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Nitrogen is the most limiting nutrient in most agro-ecosystems and thus critical for sustaining high yields. Conventional agricultural practices use synthetic fertilizers to ensure an adequate supply of nitrogen in soils, but fertilizers come at a significant monetary and environmental cost. A strategy to improve nitrogen supply in cropping systems is the inclusion of nitrogen-fixing legumes, which can provide nitrogen benefits to companion crops through belowground nitrogen transfer. However, a better understanding of the underlying mechanisms and factors that govern nitrogen transfer is important in order to determine potential areas for improving this association. Here, we review the mechanisms of belowground nitrogen transfer in managed herbaceous cropping systems, focusing on forage systems. We classify three major routes of nitrogen transfer from legumes to non-legumes: (1) decomposition of legume root tissues and uptake of mineralized nitrogen by neighboring plants, (2) exudation of soluble nitrogen compounds by legumes and uptake by non-legumes, and (3) transfer of nitrogen mediated by plant-associated mycorrhizae. Literature data shows that rates of nitrogen transfer range from 0 to 73 % from forage legumes to companion grasses in mixed stands, depending on the legume species and cultivar. We list the factors that affect nitrogen transfer including abiotic factors, e.g., water stress, temperature, light, soil available nitrogen, and application of nitrogen fertilizer, and biotic factors, e.g., root contact, plant density, growth stage, production year, defoliation, and root herbivores. While the rates of nitrogen transfer are often constrained by abiotic conditions, such as temperature and water availability, that are beyond the control of growers, agronomic practices, e.g., planting density and choice of species and cultivar, may help to increase nitrogen transfer. Ultimately, the selection of plant pairs with compatible traits offers the best path forward to improving nitrogen transfer in intercrops.

Belowground nitrogen transfer from legumes to non-legumes under managed herbaceous cropping systems. A review