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Univ Adelaide, Sch Mol & Biomed Sci, ARC NHMRC Res Network Genes & Environm Dev, ARC Special Res Ctr Mol Genet Dev, Adelaide, SA 5005, Australia Univ Adelaide, Sch Mol & Biomed Sci, ARC NHMRC Res Network Genes & Environm Dev, ARC Special Res Ctr Mol Genet Dev, Adelaide, SA 5005, Australia