Reactive oxygen species (ROS) are versatile compounds which can have toxic or signalling effects in a wide range living organisms, including seeds. They have been reported to play a pivotal role in the regulation of seed germination and dormancy but their mechanisms of action are still far from being fully understood. In this review, we sum-up the major findings that have been carried out this last decade in this field of research and which altogether shed a new light on the signalling roles of ROS in seed physiology. ROS participate in dormancy release during seed dry storage through the direct oxidation of a subset of biomolecules. During seed imbibition, the controlled generation of ROS is involved in the perception and transduction of environmental conditions that control germination. When these conditions are permissive for germination, ROS levels are maintained at a level which triggers cellular events associated with germination, such as hormone signalling. Here we propose that the spatiotemporal regulation of ROS production acts in concert with hormone signalling to regulate the cellular events involved in cell expansion associated with germination.

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The signalling role of ROS in the regulation of seed germination and dormancy.

Rapeseed yield parameters have been altered due to the increased exposure to abiotic stresses. The changing climatic conditions have reduced biomass production and seed yield, which ultimately affect the success of rapeseed crops. The severity and ability of abiotic stresses to act in tandem are a growing concern for agriculture producers across the globe. Rapeseed is one of the most vital cultivars in the world, serving as a major player in edible oil production. Abiotic stress conditions impose various effects on plant metabolism associated with growth stages, soil water storage capacity and plant physiological aspects. Modern agriculture aims to increased plant productivity to ensure global food security for the future. This can be achieved using quality seed, appropriate agricultural practices, measuring pest reduction techniques and understanding crop production challenges. Important tools, including conventional breeding and biotechnological approaches, can be employed to develop abiotic stress tolerance rapeseed. This review highlights the physiological and biochemical responses of rapeseed to abiotic stresses. The review also addresses conventional and modern stress mitigation strategies for the development of climate-smart rapeseed.

Eco-physiological and Biochemical Responses of Rapeseed ( Brassica napus L.) to Abiotic Stresses: Consequences and Mitigation Strategies

Photoacoustic imaging with fiber optic technology: A review

In the coming decades, maintaining a steady food supply for the increasing world population will require high-yielding crop plants which can be productive under increasingly variable conditions. Maintaining high yields will require the successful and uniform establishment of plants in the field under altered environmental conditions. Seed vigor, a complex agronomic trait that includes seed longevity, germination speed, seedling growth, and early stress tolerance, determines the duration and success of this establishment period. Elevated temperature during early seed development can decrease seed size, number, and fertility, delay germination and reduce seed vigor in crops such as cereals, legumes, and vegetable crops. Heat stress in mature seeds can reduce seed vigor in crops such as lettuce, oat, and chickpea. Warming trends and increasing temperature variability can increase seed dormancy and reduce germination rates, especially in crops that require lower temperatures for germination and seedling establishment. To improve seed germination speed and success, much research has focused on selecting quality seeds for replanting, priming seeds before sowing, and breeding varieties with improved seed performance. Recent strides in understanding the genetic basis of variation in seed vigor have used genomics and transcriptomics to identify candidate genes for improving germination, and several studies have explored the potential impact of climate change on the percentage and timing of germination. In this review, we discuss these recent advances in the genetic underpinnings of seed performance as well as how climate change is expected to affect vigor in current varieties of staple, vegetable, and other crops. 

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Seed germination and vigor: ensuring crop sustainability in a changing climate

Rapeseed (Brassica napus L.) is an important oilseed crop in the world. Its productivity is significantly influenced by numerous abiotic stresses, including cold stress (CS). Consequently, enhancement in CS tolerance is becoming an important area for agricultural investigation and crop improvement. Therefore, the current study aimed to identify the stress-responsive genes, metabolites, and metabolic pathways based on a combined transcriptome and metabolome analysis to understand the CS responses and tolerance mechanisms in the cold-tolerant (C18) and cold-sensitive (C6) rapeseed varieties. Based on the metabolome analysis, 31 differentially accumulated metabolites (DAMs) were identified between different comparisons of both varieties at the same time points. From the transcriptome analysis, 2,845, 3,358, and 2,819 differentially expressed genes (DEGs) were detected from the comparison of C6-0 vs. C18-0, C6-1 vs. C18-1, and C6-7 vs. C18-7. By combining the transcriptome and metabolome data sets, we found that numerous DAMs were strongly correlated with several differentially expressed genes (DEGs). A functional enrichment analysis of the DAMs and the correlated DEGs specified that most DEGs and DAMs were mainly enriched in diverse carbohydrates and amino acid metabolisms. Among them, starch and sucrose metabolism and phenylalanine metabolism were significantly enriched and played a vital role in the CS adaption of rapeseed. Six candidate genes were selected from the two pathways for controlling the adaption to low temperature. In a further validation, the T-DNA insertion mutants of their Arabidopsis homologous, including 4cl3, cel5, fruct4, ugp1, axs1, and bam2/9, were characterized and six lines differed significantly in levels of freezing tolerance. The outcome of the current study provided new prospects for the understanding of the molecular basis of CS responses and tolerance mechanisms in rapeseed and present a set of candidate genes for use in improving CS adaptability in the same plant.

