When and where soil is important to modify the carbon and water economy of leaves

Photosynthetic ‘least-cost’ theory posits that the optimal trait combination for a given environment is that where the summed costs of photosynthetic water and nutrient acquisition/use are minimised. The effects of soil water and nutrient availability on photosynthesis should be stronger as climate-related costs for both resources increase. 

Two independent datasets of photosynthetic traits, Globamax (1509 species, 288 sites) and Glob13C (3645 species, 594 sites), were used to quantify biophysical and biochemical limitations of photosynthesis and the key variable C_i/C_a (CO₂ drawdown during photosynthesis). Climate and soil variables were associated with both datasets. 

The biochemical photosynthetic capacity was higher on alkaline soils. This effect was strongest at more arid sites, where water unit-costs are presumably higher. Higher values of soil silt and depth increased C_i/C_a, likely by providing greater H₂O supply, alleviating biophysical photosynthetic limitation when soil water is scarce. 

Climate is important in controlling the optimal balance of H₂O and N costs for photosynthesis, but soil properties change these costs, both directly and indirectly. In total, soil properties modify the climate-demand driven predictions of C_i/C_a by up to 30% at a global scale.