The Photo-3 model: A Python-based model for C3, C4, and CAM photosynthesis coupled with environmental conditions

The ability to depict all three photosynthetic types (C3, C4, and CAM) has important implications for the study of both natural and agroecosystems. Currently no model exists which covers all types of photosynthesis in a consistent way and which can be fully integrated with environmental conditions. This is partially because, despite the fact that Crassulacean acid metabolism (CAM) photosynthesis is prevalent in many plants in arid and semi-arid ecosystems, where it may comprise nearly 50% of all plant biomass, CAM modelling remains understudied. The Photo-3 model takes advantage of recent advances in mechanistic modeling of CAM photosynthesis to provide a direct comparison of CAM functioning with C3 and C4 functioning under a wide range of soil and atmospheric conditions. The model is based on a core Farquhar photosynthetic model with additional functions to represent the spatial and temporal separations of carbon uptake and assimilation in the case of C4 and CAM photosynthesis. We have parameterized the model for one representative species of each photosynthetic type: Opuntia ficus-indica (CAM), Sorghum bicolor (C4), and Triticum aestivum (C3). Results agree well with experimental data on carbon assimilation and water use for the three species. Model runs using climate data from Temple, TX; Sicily, Italy; Zacatecas, Mexico; Pernambuco, Brazil and Adias Ababa, Ethiopia illustrate the high water use efficiency of CAM plants and its cumulative effects on long-term productivity in water-limited environments. The Photo-3 model, which is written in Python, will be made publicly available on GitHub and its outputs may be coupled to existing models of plant growth and phenology. The model may be used to evaluate potential productivity and water use for C3, C4, and CAM plants, and to devise optimal strategies for cropping systems and irrigation in water-limited environments.