The C 4 photosynthetic pathway is nearly always underpinned by characteristic Kranz anatomy, whereby vascular bundles are closely spaced and are often separated by only two mesophyll (M) cells (reviewed in Sedelnikova, Hughes, & Langdale, 2018). This cellular arrangement enables function of the C 4 cycle, in which CO 2 is initially fixed in the outer M cells by phosphoenolpyruvate carboxylase (PEPC) into the 4-carbon compound oxaloacetate. Although there are at least three distinct C 4 enzymatic cycles (Furbank, 2011), all involve a 4-carbon derivative of oxaloacetate being shuttled to the inner bundle-sheath (BS) cells to be decarboxylated. The CO 2 released in the BS cells is refixed by ribulose bisphosphate carboxylase/oxygenase (RuBisCO) in the Calvin-Benson (C 3) cycle. Importantly, refixation occurs in a cellular environment where local levels of CO 2 are high enough to suppress the competing oxygenation reaction and thus to prevent the energetically wasteful process of photorespiration. As compared to