An active DNA transposon nDart causing leaf variegation and mutable dwarfism and its related elements in rice

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Abstract
While characterized mutable alleles caused by DNA transposons have been abundant in maize since the discovery of Dissociation conferring variegation by Barbara McClintock, only a few mutable alleles have been described in rice even though the rice genome contains various transposons. Here, we show that a spontaneous mutable virescent allele, pyl‐v, is caused by the disruption of the nuclear‐coded essential chloroplast protease gene, OsClpP5, due to insertion of a 607‐bp non‐autonomous DNA transposon, non‐autonomous DNA‐based active rice transposon one (nDart1), belonging to the hAT superfamily. The transposition of nDart1 can be induced by crossing with a line containing an autonomous element, aDart, and stabilized by segregating out of aDart. We also identified a novel mutable dwarf allele thl‐m caused by an insertion of nDart1. The japonica cultivar Nipponbare carries no aDart, although it contains epigenetically silenced Dart element(s), which can be activated by 5‐azacytidine. Nipponbare bears four subgroups of about 3.6‐kb Dart‐like sequences, three of which contain potential transposase genes, and around 3.6‐kb elements without an apparent transposase gene, as well as three subgroups of about 0.6‐kb nDart1‐related elements that are all internal deletions of the Dart‐like sequences. Both nDart1 and 3.6‐kb Dart‐like elements were also present in indica varieties 93‐11 and Kasalath. nDart1 appears to be the most active mutagen among nDart1‐related elements contributing to generating natural variations. A candidate for an autonomous element, aDart, and a possible application of nDart1 for transposon tagging are discussed.