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Phosphorylation of H2A at serine 95 (H2AS95ph) mediated by MLK4 promotes flowering and H2A.Z deposition. However, little is known about MLK1, MLK2, and MLK3 during the flowering time. Here, we systemically analyze the functions of MLK family in flowering time and development. Mutation in  MLK3 , but not  MLK1  and  MLK2 , displayed late-flowering phenotype. Loss of  MLK3  function enhanced the late-flowering phenotype of  mlk4  mutant, but not reinforced the late-flowering phenotype of  mlk1 mlk2  double mutants. MLK3 displayed the kinase activity to histone H2AS95ph  in vitro . The global H2AS95ph levels were reduced in  mlk3 mlk4 , but not in  mlk3  and  mlk4  single mutant and  mlk1 mlk2  double mutant, and the H2AS95ph levels in  mlk1 mlk3 mlk4  and  mlk2 mlk3 mlk4  were similar to those in  mlk3 mlk4  double mutant. MLK3 interacted with CCA1, which binds to the promoter of  GI . Correspondingly, the transcription levels and H2AS95ph levels of  GI  were reduced in  mlk3  and  mlk4  single mutant, and greatly decreased in  mlk3 mlk4  double mutant, but not further attenuated in  mlk1 mlk3 mlk4  and  mlk2 mlk3 mlk4  triple mutant. Together, our results suggested that H2AS95ph deposition mediated by MLK3 and MLK4 is essential for flowering time in  Arabidopsis .

Phosphorylation of Histone H2A at Serine 95 Is Essential for Flowering Time and Development in Arabidopsis