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Notogenesis, neurulation, somitogenesis Development 148 Gastrulation Notogenesis, neurulation, somitogenesis The embryonic axial structures development : 1. primitive streak 2. notochord 3. neural tube 4. somites 3 week development of primitive streak as crucial structure in transformation of bilaminar into trilaminar embryo 1 week from zygote to beginning of implantation Implantation beginning - 7ED endometrium, LP endometrium, epithelium syncytiotrophoblast cytotrophoblast Blastocyst cavity 2 week implantation continues Embryo is at bilaminar blastoderm stage. The period of blastogenesis terminates by bilaminar blastoderm formation and the period of embryogenesis begins. Implantation is completed on ED14 . 3 week development of bilaminar into trilaminar blastoderm/embryo consisting of 3 germ layers: ectoderm, mesoderm, endoderm Epiblast and primitive streak formation Mechanism of embryonic mesoderm and endoderm development Primitive streak morphogenetic centre in mesoderm, endoderm and ectoderm development • It regulates the epiblast cells migration and differentiation, they invaginate in its axis • mechanism? • production of particular growth factor s/morphogenes • FGF8- fibroblast growth factor • mechanism of its activity? • E-cadherin expression decreases in epiblast cells All 3 germ layers develop from EPIBLAST • ectoderm and endoderm are arranged as epithelium • CAM + • mesoderm is not arranget as epithelium, histologically it is connective tissue – primary mesenchyme • CAM - Development of notochord / chorda dorsalis epiblast precursors, invaginating in Hensen´s primitive node, migrate toward the prechordal plate, - creating the cord-like cellular axial structure, s.c. notochord Axial orientation establishment Body axis cranio-caudal, dorso-ventral , right-left established during 3rd week Cranio-caudal axis 1) Cell signalling from the area of rostral/cranial end of embryo, anterior visceral mesoderm, AVM genes OTX2, LIM1, HESX1 and factor Cerebrus - determine the head end (before the primitive streak formation) PP, primitive streak - factor Nodal, belongs to TGFb - transforming growth factor 2) Cell - signalling from primitive node and notochord, gen Brachyury – dorsal mesoderm formation in the middle and caudal part of embryo gene defect – longitudinal axis shortening – caudal dysgenesis /regression Dorso-ventral axis 1) mesoderm ventralisation BMP – bone morphogenetic protein FGF8 – fibroblast growth factor – determine, what will differentiate in ventral mesoderm – kidneys, gonads, blood vessels 2) mesoderm dorsalisation – Noggin, Chordin, Follistatin block the BMP , that results in notochord and somites formation, neuroectoderm differentiation from ectoderm, differentiation of notochord and paraxial mesoderm Signalling molecules and transription factors in primitive streak First, cranial part/ head process of notochord is established. Caudal part is established by caudal morphogenetic system (caudal end of notochord, intensively proliferating, and ectoderm close to primitive node), followed by gradual primitive streak shortening . Neural plate induction mechanism: interaction of notochord with adjacent ectoderm Noggin, Chordin, Follistatin, neural inductors, for ectoderm to neuroectoderm in the region of forebrain and midbrain Hindbrain and spinal cord are induced by WNT and FGF expressed in the region of the caudal morphogenetic system. Neural plate induction and neural tube development (interaction notochord/adjacent ectoderm) neural folds neural groove neural tube Neural tube development (by activity of caudal morphogenetic systém) results in the formation of hindbrain and spinal cord • caudal morphogenetic centre – intensive proliferating caudal end of notochord • caudal end of notochord gets beneath the primitive streak/primitive groove, that becomes determined to neuroepithelium at that region • primitive streak shortening • neural plate lengthening • neurulation (plate to tube development) Neuroporus anterior ED 25 Neuroporus posterior ED 28 Histogenesis of neural tube wall Neuroepithelium differentiation Neural crest formation and its derivates Neural tube development and intraembryonic mesoderm differentiation 4 week neural tube , somites Somites, 42 – 44 pairs of paraxial mesoderm, 1st pair in occipital region, 20ED Segmentation: Notch, WNT, segmentational genes, retinoic acid, FGF8, craniocaudal gradient