Vertebrate Embryonic Patterning 6 Limb Formation.

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Presentation transcript:

Vertebrate Embryonic Patterning 6 Limb Formation

Tetrapod Limb anterior posterior proximal distal ventral dorsal

Limb Bud Formation

Establishment of Limb Fields

Hox Gene Expression in Trunk During Limb Bud Development Fig Analysis of Biological Development 2 nd ed, Kalthoff

FGF10 Induces FGF8 supernumerary limbs from implanting FGF10 beads

Positioning of the Limb Field Forelimb position lies near anterior expression boundary of hox b8, b9, c6, c9, & d9 FGF8 expression in intermediate meso + hox genes induces FGF10 in the lateral plate meso Wnts in the lateral plate maintain FGF expression in regions of hoxd9 expression Between the fore & hindlimb buds, the developing kidney (mesonephros) inhibits FGF expression Exogenous FGF can induce supernumerary limbs

Forelimb vs Hindlimb

Forelimb vs Hindlimb Correlates with Tbx5 & Tbx4 Expression Respectively Tbx5 Tbx4

Tbx4 Gain of Function Study Tbx4 expressing retrovirus infected embryo

Proximal Distal Limb Axis AER –overlying ectoderm –responsible for maintaining outgrowth of limb Progress zone –underlying mesoderm –region of cell division –maintained in active mitotic state by AER

AER – Apical Ectodermal Ridge

AER Necessary for Limb Bud Outgrowth Removal of AER at various times prematurely truncates limb

Reimplantation of older AER into younger bud causes loss of proximal structures AER Necessary for Limb Bud Outgrowth Reimplantation of younger AER into older bud causes duplication of distal structures

Interactions of AER and PZ

FGF8 in the AER FGF in ectodermAER 3 day embryo limb buds

Hox Code of the Limb ProximalDistal Hoxa & d = forelimb; Hoxc & d = hindlimb

Mammalian Hox Genes

Hox Genes Required for Proper Limb Development Hoxa-11/Hoxd-11 double knockout wild-type human with synpolydactyly resulting from homozygous HOXD-13 null mutation

Proximal-Distal Structures

Anterior-Posterior Limb Axis Zone of Polarizing Activity ZPA

Classic Experiments: Retinoic Acid Induced Limb Duplication Implanting RA soaked beads caused duplication of zeugopod/autopod structures Depletion of RA reduces the normal structures

Shh is Expressed in ZPA

Demonstration that Sonic Hedgehog is Primary ZPA Component

Dorsal-Ventral Gene Patterning in Limb Bud Fig Analysis of Biological Development 2 nd ed, Kalthoff

Dorsal-Ventral Limb Axis Lmx 1.Fgf-8 induces Shh 2.Shh & Fgf-8 induce Fgf-4 3.All these induce Lmx-1 4.Wnt-7a in the ectoderm represses Lmx-1 and induces Shh BMPs

Molecular Interactions in Limb Bud

Dorsal-to-Ventral Transformations WT mouse autopod Wnt7a -/- mouse autopod

Homology of Process: A-P Axis

Homology of Process: D-V