Biology 624 - Developmental Genetics Lecture #4 – Gastrulation Movements GASTRULATION is a complex series of cell movements that: --rearranges cells, giving them new neighbors --results in the formation of 3 GERM LAYERS that will form the subsequent embryo: ectoderm, endoderm and mesoderm --forms the main body axes of the embryo (they were specified earlier but not formed) cleavage * 1.5h 0h organogenesis GASTRULATION 16h 6h * organogenesis cleavage GASTRULATION 2.5h 0h cleavage 2.8h 24h 10h cleavage 3.5h dorsal closure *GASTRULATION neurulation organogenesis 36h 18h * GASTRULATION segmentation 5h 10h organogenesis 24h 48-72h * GASTRULATION cleavage organogenesis segmentation 24h first cleavage 24h 4d 21d 7d * implantation GASTRULATION organogenesis 4.5d 6.5d segmentation 14d 20d Invagination- the infolding of a region of cells Involution- the inturning of an expanding outer layer of cells Ingression- the migration of individual cells from surface to interior Delamination- the splitting of one sheet of cells into two parallel sheets of cells Epiboly- the movement of epithelial sheets that spread as a unit to cover the embryo Convergent Extension Convergent extension- the lengthening of a group of cells in the long axis while shortening in the perpendicular axis Cell Movements during Convergent Extension in Zebrafish Fate map of the fly embryo at cellular blastoderm stage Early Gastrulation in Drosophila Figure 9.40 Gastrulation in Drosophila Schematic Representation of Mid-Gastrulation in Drosophila Germ Band Extension in Drosophila GERM BAND EXTENDED PROCESSES: --organogenesis – e.g. nervous system forms by neuroblast ingression from ectoderm --segmentation – A-P segment unique identity set up --imaginal disc specification – cells set aside that will survive pupation Germ Band Retraction in Drosophila Gastrulation in Drosophila - lateral view Gastrulation in Drosophila – Vade Mecum Movie FGF Signaling is Critical to Gastrulation in Drosophila Stage 7 Stage 8 early gastrulation midgastrulation Stage 9 WildType Mutant Neurotactin, membrane marker Twist, mesoderm marker extended germ band (Gryzik and Muller, 2004) Model for FGF signaling in Drosophila gastrulation (Leptin and Affolter, 2004) Fate Map of the Blastula of the Frog Xenopus laevis Cell Movements During Frog Gastrulation - Early Cell Movements During Frog Gastrulation - Mid Cell Movements During Frog Gastrulation - Late Epiboly of the Ectoderm Is Accomplished by Cell Division and Intercalation Convergent Extension Convergent extension- the lengthening of a group of cells in the long axis while shortening in the perpendicular axis Movie of Xenopus Gastrulation Xenopus Gastrulation – Vade Mecum Fibronectin and Amphibian Gastrulation Normal FN block peptides Blastocoel Roof Explants deposit matrix that directs the migration of mesoderm from Nagel et al, 2004 FGF Expression during avian gastrulation and the putative role of Sprouty/spred from Keller, 2005 Wnt Signaling Mediates Two Distinct Aspects of Gastrulation Convergent Extension Tissue Separation (Huelsken and Birchmeier, 2001) Amphibian Convergence Extension Requires Noncanonical Wnt Signaling (Habas et al, 2003) CE in Zebrafish Requires Oriented Cell Division (Gong et al, 2004) This is the dorsal ectoderm of embryos in gastrulation, Xdd is a Disheveled mt, animal pole is up papc Expression is Defective in spt Mutant Embryos WT spt- At the end of gastrulation---the basic body plan is set up --the three germ layers have formed and will produce all embryonic structures except the germline ECTODERM---skin, nervous system, neural crest MESODERM---muscle, bones, blood and vessels, kidney, gonads, connective tissue ENDODERM---gut, liver, lungs, pancreas C. elegans embryogenesis cleavage 0h * GASTRULATION 1.5h 6h organogenesis 16h Drosophila embryogenesis * organogenesis cleavage GASTRULATION 0h 2.5h dorsal closure 10h 24h Frog embryogenesis * GASTRULATION 3.5h 18h cleavage neurulation organogenesis 36h Zebrafish embryogenesis * GASTRULATION segmentation 2.8h 5h 10h 24h cleavage organogenesis 48-72h Chick embryogenesis Egg laid * GASTRULATION cleavage organogenesis 24h segmentation 4d 7d 21d Mouse embryogenesis first cleavage 24h implantation 4.5d * GASTRULATION organogenesis 6.5d segmentation 14d 20d