Role of Microbes in Alleviating Crop Drought Stress: A Review.

Drought stress is an annual global phenomenon that has devastating effects on crop production, so numerous studies have been conducted to improve crop drought resistance. Plant-associated microbiota play a crucial role in crop health and growth; however, we have a limited understanding of the key processes involved in microbiome-induced crop adaptation to drought stress. In this review, we summarize the adverse effects of drought stress on crop growth in terms of germination, photosynthesis, nutrient uptake, biomass, and yield, with a focus on the response of soil microbial communities to drought stress and plant-microbe interactions under drought stress. Moreover, we review the morpho-physiological, biochemical, and molecular mechanisms underlying the mitigation effect of microbes on crop drought stress. Finally, we highlight future research directions, including the characterization of specific rhizosphere microbiome species with corresponding root exudates and the efficiency of rhizobacteria inoculants under drought conditions. Such research will advance our understanding of the complex interactions between crops and microbes and improve crop resistance to drought stress through the application of beneficial drought-adaptive microbes.

Nitrate Stabilizes the Rhizospheric Fungal Community to Suppress Fusarium Wilt Disease in Cucumber.

Nitrogen forms can regulate soil-borne Fusarium wilt suppression, but the related mechanisms are largely unknown, especially possible action via the rhizospheric microbial community. Soil analysis, MiSeq high-throughput sequencing analysis, community diversity, and network analysis were used to characterize the impact of different nitrogen forms (nitrate and ammonium) on rhizospheric fungal communities and the contribution of nitrate to the suppression to Fusarium oxysporum f. sp. cucumerinum compared with ammonium. Nitrate-grown cucumber showed a lower disease index and F. oxysporum f. sp. cucumerinum abundance in the rhizosphere. In comparisons with ammonium nutrients, nitrate-fed plants maintained a higher soil rhizosphere pH, microbial biomass carbon content, microbial biomass nitrogen content, as well as fungal community richness and diversity following F. oxysporum f. sp. cucumerinum incubation. All these factors were negatively related with disease index. Network analysis showed fewer nodes and edges in the ammonium treatments compared with nitrate treatments. The relative abundance of Pathotroph-Saprotroph, Pathotroph-Saprotroph-Symbiotroph, and Saprotroph fungi explained 82% of the variability of rhizosphere F. oxysporum f. sp. cucumerinum abundance. In conclusion, after pathogen inoculation under nitrate nutrition, the less-affected microbial composition, community diversity, and community internal relations, which resulted from the more diverse and robust microbial population, potentially contributed to greater Fusarium wilt suppression.

Plant Primary Metabolism Regulated by Nitrogen Contributes to Plant-Pathogen Interactions.

Nitrogen contributes to plant defense responses by the regulation of plant primary metabolism during plant-pathogen interactions. Based on biochemical, physiological, bioinformatic and transcriptome approaches, we investigated how different nitrogen forms (ammonium vs. nitrate) regulate the physiological response of cucumber (Cucumis sativus) to Fusarium oxysporum f. sp. cucumerinum (FOC) infection. The metabolic profile revealed that nitrate-grown plants accumulated more organic acids, while ammonium-grown plants accumulated more amino acids; FOC infection significantly increased levels of both amino acids and organic acids in the roots of ammonium-grown plants. Transcriptome analysis showed that genes related to carbon metabolism were mostly up-regulated in plants grown with nitrate, whereas in ammonium-grown plants the up-regulated genes were mostly those that were related to primary nitrogen metabolism. Root FOC colonization and disease incidence were positively correlated with levels of root amino acids and negatively correlated with levels of root organic acids. In conclusion, organic acid metabolism and expression of related genes increased under nitrate, whereas ammonium increased the level of amino acids and expression of related genes; these altered levels of organic acids and amino acids resulted in different tolerances to FOC infection depending on the nitrogen forms supplied.

Wilted cucumber plants infected by Fusarium oxysporum f. sp. cucumerinum do not suffer from water shortage.

