| Summary: | Abstract Understanding the mechanisms underlying variations in carbon isotope discrimination (Δ) in C4 plants is critical for predicting the C3/C4 ratio in C3/C4 mixed grassland. The value of Δ is determined by bundle sheath leakiness (Ф) and the ratio of intercellular to ambient CO2 concentration (C i /C a ). Leaf nitrogen concentration (N leaf ) is considered a driver of Δ in C4 plants. However, little is known about how N leaf affects Ф and C i /C a , and subsequently Δ. Here leaf carbon isotope composition, N leaf , Ф, and leaf gas exchange were measured in Cleistogenes squarrosa, a dominant C4 species in the Inner Mongolia grassland. Δ remained relatively stable under variable N and water supply. Higher N supply and lower water supply increased N leaf , stimulated photosynthesis and further decreased C i /C a . High N supply increased Ф, which responded weakly to water supply. N leaf exerted similar effects on C i /C a and on Ф in the field and pot experiments. Pooling all the data, N leaf explained 73% of the variation in C i /C a . Overall, both Ф and C i /C a determined Δ; however, the contribution of Ф was stronger. N leaf influenced Δ primarily though C i /C a , rather than Ф. Ф should be considered in estimating Δ of C4 endmember.
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