Through systematic analysis of a comprehensive suite of oat mutants defective in the biosynthesis of the UV-fluorescent triterpene glycoside avenacin A-1, we characterized two genes overlapping with the avenacin pathway, and participate in the biosynthesis of avenacins. SAD4 is a broad-substrate phenol glucosyltransferase and PAL2 is a phenylalanine ammonia-lyase catalysing the first committed step in phenol biosynthesis. They are involved in avenacin biosynthesis by synthesizing the donors for its glycosylation and acylation.
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