Developing stress-tolerant, resource-efficient crops is a key strategy for addressing the challenges of climate change, global food security, and land degradation (Blum & Jordan, 1985; Hunter et al., 2017; Mickelbart et al., 2015). Maize is a critical global crop, cultivated for food, fuel, and industrial uses (FAO, 2018). In intensive agriculture systems, nitrogen (N) fertilizers are over-applied to maximize grain yield, yet over half of the applied nitrate leaches beyond the root zone and pollutes waterways, or is volatilized as greenhouse gases (Dhital & Raun, 2016; Hirel et al., 2011). In low-input subsistence agriculture, which sustains half of the global population, maize is grown