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Developmental Biology – Biology 4361 



    Gastrulation – 
 Chick, Fish, Mammal 

        November 15, 2005
Zebrafish gastrulation 
blastodisc




                                       Haffter et al., 1996 
Zebrafish gastrulation 

                                      hypoblast formation 
                                        – ingression and/or involution of deep cells 
                                                                embryonic shield 




ENL          periderm (extraembryonic protective membrane) 

epiblast      ectoderm 

hypoblast       mesendoderm (mesoderm and endoderm precursors) 

YSL          endoderm                                                    Figure 10.21 
Zebrafish embryonic shield 




                                                                           ­ driven by YSL 



Figure 10.22 



      Embryonic shield functionally equivalent to amphibian dorsal lip of the blastopore 
         ­ can organize 2° axis when transplanted
Zebrafish gastrulation 
blastodisc      yolk syncytial layer    epiblast 


                                                    hypoblast 




                                                    somite 
                                        somite

                                                    notochord 




                                                    Haffter et al., 1996 
Chick cleavage 




      area pellucida 
                             delamination 
area opaca                   migration 




                                     ­ epiblast forms all three germ layers 
                                        (plus extraembryonic membrane) 

                                     ­ hypoblast forms extraembryonic endoderm 
                                         ­ does not form any embryonic 
                                              endoderm or mesoderm
                                             ­ forms primordial germ cells 
  Figure 5.16                                ­ different from fish hypoblast 
Gastrulation ­ chick 

          ­ area pellucida =  inner transparent portion of 
              the blastoderm above subgerminal space 


          ­ area opaca  = ‘opaque’ peripheral area; 
              cells in contact with yolk 




          ­ gastrulation starts with extensive cell 
             rearrangements in posterior epiblast 


          ­ cells move to midline and the forward 
             primitive streak




                                        Figure 10.23 
Gastrulation ­ chick 



                ­ elongation of primitive streak and formation of 
                     primitive groove & primitive pit 

                 ­ anterior end of primitive ridges are thickened 
                    = Hensen’s node 




  ­ major gastrulation events occur at the 
     primitive pit & 
     primitive groove 

  ­ functional equivalent of amphibian blastopore
Chick gastrulation


                             involution 
                             ingression 




                ­ future endoderm ingress first; 
                   displace hypoblast 




                                       Figure 10.24 
Chick gastrulation 

              ­ future mesoderm (axial, paraxial, lateral) cells ingress from 
                   anterior end of primitive pit 
              ­ epithelial to mesenchymal transformation 




{ 

 ­ extraembryonic membranes surround yolk 
    ­ ectoderm – spread by epiboly 
    ­ hypoblast cells spread along yolk surface 
    ­ mesoderm cells in between
Gastrulation  fate 
  map ­ chick




          Figure 10.26 
Chick – 24 h




  Figure 10.27 
Chicken development 1 




                         head region 
                         starts to lift off 
                         the yolk
Chicken  development 2 




ventral closure of the body proceeds from anterior to posterior
Chicken development 3 




        ventral closure of the body is completed
Chicken  development 


         extraembryonic membranes and extraembryonic 
         blood vessels completely surround the yolk 




          after completed ventral closure, embryo 
          rests on top of the yolk sac




         yolk sac is connected to the gut by a 
         narrow bridge 
Mammalian cleavage 

  Early mammalian development not very well studied: 
    ­ small eggs 
    ­ difficult to maintain in culture beyond blastocyst stage 
    ­ ethical issues 

  ­ holoblastic cleavage results in a blastocyst: 




                                                     inner cell mass         trophoblast 



                                                        embryo         ­ hatching from zona 
                                                                       ­ implantation in uterus 
                                                                       ­ placenta formation 



                                               ­ inner cell mass & trophoblast are equivalent to 
Figure 5.11 
                                                   epiblast & hypoblast of the chicken embryo 
Early development in mammals 
               Human: 0 ­ 6 days




                                                                      Figure 5.8 
­ embryo enters the uterus in the ‘morula’ stage 
­ blastocyst stage makes contact with uterus 
­ before implantation into uterus, the embryo sheds zona pellucida 
Early development in mammals 




hatching of a mouse               mouse blastocyst         initial implantation of the blastocyst 
   blastocyst from               entering the uterus                  (rhesus monkey)
 the zona pellucida 




                                    th 
Gilbert SF, Developmental Biology, 6  ed, Sinauer, 2000 
Early development in mammals 
                                                       Human: 7 – 11  days


                                                      ­ inner cell mass delaminates hypoblast 
                                                          cells that line the blastocoel, forming 
                                                          extraembryonic endoderm 
                                                           ­ results in epiblast & hypoblast 



                                                      ­  trophoblast divides into: 
                                                           ­ cytotrophoblast ­ anchors embryo to 
                                                               uterus tissue 
                                                           ­ syncytiotrophoblast ­ furthers 
                                                               progression of the embryo into 
                                                               uterus wall by digesting uterus tissue 



                                                       ­  amniotic cavity begins to form 

                                    th 
Gilbert SF, Developmental Biology, 7  ed, Sinauer, 2003 
Early development in mammals 
                                                               Human 9 – 11 days

                                                   ­ epiblast delaminates the amniotic ectoderm that 
                                                      surrounds the amniotic cavity 


                                                   ­ two­layered blastodisc similar to that in birds 
                                                      embryonic epiblast & hypoblast 


                                                   ­ cytotrophoblast enzymatically remodels 
                                                      maternal blood vessels (trophoblastic lacunae) 


                                                   ­ uterus sends additional blood vessels 


                                                   ­ embryo is nourished by maternal blood 


                                                   ­ extraembryonic mesoderm forms; it will develop 
                                                      into the extraembryonic coelom 


