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Third Week of Development (Trilaminar Germ Disk) The main events during this period: • • • • • Gastrulatin. Formation of the notochord. Establishment of ^ body axes. Growth of ^ embryonic disc(cephalocaudally). Further development of ^ trophoblast. Gastrulation • The process that establishes all three germ layers ( ectoderm, mesoderm, & endoderm), which begins with ^ formation of ^ primitive streak. • ^ Primitive streak: a narrow groove formed on ^ surface of ^ epiblast, which is clearly visible at 15- 16 days embryo. • ^ cephalic end of ^ streak, ^ primitive node, consists of a slightly elevated area surrounding ^ small primitive pit . • Cells of ^ epiblast migrate toward ^ primitive streak, upon their arrival, they detach from ^ epiblast & slip beneath it, this inward movement is known as invagination . • Once ^ cells have invaginated ,some displace ^ hypoblast, creating ^ embryonic endoderm. • Some cells come to lie between ^ epiblast & newly created endoderm to form mesoderm. • Cells remaining in ^ epiblast then form ectoderm. • Epiblast layer is ^ source of all ^ germ layers. • Oropharyngeal membrane at ^ cranial end of ^ disc consists of small region of tightly adherent ectoderm & endoderm without mesoderm. • Cloacal membrane at ^ caudal end of ^ disc similar to ^ orophangeal memb. • ^ Prechordal plate forms between ^ tip of ^ notochord & ^ oropharyngeal membrane & is derived from ^ 1st cells that migrate from ^ node in cephalic direction. • As more & more cells move bet. ^ epiblast & hypoblast layers, they begin to spread laterally & cranially. • Gradually, they migrate beyond ^ margin of ^ disc & establish contact with ^ extraembryonic mesoderm covering ^ yolk sac & amnion. Formation of the Notochord • Prenotochordal cells invaginating in ^ primitive node move forward cranially in ^ midline until they reach ^ prechordal plate, these cells become intercalated in ^ hypoblast for a short time at ^ midline then form ^ notochord plate. • Cells of ^ notochordal plate proliferate & detach from ^ endoderm & form a solid cord of cells , ^ definitive notochord. • ^ Cranial end forms first, & caudal regions are added as ^ primitive streak assumes a more caudal position. • ^ notochord & prenotochordal cells extend cranially to ^ prechordal plate & caudally to ^ primitive pit. • The primitive pit forms an indentation in ^ epiblast, ^ neurenteric canal temporarily connects ^ amniotic & yolk sac cavities. • When ^ cloacal memb. appears, ^ posterior wall of ^ yolk sac forms a small diverticulum that extends into ^ connecting stalk called ^ allantoenteric diverticulum, or allantois, appears around ^ 16th day of development. Establishment of the body axes • Establishment of ^ body axes, anteroposterior, dorsoventral, & left-right, takes place before & during ^ period of gastrulation. • Cephalic & caudal ends of ^ embryo are established before ^ primitive streak is formed. • There are groups of genes which control ^ process of gastrulation & establishment of ^ body axes. Fate map established during gastrulation • Regions of ^ epiblast that migrate through ^ primitive streak have been mapped, & their ultimate fates have been determined. • For ex. cells that ingress through ^ cranial region of ^ node become prechordal plate & notochord; those migrating at ^ lateral edges of ^ node& from ^ cranial end of ^ streak become paraxial mesoderm; cells migrating through ^ midstreak region become intermediate mesoderm; & those migrating through ^ more caudal part of ^ streak form lateral plate mesoderm. Growth of the embryonic disc • The embryonic disc, initially flat almost round, gradually becomes elongated, with a broad cephalic & narrow caudal end. • Continuous migration of cells from ^ primitive streak in cephalic direction leads to growth & elongation of ^ cephalic region. • The migration of cells from ^ primitive streak forward & laterally continues until ^ end of 4th week. • In ^ cephalic region, germ layers begin their differentiation by ^ middle of ^ 3rd week, whereas in ^ caudal part, differentiation begins by ^ end of ^ 4th week. Thus gastrulation (formation of germ layers) continue in caudal segments while cranial structures are differentiating, causing ^ embryo to develop cephalocaudally. Further development of ^ trophoblast • By ^ beginning of ^ 3rd week, ^ trophoblast is characterized by primary villi that consist of a cytotrophoblastic core covered by a syncytial layer. • During further development, mesodermal cells penetrate ^ core of primary villi & grow toward ^ decidua, here are called secondary villi. • By ^ end of ^ 3rd week, mesodermal cells in ^ core of ^ villus begin to differentiate into blood cells & small blood cells forming villus capillary system, ^ villi are called tertiary or definitive placental villus. • Capillaries in tertiary villi make contact with capillaries developing in ^ mesoderm of ^ chorionic plate & in ^ connecting stalk, these vessels , in turn, establish contact with intraembryonic circulatory system, connecting ^ placenta & ^ embryo. • Hence, when ^ heart begins to beat in ^ 4th week of development, ^ villus system is ready to supply ^ embryo with nutrients & oxygen. • Meanwhile, cytotrophoblast cells in ^ villi penetrate into overlying syncytium until they reach ^ maternal endometrium, here they contact with similar extensions from ^ neighboring villus stems forming a thin outer cytotrophoblast shell. • Villi that extend from ^ chorionic plate to ^ decidua basalis called stem or anchoring villi, those that branch from ^ sides of stem villi are free (terminal) villi, through which exchange of nutrients & other factors will occur. • By ^ 19th-20th day ^ embryo is attached to its trophoblastic shell by a narrow connecting stalk.