The gene for pyruvate, orthophosphate dikinase in C4 plants: structure, regulation and evolution

Pyruvate, orthophosphate dikinase (PPDK; EC 2.7.9.1) is a key enzyme in photosynthesis in plants that exploit the C4 photosynthetic pathway for the fixation of CO2. This review focuses on the structure, regulation and evolution of the C4-type ppdk gene in the maize genome. The C4-ppdk gene in maize consists of 19 exons spanning about 12 kbp. The gene is transcribed from two different initiation sites under the control of two promoters to produce two mRNAs of different sizes. The larger one contains the exon 1 sequence that encodes the chloroplast transit peptide and its product acts as C4-PPDK in chloroplasts, while the smaller one does not contain the sequence and its product may function as a C3-enzyme in the cytosol. This unusual dual promoter system is not unique to the maize C4-type ppdk gene since the same organization is also observed in the rice (C3 plant) ppdk gene and in Flaveria. Thus, the two-promoter system is common to plant ppdk genes from C3 and C4, monocot and dicot plants. A discussion is also presented of the generation of a system for regulation of the expression of the C4-type ppdk gene. A chimeric gene consisting of a reporter gene under the control of the promoter of maize C4-ppdk is exclusively expressed in photosynthetic tissues and not in roots or stems of transgenic rice. The expression of the introduced gene is also regulated by light: it is low in etiolated leaves and is enhanced by illumination. These results indicate that the regulatory system that controls ppdk expression in maize is not unique to C4 plants.