Background Knowledge of biological and climatic controls in terrestrial nitrogen (N) cycling within and across ecosystems is central to understanding global patterns of key ecosystem processes. The ratios of 15 N: 14 N in plants and soils have been used as indirect indices of N cycling parameters, yet our understanding of controls over N isotope ratios in plants and soils is still developing. Scope In this review, we provide background on the main processes that affect plant and soil N isotope ratios. In a similar manner to partitioning the roles of state factors and interactive controls in determining ecosystem traits, we review N isotopes patterns in plants and soils across a number of proximal factors that influence ecosystem properties as well as mechanisms that affect these patterns. Lastly, some remaining questions that would improve our understanding of N isotopes in terrestrial ecosystems are highlighted. Conclusion Compared to a decade ago, the global patterns of plant and soil N isotope ratios are more resolved. Additionally, we better understand how plant and soil N isotope ratios are affected by such factors as mycorrhizal fungi, climate, and microbial processing. A comprehensive understanding of the N cycle that ascribes different degrees of isotopic fractionation for each step under different conditions is closer to being realized, but a number of process-level questions still remain.

Quantifying global patterns of terrestrial nitrogen (N) cycling is central to predicting future patterns of primary productivity, carbon sequestration, nutrient fluxes to aquatic systems, and climate forcing. With limited direct measures of soil N cycling at the global scale, syntheses of the 15N:14N ratio of soil organic matter across climate gradients provide key insights into understanding global patterns of N cycling. In synthesizing data from over 6000 soil samples, we show strong global relationships among soil N isotopes, mean annual temperature (MAT), mean annual precipitation (MAP), and the concentrations of organic carbon and clay in soil. In both hot ecosystems and dry ecosystems, soil organic matter was more enriched in 15N than in corresponding cold ecosystems or wet ecosystems. Below a MAT of 9.8°C, soil δ15N was invariant with MAT. At the global scale, soil organic C concentrations also declined with increasing MAT and decreasing MAP. After standardizing for variation among mineral soils in soil C and clay concentrations, soil δ15N showed no consistent trends across global climate and latitudinal gradients. Our analyses could place new constraints on interpretations of patterns of ecosystem N cycling and global budgets of gaseous N loss.

The representation of the nitrogen (N) cycle in Earth system models (ESMs) is strongly motivated by the constraint N poses on the sequestration of anthropogenic carbon (C). Models typically implement a stoichiometric relationship between C and N in which external supply and assimilation by organisms are adjusted to maintain their internal stoichiometry. N limitation of primary productivity thus occurs if the N supply from uptake and fixation cannot keep up with the construction of tissues allowed by C assimilation. This basic approach, however, presents considerable challenges in how to faithfully represent N limitation. Here, we review how N limitation is currently implemented and evaluated in ESMs and highlight challenges and opportunities in their future development. At or near steady state, N limitation is governed by the magnitude of losses from the plant-unavailable pool vs. N fixation and there are considerable differences in how models treat both processes. In nonsteady-state systems, the accumulation of N in pools with slow turnover rates reduces N available for plant uptake and can be challenging to represent when initializing ESM simulations. Transactional N limitation occurs when N is incorporated into various vegetation and soil pools and becomes available to plants only after it is mineralized, the dynamics of which depends on how ESMs represent decomposition processes in soils. Other challenges for ESMs emerge when considering seasonal to interannual climatic oscillations as they create asynchronies between C and N demand, leading to transient alternations between N surplus and deficit. Proper evaluation of N dynamics in ESMs requires conceptual understanding of the main levers that trigger N limitation, and we highlight key measurements and observations that can help constrain these levers. Two of the biggest challenges are the mechanistic representation of plant controls on N availability and turnover, including N fixation and organic matter decomposition processes.

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