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The evolution of Crassulacean acid metabolism (CAM) is thought to be along a C 3 -CAM continuum including multiple variations of CAM such as CAM cycling and CAM idling. Here, we applied large-scale constraint-based modeling to investigate the metabolism and energetics of plants operating in C 3 , CAM, CAM cycling, and CAM idling. Our modeling results suggested that CAM cycling and CAM idling could be potential evolutionary intermediates in CAM evolution by establishing a starch/sugar-malate cycle. Our model analysis showed that by varying CO 2  exchange during the light period, as a proxy of stomatal conductance, there exists a C 3 -CAM continuum with gradual metabolic changes, supporting the notion that evolution of CAM from C 3  could occur solely through incremental changes in metabolic fluxes. Along the C 3 -CAM continuum, our model predicted changes in metabolic fluxes not only through the starch/sugar-malate cycle that is involved in CAM photosynthetic CO 2  fixation but also other metabolic processes including the mitochondrial electron transport chain and the tricarboxylate acid cycle at night. These predictions could guide engineering efforts in introducing CAM into C 3  crops for improved water use efficiency.

Metabolic Modeling of the C3-CAM Continuum Revealed the Establishment of a Starch/Sugar-Malate Cycle in CAM Evolution