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Waterlogging is one of the main abiotic stresses suffered by plants. Inhibition of aerobic respiration during waterlogging limits energy metabolism and restricts growth and a wide range of developmental processes, from seed germination to vegetative growth and further reproductive growth. Plants respond to waterlogging stress by regulating their morphological structure, energy metabolism, endogenous hormone biosynthesis, and signaling processes. In this updated review, we systematically summarize the changes in morphological structure, photosynthesis, respiration, reactive oxygen species damage, plant hormone synthesis, and signaling cascades after plants were subjected to waterlogging stress. Finally, we propose future challenges and research directions in this field.

Drought and waterlogging seriously affect the growth of plants and are considered severe constraints on agricultural and forestry productivity; their frequency and degree have increased over time due to global climate change. The morphology, photosynthetic activity, antioxidant enzyme system and hormone levels of plants could change in response to water stress. The mechanisms of these changes are introduced in this review, along with research on key transcription factors and genes. Both drought and waterlogging stress similarly impact leaf morphology (such as wilting and crimping) and inhibit photosynthesis. The former affects the absorption and transportation mechanisms of plants, and the lack of water and nutrients inhibits the formation of chlorophyll, which leads to reduced photosynthetic capacity. Constitutive overexpression of 9-cis-epoxydioxygenase (NCED) and acetaldehyde dehydrogenase (ALDH), key enzymes in abscisic acid (ABA) biosynthesis, increases drought resistance. The latter forces leaf stomata to close in response to chemical signals, which are produced by the roots and transferred aboveground, affecting the absorption capacity of CO2, and reducing photosynthetic substrates. The root system produces adventitious roots and forms aerenchymal to adapt the stresses. Ethylene (ETH) is the main response hormone of plants to waterlogging stress, and is a member of the ERFVII subfamily, which includes response factors involved in hypoxia-induced gene expression, and responds to energy expenditure through anaerobic respiration. There are two potential adaptation mechanisms of plants (“static” or “escape”) through ETH-mediated gibberellin (GA) dynamic equilibrium to waterlogging stress in the present studies. Plant signal transduction pathways, after receiving stress stimulus signals as well as the regulatory mechanism of the subsequent synthesis of pyruvate decarboxylase (PDC) and alcohol dehydrogenase (ADH) enzymes to produce ethanol under a hypoxic environment caused by waterlogging, should be considered. This review provides a theoretical basis for plants to improve water stress tolerance and water-resistant breeding.

Summary Group VII ethylene response factors (ERFVIIs) play important roles in ethylene signalling and plant responses to flooding. However, natural ERFVII variations in maize (ZmERFVIIs) that are directly associated with waterlogging tolerance have not been reported. Here, a candidate gene association analysis of the ZmERFVII gene family showed that a waterlogging‐responsive gene, ZmEREB180, was tightly associated with waterlogging tolerance. ZmEREB180 expression specifically responded to waterlogging and was up‐regulated by ethylene; in addition, its gene product localized to the nucleus. Variations in the 5ʹ‐untranslated region (5ʹ‐UTR) and mRNA abundance of this gene under waterlogging conditions were significantly associated with survival rate (SR). Ectopic expression of ZmEREB180 in Arabidopsis increased the SR after submergence stress, and overexpression of ZmEREB180 in maize also enhanced the SR after long‐term waterlogging stress, apparently through enhanced formation of adventitious roots (ARs) and regulation of antioxidant levels. Transcriptomic assays of the transgenic maize line under normal and waterlogged conditions further provided evidence that ZmEREB180 regulated AR development and reactive oxygen species homeostasis. Our study provides direct evidence that a ZmERFVII gene is involved in waterlogging tolerance. These findings could be applied directly to breed waterlogging‐tolerant maize cultivars and improve our understanding of waterlogging stress.

Peach ( Prunus persica L. Batsch) is a shallow root fruit tree with poor waterlogging tolerance. Hydrogen sulfide (H2S) is a signal molecule which regulates the adaptation of plants to adverse environments. Nevertheless, the effects of exogenous applications of H2S in fruit tree species especially in peach trees under waterlogging stress have been scarcely researched. Thus, the goal of this research was to investigate the alleviating effect of exogenous H2S on peach seedlings under waterlogging stress. In the present study, we found that the effect of exogenous H2S depended on the concentration and 0.2 mM sodium hydrosulfide (NaHS) showed the best remission effect on peach seedlings under waterlogging stress. Waterlogging significantly reduced the stomatal opening, net photosynthetic rate, and Fv/Fm of peach seedlings. The results of histochemical staining and physiological and biochemical tests showed that waterlogging stress increased the number of cell deaths and amounts of reactive oxygen species (ROS) accumulated in leaves, increased the number of root cell deaths, significantly increased the electrolyte permeability, O2.– production rate, H2O2 content and ethylene synthesis rate of roots, and significantly reduced root activity. With prolonged stress, antioxidative enzyme activity increased initially and then decreased. Under waterlogging stress, application of 0.2 mM NaHS increased the number of stomatal openings, improved the chlorophyll content, and photosynthetic capacity of peach seedlings. Exogenous H2S enhanced antioxidative system and significantly alleviate cell death of roots and leaves of peach seedlings caused by waterlogging stress through reducing ROS accumulation in roots and leaves. H2S can improve the activity and proline content of roots, reduce oxidative damage, alleviated lipid peroxidation, and inhibit ethylene synthesis. The H2S scavenger hypotaurine partially eliminated the effect of exogenous H2S on alleviating waterlogging stress of peach seedlings. Collectively, our results provide an insight into the protective role of H2S in waterlogging-stressed peach seedlings and suggest H2S as a potential candidate in reducing waterlogging-induced damage in peach seedlings.

