Nitrogen (N) is one of the most limiting nutrients for plant growth worldwide (Elser et al., 2007). It enters the biosphere primarily by biological N fixation, atmospheric reactive N deposition, and application of N fertilizers (Galloway et al., 2008;Peñuelas et al., 2013). After plant uptake, it is returned to soil in organic forms. Mineralization of organic N in soil is the main process of recycling organic N into mineral forms, which strongly drives N availability, distribution between biological and non-biological pools, and the cascading pathways of N leaving terrestrial ecosystems (Booth et al., 2005;Dai et al., 2020;Fang et al., 2015). For instance, ammonium (NH 4 + ) and nitrate (NO 3 − ) are favorable forms of N taken up by plants and microbes, and are mainly replenished by ammonification and nitrification processes, respectively, with the two processes covarying with each other and fundamentally driving organic N recycling (Figure S1 in Supporting Information S1;Booth et al., 2005). On the other hand, gaseous N emissions from denitrification and nitrification, and NO 3 − leaching, are essential ecosystem N output pathways that can strongly reduce plant N availability and the soil total nitrogen (TN) pool (Figure