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The 5-dag seedlings were also morphologically abnormal, in particular, the SAM and leaf primordia showed abnormal differentiation ( Fig 4 ). SAMs were partly fused, and the number of cells in the SAM region was increased in the DEX-treated plants ( Fig 4A ). In addition, the size of the rosette leaves varied among seedlings in the same line, although it was approximately constant among plants in a given line (Fig 4A and 4B ). The 5-dag mock-treated plants were similar to wild-type plants in terms of organ size and shape (Fig 4B ). By contrast, the SAMs of DEX-treated plants were greatly reduced in size. On average, the SAM was reduced to about one quarter of that of wild-type plants (Fig 4B, red arrow). At 5 dag, the 5-dag mock-treated plants were similar to wild-type plants (Fig 4B ). By contrast, the SAM of the DEX-treated plants at 5 dag was severely reduced (Fig 4B, red arrow). The SAMs of both mock-treated and DEX-treated plants at 5 dag were reduced to less than one fifth of those at 10 dag, but the leaf width did not differ significantly between mock- and DEX-treated plants (Fig 4C ). Thus, shorter OsWOX4 knockdown strongly affected not only the size of the SAMs but also the number of leaves in 5-dag plants.
To identify novel rice genes controlling leaf morphogenesis in rice, we used HAs from rice leaf development as baits (Microprobe AS-PR01) on a microarray cDNA library. We hybridized the microarray to compare the gene expression in leaves of 3.5-day-old plants after various treatments. The result revealed one set of genes that were upregulated by ethylene or cytokinin treatment during rice leaf development. We also identified several genes that were downregulated by ethylene or cytokinin treatments.