'Against all floods': plant adaptation to flooding stress and combined abiotic stresses.

Current climate change brings with it a higher frequency of environmental stresses, which occur in combination rather than individually leading to massive crop losses worldwide. In addition to, for example, drought stress (low water availability), also flooding (excessive water) can threaten the plant, causing, among others, an energy crisis due to hypoxia, which is responded to by extensive transcriptional, metabolic and growth-related adaptations. While signalling during flooding is relatively well understood, at least in model plants, the molecular mechanisms of combinatorial flooding stress responses, for example, flooding simultaneously with salinity, temperature stress and heavy metal stress or sequentially with drought stress, remain elusive. This represents a significant gap in knowledge due to the fact that dually stressed plants often show unique responses at multiple levels not observed under single stress. In this review, we (i) consider possible effects of stress combinations from a theoretical point of view, (ii) summarize the current state of knowledge on signal transduction under single flooding stress, (iii) describe plant adaptation responses to flooding stress combined with four other abiotic stresses and (iv) propose molecular components of combinatorial flooding (hypoxia) stress adaptation based on their reported dual roles in multiple stresses. This way, more future emphasis may be placed on deciphering molecular mechanisms of combinatorial flooding stress adaptation, thereby potentially stimulating development of molecular tools to improve plant resilience towards multi-stress scenarios.

Endoplasmic reticulum-bound ANAC013 factor is cleaved by RHOMBOID-LIKE 2 during the initial response to hypoxia in Arabidopsis thaliana.

Aerobic reactions are essential to sustain plant growth and development. Impaired oxygen availability due to excessive water availability, e.g., during waterlogging or flooding, reduces plant productivity and survival. Consequently, plants monitor oxygen availability to adjust growth and metabolism accordingly. Despite the identification of central components in hypoxia adaptation in recent years, molecular pathways involved in the very early activation of low-oxygen responses are insufficiently understood. Here, we characterized three endoplasmic reticulum (ER)-anchored Arabidopsis ANAC transcription factors, namely ANAC013, ANAC016, and ANAC017, which bind to the promoters of a subset of hypoxia core genes (HCGs) and activate their expression. However, only ANAC013 translocates to the nucleus at the onset of hypoxia, i.e., after 1.5 h of stress. Upon hypoxia, nuclear ANAC013 associates with the promoters of multiple HCGs. Mechanistically, we identified residues in the transmembrane domain of ANAC013 to be essential for transcription factor release from the ER, and provide evidence that RHOMBOID-LIKE 2 (RBL2) protease mediates ANAC013 release under hypoxia. Release of ANAC013 by RBL2 also occurs upon mitochondrial dysfunction. Consistently, like ANAC013 knockdown lines, rbl knockout mutants exhibit impaired low-oxygen tolerance. Taken together, we uncovered an ER-localized ANAC013-RBL2 module, which is active during the initial phase of hypoxia to enable fast transcriptional reprogramming.

Effects of drought, subsequent waterlogging and redrying on growth, physiology and metabolism of wheat.

With climate change, longer periods without precipitation but also heavy rains will become more frequent. Thus, understanding and predicting the implications of drought-waterlogging-redrying cycles for plants is essential. We examined the effects of such events on wheat (Triticum aestivum). We measured the impacts of subsequent water treatments (drought-waterlogging-redrying) on plant shoot and root biomass, photosynthesis and transpiration, as well as on primary metabolites and transcripts of leaves. Drought and drought followed by waterlogging severely reduced shoot and root biomass. Chlorophyll fluorescence parameters and the CO2 assimilation rate per unit leaf area were not affected by the treatments but, after the redrying phase, plants grown under the stress treatments showed a higher transpiration rate per unit leaf area and a lower instantaneous water use efficiency. Many organic acids of the citrate cycle were less concentrated in leaves of stressed plants, while most amino acids were more concentrated. Transcript analysis of genes involved in signalling and metabolism revealed different expression patterns. While some genes responded only to drought or drought followed by waterlogging, several genes were induced upon both treatments and some were still upregulated at the end of the redrying phase. We provide insights into how wheat responds to changes in water regimes, with some of the changes probably allowing the plants to cope with these stressors, at least to a certain degree.