Topography-driven differences in soil N transformation constrain N availability in karst ecosystems

Fragile karst ecosystems are characterized by complex topographic landscapes associated with high variations in vegetation restoration. Identifying the characteristics and driving factors of nitrogen (N) availability across the topographic gradient is essential to guide vegetation restoration in karst regions. In this study, we collected soil samples and plant leaves along the topographic gradient (ridge, upper slope, middle slope, and foot slope) of convex slopes in the karst fault basin of southwest China, and determined the indicators reflecting soil N availability, N transformation rates, and their controlling factors. Our results showed that foliar N content and δ¹⁵N value, soil inorganic N content and δ¹⁵N value, and foliar N:P ratio were substantially lower on the steep hillslopes than on the flat top ridge. Steep slope soils also had a lower enzyme C:N ratio but a higher enzyme N:P ratio than the flat ridge soils. Furthermore, the vector angles calculated by soil extracellular enzyme analysis were below 45^o in all studied soils and decreased significantly with increasing slope, indicating that microbial growth was generally limited by N. These results jointly suggest the declines in soil N availability across the topographic gradient, which are further explained by the changes in soil inherent N transformation processes. As the slope became steeper, soil mineralization and autotrophic nitrification (O_NH4) rates decreased significantly, while ratio of microbial NH₄⁺ immobilization to O_NH4 and NH₄⁺ adsorption rate increased significantly, indicating the decrease in soil inorganic N supply capacity. We further found that deteriorated soil structure, decreased soil organic matter and calcium content, altered microbial abundance, and increased ratios of fungi to bacteria and gram-positive bacteria to gram-negative bacteria were the primary drivers of reduced N transformation rates and N availability across the topographic gradient. Overall, this study highlights the critical role of the topography in controlling soil N availability by regulating N transformation processes in karst regions. The topography should be considered an important factor affecting the functions and services of karst ecosystems.