                                    th 
Gilbert SF, Developmental Biology, 7  ed, Sinauer, 2003 
Human blastocyst – 12 d




Figure 10.31 
Gastrulation in mammals 




     cells ingress and involute along the primitive groove and at Hensen’s node 
        ­ form endoderm (replaces hypoblast) and mesoderm

                                    th 
Gilbert SF, Developmental Biology, 6  ed, Sinauer, 2000 
Human gastrulation – 16 d




                            Figure 10.32 
Primate germ layer derivatives 


embryo proper




                                             Figure 10.30 

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Nps3 c

  • 1. Developmental Biology – Biology 4361  Gastrulation –  Chick, Fish, Mammal  November 15, 2005
  • 2. Zebrafish gastrulation  blastodisc Haffter et al., 1996 
  • 3. Zebrafish gastrulation  hypoblast formation  – ingression and/or involution of deep cells  embryonic shield  ENL  periderm (extraembryonic protective membrane)  epiblast  ectoderm  hypoblast  mesendoderm (mesoderm and endoderm precursors)  YSL  endoderm Figure 10.21 
  • 4. Zebrafish embryonic shield  ­ driven by YSL  Figure 10.22  Embryonic shield functionally equivalent to amphibian dorsal lip of the blastopore  ­ can organize 2° axis when transplanted
  • 5. Zebrafish gastrulation  blastodisc  yolk syncytial layer  epiblast  hypoblast  somite  somite notochord  Haffter et al., 1996 
  • 6. Chick cleavage  area pellucida  delamination  area opaca  migration  ­ epiblast forms all three germ layers  (plus extraembryonic membrane)  ­ hypoblast forms extraembryonic endoderm  ­ does not form any embryonic  endoderm or mesoderm ­ forms primordial germ cells  Figure 5.16  ­ different from fish hypoblast 
  • 7. Gastrulation ­ chick  ­ area pellucida =  inner transparent portion of  the blastoderm above subgerminal space  ­ area opaca  = ‘opaque’ peripheral area;  cells in contact with yolk  ­ gastrulation starts with extensive cell  rearrangements in posterior epiblast  ­ cells move to midline and the forward  primitive streak Figure 10.23 
  • 8. Gastrulation ­ chick  ­ elongation of primitive streak and formation of  primitive groove & primitive pit  ­ anterior end of primitive ridges are thickened  = Hensen’s node  ­ major gastrulation events occur at the  primitive pit &  primitive groove  ­ functional equivalent of amphibian blastopore
  • 9. Chick gastrulation involution  ingression  ­ future endoderm ingress first;  displace hypoblast  Figure 10.24 
  • 10. Chick gastrulation  ­ future mesoderm (axial, paraxial, lateral) cells ingress from  anterior end of primitive pit  ­ epithelial to mesenchymal transformation  {  ­ extraembryonic membranes surround yolk  ­ ectoderm – spread by epiboly  ­ hypoblast cells spread along yolk surface  ­ mesoderm cells in between
  • 11. Gastrulation  fate  map ­ chick Figure 10.26 
  • 13. Chicken development 1  head region  starts to lift off  the yolk
  • 15. Chicken development 3  ventral closure of the body is completed
  • 16. Chicken  development  extraembryonic membranes and extraembryonic  blood vessels completely surround the yolk  after completed ventral closure, embryo  rests on top of the yolk sac yolk sac is connected to the gut by a  narrow bridge 
  • 17. Mammalian cleavage  Early mammalian development not very well studied:  ­ small eggs  ­ difficult to maintain in culture beyond blastocyst stage  ­ ethical issues  ­ holoblastic cleavage results in a blastocyst:  inner cell mass trophoblast  embryo  ­ hatching from zona  ­ implantation in uterus  ­ placenta formation  ­ inner cell mass & trophoblast are equivalent to  Figure 5.11  epiblast & hypoblast of the chicken embryo 
  • 18. Early development in mammals  Human: 0 ­ 6 days Figure 5.8  ­ embryo enters the uterus in the ‘morula’ stage  ­ blastocyst stage makes contact with uterus  ­ before implantation into uterus, the embryo sheds zona pellucida 
  • 19. Early development in mammals  hatching of a mouse  mouse blastocyst  initial implantation of the blastocyst  blastocyst from  entering the uterus  (rhesus monkey) the zona pellucida  th  Gilbert SF, Developmental Biology, 6  ed, Sinauer, 2000 
  • 20. Early development in mammals  Human: 7 – 11  days ­ inner cell mass delaminates hypoblast  cells that line the blastocoel, forming  extraembryonic endoderm  ­ results in epiblast & hypoblast  ­  trophoblast divides into:  ­ cytotrophoblast ­ anchors embryo to  uterus tissue  ­ syncytiotrophoblast ­ furthers  progression of the embryo into  uterus wall by digesting uterus tissue  ­  amniotic cavity begins to form  th  Gilbert SF, Developmental Biology, 7  ed, Sinauer, 2003 
  • 21. Early development in mammals  Human 9 – 11 days ­ epiblast delaminates the amniotic ectoderm that  surrounds the amniotic cavity  ­ two­layered blastodisc similar to that in birds  embryonic epiblast & hypoblast  ­ cytotrophoblast enzymatically remodels  maternal blood vessels (trophoblastic lacunae)  ­ uterus sends additional blood vessels  ­ embryo is nourished by maternal blood  ­ extraembryonic mesoderm forms; it will develop  into the extraembryonic coelom  th  Gilbert SF, Developmental Biology, 7  ed, Sinauer, 2003 
  • 23. Gastrulation in mammals  cells ingress and involute along the primitive groove and at Hensen’s node  ­ form endoderm (replaces hypoblast) and mesoderm th  Gilbert SF, Developmental Biology, 6  ed, Sinauer, 2000