Drought and flooding are two major causes of severe yield loss in soybean worldwide. A lack of knowledge of the molecular mechanisms involved in drought and flood stress has been a limiting factor for the effective management of soybeans; therefore, it is imperative to assess the expression of genes involved in response to flood and drought stress. In this study, differentially expressed genes (DEGs) under drought and flooding conditions were investigated using Illumina RNA-Seq transcriptome profiling. A total of 2724 and 3498 DEGs were identified under drought and flooding treatments, respectively. These genes comprise 289 Transcription Factors (TFs) representing Basic Helix-loop Helix (bHLH), Ethylene Response Factors (ERFs), myeloblastosis (MYB), No apical meristem (NAC), and WRKY amino acid motif (WRKY) type major families known to be involved in the mechanism of stress tolerance. The expression of photosynthesis and chlorophyll synthesis related genes were significantly reduced under both types of stresses, which limit the metabolic processes and thus help prolong survival under extreme conditions. However, cell wall synthesis related genes were up-regulated under drought stress and down-regulated under flooding stress. Transcript profiles involved in the starch and sugar metabolism pathways were also affected under both stress conditions. The changes in expression of genes involved in regulating the flux of cell wall precursors and starch/sugar content can serve as an adaptive mechanism for soybean survival under stress conditions. This study has revealed the involvement of TFs, transporters, and photosynthetic genes, and has also given a glimpse of hormonal cross talk under the extreme water regimes, which will aid as an important resource for soybean crop improvement.

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Identification and Comparative Analysis of Differential Gene Expression in Soybean Leaf Tissue under Drought and Flooding Stress Revealed by RNA-Seq.

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.

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Mechanisms of Waterlogging Tolerance in Plants: Research Progress and Prospects

We projected meta-QTL (MQTL) for drought, salinity, and waterlogging tolerance to the physical map of barley through meta-analysis. The positions of these MQTL were refined and candidate genes were identified.
 Drought, salinity and waterlogging are three major abiotic stresses limiting barley yield worldwide. Breeding for abiotic stress-tolerant crops has drawn increased attention, and a large number of quantitative trait loci (QTL) for drought, salinity, and waterlogging tolerance in barley have been detected. However, very few QTL have been successfully used in marker-assisted selection (MAS) in breeding. In this study, we summarized 632 QTL for drought, salinity and waterlogging tolerance in barley. Among all these QTL, only 195 major QTL were used to conduct meta-analysis to refine QTL positions for MAS. Meta-analysis was used to map the summarized major QTL for drought, salinity, and waterlogging tolerance from different mapping populations on the barley physical map. The positions of identified meta-QTL (MQTL) were used to search for candidate genes for drought, salinity, and waterlogging tolerance in barley. Both MQTL3H.4 and MQTL6H.2 control drought tolerance in barley. Fine-mapped QTL for salinity tolerance, HvNax4 and HvNax3, were validated on MQTL1H.4 and MQTL7H.2, respectively. MQTL2H.1 and MQTL5H.3 were also the target regions for improving salinity tolerance in barley. MQTL4H.4 is the main region controlling waterlogging tolerance in barley with fine-mapped QTL for aerenchyma formation under waterlogging conditions. Detected and refined MQTL and candidate genes are crucial for future successful MAS in barley breeding.

Meta-analysis of major QTL for abiotic stress tolerance in barley and implications for barley breeding.

