Spatial resolution of an integrated C₄+CAM photosynthetic metabolism

C₄ and CAM photosynthesis have repeatedly evolved in plants over the past 30 million years. Because both repurpose the same set of enzymes but differ in their spatial and temporal deployment, they have long been considered as distinct and incompatible adaptations. Portulaca contains multiple C₄ species that perform CAM when droughted. Spatially explicit analyses of gene expression reveal that C₄ and CAM systems are completely integrated in Portulaca oleracea, with CAM and C₄ carbon fixation occurring in the same cells and CAM-generated metabolites likely incorporated directly into the C₄ cycle. Flux balance analysis corroborates the gene expression findings and predicts an integrated C₄+CAM system under drought. This first spatially explicit description of a C₄+CAM photosynthetic metabolism presents a potential new blueprint for crop improvement.