged myotome contributes to the muscles of the back ( epaxial musculature) The dermatomes disperse to form the dermis and subcutaneous tissue of the skin Each myotome and dermatome retains its innervation from its segment of origin *Hence each somite forms its own sclerotome(the cartilage and bone), myotome (muscle) and dermatome ( the segmental skin component).* Each myotome and dermatome has its own segmental nerve component.

Intermediate Mesoderm
Which temporarily connects paraxial mesoderm with lateral plate mesoderm It differentiates into urogenital structures

Lateral plate Mesoderm

-It splits into parietal and visceral layers, which respectively line the intraembryonic cavity and surround the organs - Parietal mesoderm, together with overlying ectoderm,will form lateral and ventral body wall - Visceral mesoderm and embryonic endoderm will form the wall of the gut - Mesodermal cells of the parietal will form mesothelial membranes, or serous membranes (lining of the pleural, peritoneal and pericardial cavities)

Summary Following tissues and organs are developed from the mesoderm:Supporting tissues- connective tissue, cartilage and bone striated and smooth musculatures The walls of the heart,blood and lymph vessels Blood and lymph cells Kidney, Gonads and their corresponding ducts The cortex of the suprarenal gland The spleen

Derivatives of the Endoderm

The gastrointestinal tract is the main organ system develop from the endoderm. The endoderm covers the ventral surface of the embryo and forms the roof of the yolk sac With the development and growth of the brain vesicles, the embryonic disc begins to bulge into the amniotic cavity and to fold cephalo-caudally. The head and tail folds are formed because of folding. As a result of cephalocaudal folding, a large portion of the endoderm lined cavity is incorporated into the body of the embryo proper In the anterior part it forms foregut; in the tail region it forms the hind gut. The part between fore and hind gut is the midgut

The midgut temporarily communicates with the yolk sac by Vitelline duct. This duct is wide initially,but later on it becomes narrow and longer

A- presomite embry

B- 7 somites embryo

At its cephalic end, the foregut is bounded by the buccopharyngeal membrane ( an ectodermalendodermal membrane In the 4th week the buccopharyngeal membrane ruptures, establishing an open connection between the amniotic cavity and the primitive gut The cloacal membrane temporarily separates the hind gut from the amniotic cavity. The cloacal membrane breaks down in the 7th week to create the opening for the anus As a result of cephalocaudal and lateral folding, allantois is partially incorporated into the body of embryo, where it forms cloaca. The distal portion of the allantois remains in the connecting stalk

By the 5th week, the yolk sac duct, allantois and umbilical vessels are restricted to the umbilical ring. In humans the yolk sac is a vestigial and probably has nutritive role in the early stages
Summary:- following tissues and organs are developed from the endoderm The epithelial lining of the respiratory tract The epithelial lining of the Gastrointestinal tract The parenchyma of the thyroid, parathyroids, liver & pancreas The reticular stroma of the tonsils and thymus The epithelial lining of the urinary bladder and urethra The epithelial lining of the tympanic cavity and auditory tube