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retinoic acid vitamin ra all-trans-retinoic metabolite all-trans-retinol tautomer isomer stereoisomer embryonic embryo intercellular signaling teratogenicity teratogen acid receptor retinoid receptor rxr rar transcription morphogen signaling molecule brain somitogenesis digeorge syndrome somites metabolism biosynthesis cycle mechanism biosynthetic intermediate biogenesis substrate precursor synthesis derivative bioprecursor metabolic endogenous contractor molecular formula skeletal structural structure 3d