Despite the heat-related physiology and heat-shock proteins in maize have been extensively studied, little is known about the transcriptome profiling of how the maize varieties with different genotypes responding to high temperatures. Seedling mortality of Xiantian 5 (XT) is significantly lower than that of Zhefengtian (ZF) when exposed to high temperature (42 °C for 6 h) and followed by a recovery growth (25 °C for one week). Therefore, we performed a transcriptome analysis using the total RNA extracted from the leaves of XT and ZF that were previously subjected to heat stress at 42 °C for 0 h, 0.5 h, and 3 h, respectively. A total of 516 commonly up-regulated and 1,261 commonly down-regulated genes were identified among XT/ZF, XT0.5/ZF0.5 and XT3/ZF3 using transcriptome analysis. Gene Ontology classification of the 516 up-regulated genes showed that their encoded proteins were significantly assigned to 18 cellular components, and were classified into 9 functional categories, and were involved in 9 biological processes. Most of proteins encoded by up-regulated genes were localized in chloroplast and its structural components, and involved in multiple biological processes associated with photosynthesis, indicating that these chloroplast proteins play an important role in increasing heat tolerance in sweet maize. While the proteins encoded by 1,261 down-regulated genes were significantly assigned to 31 cellular components, and were classified into 3 functional categories, and were involved in 9 biological processes. Interestingly, these proteins were involved in a series of biological processes from gene expression to translation, suggesting that lowering these processes may contribute to improved heat resistance in sweet maize. The up-regulated genes were identified to be involved in 36 distinct metabolic pathways, of which the most significant ones was secondary metabolite biosynthetic pathway. While the down-regulated genes were identified to be involved in 23 distinct metabolic pathways, of which the most significant ones were found in ribosome. Quantitative real-time PCR analysis demonstrated that 5 genes involved in the biosynthesis of secondary metabolites and photosynthesis in XT have higher abundance than those in ZF, whereas 5 ribosome genes in XT showed lower abundance than those in ZF. In addition, heat-tolerant sweet maize may keep at lower growth level than heat-sensitive one through dowregulating expression of genes related to zeatin and brassinosteroid biosynthesis to better regulate heat stress responses. Comparative transcriptomic profiling reveals transcriptional alterations in heat-resistant and heat-sensitive sweet maize varieties under heat stress, which provides a new insight into underlying molecular mechanism of maize in response to heat stress.

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Comparative transcriptome analysis reveals the transcriptional alterations in heat-resistant and heat-sensitive sweet maize (Zea mays L.) varieties under heat stress.

A major problem of climate change is the increasing duration and frequency of heavy rainfall events. This leads to soil flooding that negatively affects plant growth, eventually leading to death of plants if the flooding persists for several days. Most crop plants are very sensitive to flooding, and dramatic yield losses occur due to flooding each year. This review summarizes recent progress and approaches to enhance crop resistance to flooding. Most experiments have been done on maize, barley, and soybean. Work on other crops such as wheat and rape has only started. The most promising traits that might enhance crop flooding tolerance are anatomical adaptations such as aerenchyma formation, the formation of a barrier against radial oxygen loss, and the growth of adventitious roots. Metabolic adaptations might be able to improve waterlogging tolerance as well, but more studies are needed in this direction. Reasonable approaches for future studies are quantitative trait locus (QTL) analyses or genome-wide association (GWA) studies in combination with specific tolerance traits that can be easily assessed. The usage of flooding-tolerant relatives or ancestral cultivars of the crop of interest in these experiments might enhance the chances of finding useful tolerance traits to be used in breeding.

Improving Flooding Tolerance of Crop Plants

Waterlogging stress caused a decrease of leaf chlorophyll content and premature leaf senescence, which are associated with dramatic changes in gene expression profiles in the aerial leaves of root-waterlogged rape seedlings.
 Soil waterlogging is a serious constraint to crop production. We investigated the physiological responses of rape (Brassica napus L.) seedlings to waterlogging stress and analyzed global gene transcription responses in the aerial leaves of waterlogged rape seedlings. Seedlings of ‘Tammi’ and ‘Youngsan’ cultivars were subjected to waterlogging for 3 and 6 days and recovery for 5 days. Waterlogging stress caused a significant decrease in leaf chlorophyll content and premature senescence of the leaves. Maximal quantum efficiency of PSII (F
 v/F
 m) decreased in the waterlogged seedlings compared with the control plants. To evaluate whether the observed physiological changes in the leaves are associated with the differential regulation of gene expression in response to waterlogging stress, we analyzed the global transcriptional profile of leaves of ‘Tammi’ seedlings that were exposed to waterlogging for a short period (36 and 72 h). SolexaQA RNA-seq analysis revealed that a total of 4,484 contigs (8.5 %) of all contigs assayed (52,747) showed a twofold change in expression after 36 h of the start of waterlogging and 9,659 contigs (18.3 %) showed a twofold change after 72 h. Major genes involved in leaf photosynthesis, including light reactions and carbon-fixing reactions, were downregulated, while a number of genes involved in the scavenging of reactive oxygen species, degradation (proteins, starch, and lipids), premature senescence, and abiotic stress tolerance were upregulated. Transcriptome analysis data suggested that the aerial leaves of waterlogged rape seedlings respond to hypoxia by regulating the expression of diverse genes in the leaves.

Ji-Hye Hwang

Papers

Global gene expression responses to waterlogging in leaves of rape seedlings