This paper reviews the interactions between water and nitrogen from physiological, agronomic, economic, breeding and modelling perspectives. Our primary focus is wheat; we consider forage crops, sorghum and legumes where relevant aspects of water–nitrogen interactions have been advanced.
From a physiological perspective, we ask: How does nitrogen deficit influence the water economy of the crop? How does water deficit influence the nitrogen economy of the crop? How do combined water and nitrogen deficit affect crop growth and yield? We emphasise synergies, and the nitrogen-driven trade-off between the efficiency in the use of water and nitrogen. The concept of nitrogen–water co-limitation is discussed briefly.
From agronomic and economic perspectives, the need to match supply of nitrogen and water is recognised, but this remains a challenge in dryland systems with uncertain rainfall. Under-fertilisation commonly causes gaps between actual and water-limited potential yield. We discuss risk aversion and the role of seasonal rainfall forecasts to manage risk.
From a breeding perspective, we ask how selection for yield has changed crop traits relating to water and nitrogen. Changes in nitrogen traits are more common and profound than changes in water-related traits. Comparison of shifts in the wheat phenotype in Australia, UK, Argentina and Italy suggests that improving yield per unit nitrogen uptake is straightforward; it requires selection for yield and allowing grain protein concentration to drift unchecked. A more interesting proposition is to increase nitrogen uptake to match yield gains and conserve protein in grain. Increased stomatal conductance is a conspicuous response to selection for yield which partially conflicts with the perception that reduced conductance at high vapour pressure deficit is required to increase water- use efficiency; but high stomatal conductance at high vapour pressure deficit may be adaptive for thermal stress.
From a modelling perspective, water and nitrogen are linked in multiple ways. In crops where water limits growth, reduced biomass reduces nitrogen demand. Reciprocally, nitrogen limitation during crop expansion reduces leaf area index and increases the soil evaporation : transpiration ratio. Water–nitrogen interactions are also captured in the water-driven uptake of nitrogen by mass flow and diffusion and in the water-driven processes of nitrogen in soil (e.g. mineralisation).
The paper concludes with suggestions for future research on water-nitrogen interactions.
