Elevated CO2 improves yield and N2 fixation but not grain N concentration of faba bean (Vicia faba L.) subjected to terminal drought

Abstract

Legumes grown in Mediterranean environments frequently experience terminal drought which reduces yield and N2 fixation processes. Decreased N2 fixation during reproductive phases may constrain seed nitrogen concentrations ([N]), reducing protein concentration of grain legumes. Plants grown under elevated atmospheric CO2 concentrations ([CO2]) have greater water use efficiency. This may result in reduced use of conserved/stored soil water, potentially helping to reduce soil water deficits later during grain filling. The extent that this process applies to drought sensitive grain legumes, which are extensively cultivated in Mediterranean environments is unclear. The objectives of this study were to investigate yield, N2 fixation and seed N response of faba bean (Vicia faba L. cv. ‘Fiesta’) grown in a dryland Mediterranean-type environment under elevated [CO2]. Plants were grown in soil columns under ambient [CO2] (˜400 ppm) or elevated [CO2] (e[CO2], ˜550 ppm) in a Free-Air CO2 Enrichment (FACE) facility in the field. One sub-group was continuously well-watered (80% field capacity, FC), whereas a second sub-group was exposed to a drought treatment (water was withheld until 30% FC was reached, which was then maintained during the reproductive phases). Biomass, gas exchange, 13C isotopic discrimination, N2 fixation by the natural abundance 15N method, nodulation and soil water content were assessed throughout the crop developmental stages.Initially, plants grown under elevated [CO2] depleted soil water more slowly in the drought treatment than those under ambient [CO2], but as plants grown under elevated [CO2] produced more biomass they used soil water more rapidly, especially towards the critical pod-filling phase. Water savings during the first phase of the drought treatment, through flowering up to the start of pod-filling, were associated with increased yield (+25%) and N2 fixation (+15%) under drought. Elevated [CO2]-induced stimulation of nodulation and nodule density helped maintain N2 fixation under drought, even though nodule activity decreased under the combined effect of e[CO2] and drought from pod-filling onwards. This later stage decrease was associated with decreased carbohydrate and increased amino acid concentrations in nodules, indicating a down-regulation of N2 fixation. Associated with the decrease of N2 fixation during pod-filling, seed N concentration was lower under the combination of e[CO2] and drought. We propose a conceptual model to explain the importance of N2 fixation during the grain filling stage to maintain seed N concentration under e[CO2]. These findings suggest that e[CO2]-induced savings in soil water may mitigate negative effects of drought on yield and N2 fixation of faba bean, without fully compensating the effect of prolonged drought on seed N concentration.