In the first half of the 1990s, early evidence on Tyr phosphorylation in plants was reported. Cloning, expression in E. coli, and biochemical characterization of the product of A. thaliana APK1 gene allowed authors to conclude that this novel type of protein kinase could phosphorylate Tyr, Ser, and Thr residues, though Tyr phosphorylation seemed to occur only on limited substrates . Then, further evidence was obtained for the presence in plants of DSKs , and the dual phosphorylation of the Thr-x-Tyr motif in the activation loop of MAPKs was found to be conserved also in plants . The first example of Tyr phosphorylation in plant organelles also represented the first indication of protein phosphorylation as part of a redox signaling mechanism in mitochondria .
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