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Background  The key enzymes of photosynthetic carbon assimilation in C 4  plants have evolved independently several times from C 3  isoforms that were present in the C 3  ancestral species. The C 4  isoform of phosphoenolpyruvate carboxylase (PEPC), the primary CO 2 -fixing enzyme of the C 4  cycle, is specifically expressed at high levels in mesophyll cells of the leaves of C 4  species. We are interested in understanding the molecular changes that are responsible for the evolution of this C 4 -characteristic PEPC expression pattern, and we are using the genus  Flaveria  (Asteraceae) as a model system. It is known that  cis -regulatory sequences for mesophyll-specific expression of the  ppcA1  gene of  F. trinervia  (C 4 ) are located within a distal promoter region (DR).    Results  In this study we focus on the proximal region (PR) of the  ppcA1  promoter of  F. trinervia  and present an analysis of its function in establishing a C 4 -specific expression pattern. We demonstrate that the PR harbours  cis -regulatory determinants which account for high levels of PEPC expression in the leaf. Our results further suggest that an intron in the 5' untranslated leader region of the PR is not essential for the control of  ppcA1  gene expression.    Conclusion  The allocation of  cis -regulatory elements for enhanced expression levels to the proximal region of the  ppcA1  promoter provides further insight into the regulation of PEPC expression in C 4  leaves.

Evolution of the C4 phosphoenolpyruvate carboxylase promoter of the C4 species Flaveria trinervia: the role of the proximal promoter region