The effects of mineral nitrogen limitation, competition, arbuscular mycorrhiza, and their respective interactions, on morphological and chemical plant traits of Plantago lanceolata

Plants are sessile organisms that suffer from a multitude of challenges such as abiotic stress or the interactions with competitors, antagonists and symbionts, which influence their performance as well as their eco-physiological and biochemical responses in complex ways. In particular, the combination of different stressors and their impact on plant biomass production and the plant's ability to metabolically adjust to these challenges are less well understood. To study the effects of mineral nitrogen (N) availability, interspecific competition and the association with arbuscular mycorrhizal fungi (AMF) on biomass production, biomass allocation patterns (root/shoot ratio, specific leaf area) and metabolic responses, we chose the model organism Plantago lanceolata L. (Plantaginaceae). Plants were grown in a full factorial experiment. Biomass production and its allocation patterns were assessed at harvest, and the influence of the different treatments and their interactions on the plant metabolome were analysed using a metabolic fingerprinting approach with ultra-high performance liquid chromatography coupled with time-of-flight-mass spectrometry. Limited supply of mineral N caused the most pronounced changes with respect to plant biomass and biomass allocation patterns, and altered the concentrations of more than one third of the polar plant metabolome. Competition also impaired plant biomass production, yet affected the plant metabolome to a much lesser extent than limited mineral N supply. The interaction of competition and limited mineral N supply often caused additive changes on several traits. The association with AMF did not enhance biomass production, but altered biomass allocation patterns such as the root/shoot ratio and the specific leaf area. Interestingly, we did not find significant changes in the plant metabolome caused by AMF. A targeted analysis revealed that only limited mineral N supply reduced the concentrations of one of the main target defence compounds of P. lanceolata, the iridoid glycoside catalpol. In general, the interaction of competition and limited mineral N supply led to additive changes, while the association with AMF in any case alleviated the observed stress responses. Our results show that the joint analysis of biomass/allocation patterns and metabolic traits allows a more comprehensive interpretation of plant responses to different biotic and abiotic challenges; specifically, when multiple stresses interact. (C) 2015 Elsevier Ltd. All rights reserved.