miR399 is the first miRNA demonstrated to be up-regulated specifically by P deficiency and rapidly decreased upon P readdition ( Fujii et al., 2005 ). The Arabidopsis ( Arabidopsis thaliana ) genome encodes six MIR399 genes ( MIR399A to -F ), which are all induced by P starvation to different extents. Transgenic Arabidopsis plants overexpressing miR399 exhibit increased Pi uptake and allocation to the shoot, resulting in excessive shoot Pi when grown under P-sufficient conditions ( Fujii et al., 2005 ; Aung et al., 2006 ; Bari et al., 2006 ; Chiou et al., 2006 ). The high Pi level is accompanied by chlorosis and necrosis at the tips of mature leaves, which are characteristics of Pi toxicity ( Delhaize and Randall, 1995 ). Remobilization of Pi from old to young leaves during P starvation is impaired in the transgenic lines ( Chiou et al., 2006 ). Similar phenotypes of Pi toxicity and defective Pi allocation are observed in rice ( Oryza sativa ) transgenic plants overexpressing the miR399 homologs ( Hu et al., 2011 ). Overexpression of Arabidopsis miR399 in tomato ( Solanum lycopersicum ) not only results in increased accumulation of Pi but also enhances the
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