Within leaf nitrogen allocation regulates the photosynthetic behavior of xerophytes in response to increased soil rock fragment content

There is limited information on how plant functional traits vary with soil rock fragment content (RFC), especially for xerophytes growing in stony soils. We examined leaf functional traits of three xerophytes (Sophora davidii; Cotinus szechuanensi; and Artemisia vestita) grown under an RFC gradient in a heavy loamy soil. Our results show that photosynthetic capacity increased linearly with RFC in S. davidii, whereas unimodal patterns were observed for the other two species. The RFC that maximized photosynthetic capacity (Asat) and photosynthetic N use efficiency (PNUE) were achieved by allocating more N to photosynthetic apparatus at the expense of cell walls. For C. szechuanensis, the increased fraction of photosynthetic N allocated to carboxylation (PC) bioenergetics (PB), and thylakoid light-harvesting components (PL) together contributed to the higher Asat and PNUE values. As for S. davidii, both PC and PB mainly contributed to higher Asat and PNUE, whereas for A. vestita only PB was the main contributor. Our results suggest that increased non-capillary porosity of high RFC soil conditions through promoting the root growth of S. davidii and C. szechuanensis ensures sufficient water and N supply for photosynthetic capacity. In shallow-rooted species A. vestita, low RFC soil maintained higher nitrate N in the topsoil, enhancing leaf photosynthetic capacity. We conclude that rock fragments promoted leaf photosynthetic capacity in the studied loamy soil system, but the promoting effect was species-specific. The results highlight the relevance of consideration of soil rock fraction in evaluation of photosynthetic behavior of xerophytes in heterogeneous rocky soils.