Background and Aims: Fusarium wilt is primarily a soil-borne disease and results in yield loss and quality decline in cucumber (Cucumis sativus). The main symptom of fusarium wilt is the wilting of entire plant, which could be caused by a fungal toxin(s) or blockage of water transport. To investigate whether this wilt arises from water shortage, the physiological responses of hydroponically grown cucumber plants subjected to water stress using polyethylene glycol (PEG, 6000) were compared with those of plants infected with Fusarium oxysporum f. sp. cucumerinum (FOC). Methods: Parameters reflecting plant water status were measured 8d after the start of treatment. Leaf gas exchange parameters and temperature were measured with a LI-COR portable open photosynthesis system and by thermal imaging. Chlorophyll fluorescence and chloroplast structures were assessed by imaging pulse amplitude-modulated fluorometry and transmission electron microscopy, respectively. Key Results: Cucumber water balance was altered after FOC infection, with decreased water absorption and hydraulic conductivity. However, the responses of cucumber leaves to FOC and PEG differed in leaf regions. Under water stress, measures of lipid peroxidation (malondialdehyde) and chlorophyll fluorescence indicated that the leaf edge was more seriously injured, with a higher leaf temperature and disrupted leaf water status compared with the centre. Here, abscisic acid (ABA) and proline were negatively correlated with water potential. In contrast, under FOC infection, membrane damage and a higher temperature were observed in the leaf centre while ABA and proline did not vary with water potential. Cytologically, FOC-infected cucumber leaves exhibited circular chloroplasts and swelled starch grains in the leaf centre, in which they again differed from PEG-stressed cucumber leaves. Conclusions: This study illustrates the non-causal relationship between fusarium wilt and water transport blockage. Although leaf wilt occurred in both water stress and FOC infection, the physiological responses were different, especially in leaf spatial distribution.

Nitrate Increased Cucumber Tolerance to Fusarium Wilt by Regulating Fungal Toxin Production and Distribution.

Cucumber Fusarium wilt, induced by Fusarium oxysporum f. sp. cucumerinum (FOC), causes severe losses in cucumber yield and quality. Nitrogen (N), as the most important mineral nutrient for plants, plays a critical role in plant-pathogen interactions. Hydroponic assays were conducted to investigate the effects of different N forms (NH4⁺ vs. NO3‒) and supply levels (low, 1 mM; high, 5 mM) on cucumber Fusarium wilt. The NO3‒-fed cucumber plants were more tolerant to Fusarium wilt compared with NH4⁺-fed plants, and accompanied by lower leaf temperature after FOC infection. The disease index decreased as the NO3‒ supply increased but increased with the NH4⁺ level supplied. Although the FOC grew better under high NO3- in vitro, FOC colonization and fusaric acid (FA) production decreased in cucumber plants under high NO3- supply, associated with lower leaf membrane injury. There was a positive correlation between the FA content and the FOC number or relative membrane injury. After the exogenous application of FA, less FA accumulated in the leaves under NO3- feeding, accompanied with a lower leaf membrane injury. In conclusion, higher NO3- supply protected cucumber plants against Fusarium wilt by suppressing FOC colonization and FA production in plants, and increasing the plant tolerance to FA.

Nitrate Protects Cucumber Plants Against Fusarium oxysporum by Regulating Citrate Exudation.

Fusarium wilt causes severe yield losses in cash crops. Nitrogen plays a critical role in the management of plant disease; however, the regulating mechanism is poorly understood. Using biochemical, physiological, bioinformatic and transcriptome approaches, we analyzed how nitrogen forms regulate the interactions between cucumber plants and Fusarium oxysporum f. sp. cucumerinum (FOC). Nitrate significantly suppressed Fusarium wilt compared with ammonium in both pot and hydroponic experiments. Fewer FOC colonized the roots and stems under nitrate compared with ammonium supply. Cucumber grown with nitrate accumulated less fusaric acid (FA) after FOC infection and exhibited increased tolerance to chemical FA by decreasing FA absorption and transportation in shoots. A lower citrate concentration was observed in nitrate-grown cucumbers, which was associated with lower MATE (multidrug and toxin compound extrusion) family gene and citrate synthase (CS) gene expression, as well as lower CS activity. Citrate enhanced FOC spore germination and infection, and increased disease incidence and the FOC population in ammonium-treated plants. Our study provides evidence that nitrate protects cucumber plants against F. oxysporum by decreasing root citrate exudation and FOC infection. Citrate exudation is essential for regulating disease development of Fusarium wilt in cucumber plants.