Four mutant alleles elucidate the role of the G2 protein in the development of C(4) and C(3) photosynthesizing maize tissues.

Maize leaf blades differentiate dimorphic photosynthetic cell types, the bundle sheath and mesophyll, between which the reactions of C(4) photosynthesis are partitioned. Leaf-like organs of maize such as husk leaves, however, develop a C(3) pattern of differentiation whereby ribulose bisphosphate carboxylase (RuBPCase) accumulates in all photosynthetic cell types. The Golden2 (G2) gene has previously been shown to play a role in bundle sheath cell differentiation in C(4) leaf blades and to play a less well-defined role in C(3) maize tissues. To further analyze G2 gene function in maize, four g2 mutations have been characterized. Three of these mutations were induced by the transposable element Spm. In g2-bsd1-m1 and g2-bsd1-s1, the element is inserted in the second intron and in g2-pg14 the element is inserted in the promoter. In the fourth case, g2-R, four amino acid changes and premature polyadenylation of the G2 transcript are observed. The phenotypes conditioned by these four mutations demonstrate that the primary role of G2 in C(4) leaf blades is to promote bundle sheath cell chloroplast development. C(4) photosynthetic enzymes can accumulate in both bundle sheath and mesophyll cells in the absence of G2. In C(3) tissue, however, G2 influences both chloroplast differentiation and photosynthetic enzyme accumulation patterns. On the basis of the phenotypic data obtained, a model that postulates how G2 acts to facilitate C(4) and C(3) patterns of tissue development is proposed.