Background Sesame ( Sesamum indicum L.) is one of the world’s most important oil crops. However, it is susceptible to abiotic stresses in general, and to waterlogging and drought stresses in particular. The molecular mechanisms of abiotic stress tolerance in sesame have not yet been elucidated. The WRKY domain transcription factors play significant roles in plant growth, development, and responses to stresses. However, little is known about the number, location, structure, molecular phylogenetics, and expression of the WRKY genes in sesame. Results We performed a comprehensive study of the WRKY gene family in sesame and identified 71 SiWRKYs . In total, 65 of these genes were mapped to 15 linkage groups within the sesame genome. A phylogenetic analysis was performed using a related species ( Arabidopsis thaliana ) to investigate the evolution of the sesame WRKY genes. Tissue expression profiles of the WRKY genes demonstrated that six SiWRKY genes were highly expressed in all organs, suggesting that these genes may be important for plant growth and organ development in sesame. Analysis of the SiWRKY gene expression patterns revealed that 33 and 26 SiWRKYs respond strongly to waterlogging and drought stresses, respectively. Changes in the expression of 12 SiWRKY genes were observed at different times after the waterlogging and drought treatments had begun, demonstrating that sesame gene expression patterns vary in response to abiotic stresses. Conclusions In this study, we analyzed the WRKY family of transcription factors encoded by the sesame genome. Insight was gained into the classification, evolution, and function of the SiWRKY genes, revealing their putative roles in a variety of tissues. Responses to abiotic stresses in different sesame cultivars were also investigated. The results of our study provide a better understanding of the structures and functions of sesame WRKY genes and suggest that manipulating these WRKYs could enhance resistance to waterlogging and drought.

Waterlogging stress (WS) induces ethylene (ET) and polyamine (spermine, putrescine, and spermidine) production in plants, but their reprogramming is a decisive element for determining the fate of the plant upon waterlogging-induced stress. WS can be challenged by exploring symbiotic microbes that improve the plant’s ability to grow better and resist WS. The present study deals with identification and application of 1-aminocyclopropane-1-carboxylic acid (ACC) deaminase-producing fungal endophyte Trichoderma asperellum (strain MAP1), isolated from the roots of Canna indica L., on wheat growth under WS. MAP1 positively affected wheat growth by secreting phytohormones/secondary metabolites, strengthening the plant’s antioxidant system and influencing the physiology through polyamine production and modulating gene expression. MAP1 inoculation promoted yield in comparison to non-endophyte inoculated waterlogged seedlings. Exogenously applied ethephon (ET synthesis inducer) and 1-aminocyclopropane carboxylic acid (ACC; ET precursor) showed a reduction in growth, compared to MAP1-inoculated waterlogged seedlings, while amino-oxyacetic acid (AOA; ET inhibitor) application reversed the negative effect imposed by ET and ACC, upon waterlogging treatment. A significant reduction in plant growth rate, chlorophyll content, and stomatal conductance was noticed, while H 2 O 2 , MDA production, and electrolyte leakage were increased in non-inoculated waterlogged seedlings. Moreover, in comparison to non-inoculated waterlogged wheat seedlings, MAP1-inoculated waterlogged wheat exhibited antioxidant–enzyme activities. In agreement with the physiological results, genes associated with the free polyamine (PA) biosynthesis were highly induced and PA content was abundant in MAP1-inoculated seedlings. Furthermore, ET biosynthesis/signaling gene expression was reduced upon MAP1 inoculation under WS. Briefly, MAP1 mitigated the adverse effect of WS in wheat, by reprogramming the PAs and ET biosynthesis, which leads to optimal stomatal conductance, increased photosynthesis, and membrane stability as well as reduced ET-induced leaf senescence.

This study aimed to explore the effects of waterlogging priming during the seedling stage on waterlogging tolerance in wheat at the booting stage. Greenhouse pot experiments were conducted for two consecutive years on two wheat cultivars with different response types to waterlogging priming throughout their entire growth period using four treatments: no priming and no waterlogging stress (NC), priming and no waterlogging stress (PC), no priming with waterlogging stress (NW), and priming with waterlogging stress (PW). Through an investigation of the yield, biomass, aboveground photosynthesis, root morphological structure and physiological indices, we confirmed that the effectiveness of waterlogging priming varied by cultivars. For Zhenmai10, seedling-stage waterlogging priming significantly alleviated the yield loss caused by waterlogging at the booting stage. The aboveground photosynthetic efficiency and root growth were maintained. Antioxidant system was further activated to regulate reactive oxygen species homeostasis. The anaerobic respiration process and aerenchyma formation were promoted to improve oxygen supply capacity. However, all these effects appeared to be weak in Yangmai22. Generally, we offered a new perspective on physiological mechanisms of the long-term effects of waterlogging priming from two aspects: enhancing waterlogging tolerance of plants and accelerating the physiological response of roots to waterlogging stress.

Waterlogging stands as a common environmental challenge, significantly affecting plant growth, yield, and, in severe cases, survival. In response to waterlogging stress, plants exhibit a series of intricate physiologic, metabolic, and morphologic adaptations. Notably, the gaseous phytohormone ethylene is rapidly accumulated in the plant submerged tissues, assuming an important regulatory factor in plant-waterlogging tolerance. In this review, we summarize recent advances in research on the mechanisms of ethylene in the regulation of plant responses to waterlogging stress. Recent advances found that both ethylene biosynthesis and signal transduction make indispensable contributions to modulating plant adaptation mechanisms to waterlogged condition. Ethylene was also discovered to play an important role in plant physiologic metabolic responses to waterlogging stress, including the energy mechanism, morphologic adaptation, ROS regulation and interactions with other phytohormones. The comprehensive exploration of ethylene and its associated genes provides valuable insights into the precise strategies to leverage ethylene metabolism for enhancing plant resistance to waterlogging stress.

Waterlogging has negative effects on crop yield. Physiological and transcriptome data of two peanut cultivars [Zhongkaihua 1 (ZKH 1) and Huayu 39 (HY 39)] were studied under normal water supply and waterlogging stress for 5 or 10 days at the flowering stage. The results showed that the main stem height, the number of lateral branches, lateral branch length, and the stem diameter increased under waterlogging stress, followed by an increase in dry matter accumulation, which was correlated with the increase in the soil and plant analysis development (SPAD) and net photosynthetic rate (Pn) and the upregulation of genes related to porphyrin and chlorophyll metabolism and photosynthesis. However, the imbalance of the source–sink relationship under waterlogging was the main cause of yield loss, and waterlogging caused an increase in the sucrose and soluble sugar contents and a decrease in the starch content; it also decreased the activities of sucrose synthetase (SS) and sucrose phosphate synthetase (SPS), which may be due to the changes in the expression of genes related to starch and sucrose metabolism. However, the imbalance of the source–sink relationship led to the accumulation of photosynthate in the stems and leaves, which resulted in the decrease of the ratio of pod dry weight to total dry weight (PDW/TDW) and yield. Compared with ZKH 1, the PDW of HY 39 decreased more probably because more photosynthate accumulated in the stem and leaves of HY 39 and could not be effectively transported to the pod.

Background Morphological plasticity is one of the capacities of plants to modify their morphological appearance in response to external stimuli. A plant’s morphology and physiology are constantly tuned to its variable surroundings by complex interactions between environmental stimuli and internal signals. In most of plant species,, such phenotypic and physiological expression varies among different varieties based on their levels of particular environmental stress conditions. However, the morphological and yield responses of common bean varieties to different environmental conditions are not well known. The purpose of the study was to evaluate morphological and yield response of common bean to soil moisture stress and to investigate the morphological mechanism by which common bean varieties tolerate fluctuations in moisture stress. Methods A pot experiment was carried out to investigate the effects of different moisture levels on the phenotypic and yield responses of common bean varieties. A factorial combination of five common bean varieties (Hirna, kufanzik, Awash-1, Ado, and Chercher) and three moisture levels (control, waterlogging stress, and moisture deficit stress) was used in three replications. Moisture stress treatments were started 20 days after planting, at the trifoliate growth stage. To evaluate the response of each variety, morphological and yield data were collected at week intervals. Main results The results indicated that moisture levels and varieties had a significant influence on all growth parameters. Crop phenology was significantly influenced by the interaction effect of moisture level and variety. Exposing Hirna variety to moisture stress led to extended flowering and pod setting by 23 and 24 days, respectively, compared to the other treatments. The results showed that the phenotypic responses to moisture deficit and waterlogging stress varied between varieties. Waterlogging stress had a stronger reduction effect on the fresh weight, dry weight and leaf area of common bean varieties than moisture deficit and the control. Pods per plant, seeds per plant, grain yield per plant, and harvest index were significantly influenced by the varieties, moisture stress levels and their interaction. Except for Chercher and Hirna. However, varieties Ado, kufanzik and Awasha-1 did not show significant differences on the time of flower initiation due to moisture level. Biomass and growth in leaf fresh weight, leaf dry weight, leaf area, leaf number and plant height were significantly influenced by moisture level. When moisture deficit and waterlogging stress occurred, Ado and Awash-1 were more responsive to moisture stress than Hirna, Chercher, and Kufanzik. Conclusion Hence, Hirna and Kufanzik varieties were found to be tolerant because they produced higher yields than the Chercher, Awash-1, and Ado varieties.