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Integrated Analysis of Metabolome and Transcriptome Reveals Insights for Cold Tolerance in Rapeseed (Brassica napus L.)

Slow germination speed caused by low temperature stress intensifies the risk posed by adverse environmental factors, contributing to low germination rate and reduced production of rapeseed. The purpose of this study was to understand the transcriptional regulation mechanism for rapid germination of rapeseed. The results showed that seed components and size do not determine the seed germination speed. Different temporal transcriptomic profiles were generated under normal and low temperature conditions in genotypes with fast and slow germination speeds. Using weight gene co-expression network analysis, 37 823 genes were clustered into 15 modules with different expression patterns. There were 10 233 and 9111 differentially expressed genes found to follow persistent tendency of up- and down-regulation, respectively, which provided the conditions necessary for germination. Hub genes in the continuous up-regulation module were associated with phytohormone regulation, signal transduction, the pentose phosphate pathway, and lipolytic metabolism. Hub genes in the continuous down-regulation module were involved in ubiquitin-mediated proteolysis. Through pairwise comparisons, 1551 specific upregulated DEGs were identified for the fast germination speed genotype under low temperature stress. These DEGs were mainly enriched in RNA synthesis and degradation metabolisms, signal transduction, and defense systems. Transcription factors, including WRKY, bZIP, EFR, MYB, B3, DREB, NAC, and ERF, are associated with low temperature stress in the fast germination genotype. The aquaporin NIP5 and late embryogenesis abundant (LEA) protein genes contributed to the water uptake and transport under low temperature stress during seed germination. The ethylene/H2O2-mediated signal pathway plays an important role in cell wall loosening and embryo extension during germination. The ROS-scavenging system, including catalase, aldehyde dehydrogenase, and glutathione S-transferase, was also upregulated to alleviate ROS toxicity in the fast germinating genotype under low temperature stress. These findings should be useful for molecular assisted screening and breeding of fast germination speed genotypes for rapeseed.

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Associating transcriptional regulation for rapid germination of rapeseed (Brassica napus L.) under low temperature stress through weighted gene co-expression network analysis.

High Yield Achieved by Early-Maturing Rapeseed with High Light Energy and Temperature Production Efficiencies under Ideal Planting Density

Evaluation of the Low‐Temperature Tolerance of Rapeseed Genotypes at the Germination and Seedling Emergence Stages

Low temperature inhibits rapid germination and successful seedling establishment of rapeseed (Brassica napus L.), leading to significant productivity losses. Little is known about the genetic diversity for seed vigor under low-temperature conditions in rapeseed, which motivated our investigation of 13 seed germination- and emergence-related traits under normal and low-temperature conditions for 442 diverse rapeseed accessions. The stress tolerance index was calculated for each trait based on performance under non-stress and low-temperature stress conditions. Principal component analysis of the low-temperature stress tolerance indices identified five principal components that captured 100% of the seedling response to low temperature. A genome-wide association study using ~8 million SNP (single-nucleotide polymorphism) markers identified from genome resequencing was undertaken to uncover the genetic basis of seed vigor related traits in rapeseed. We detected 22 quantitative trait loci (QTLs) significantly associated with stress tolerance indices regarding seed vigor under low-temperature stress. Scrutiny of the genes in these QTL regions identified 62 candidate genes related to specific stress tolerance indices of seed vigor, and the majority were involved in DNA repair, RNA translation, mitochondrial activation and energy generation, ubiquitination and degradation of protein reserve, antioxidant system, and plant hormone and signal transduction. The high effect variation and haplotype-based effect of these candidate genes were evaluated, and high priority could be given to the candidate genes BnaA03g40290D, BnaA06g07530D, BnaA09g06240D, BnaA09g06250D, and BnaC02g10720D in further study. These findings should be useful for marker-assisted breeding and genomic selection of rapeseed to increase seed vigor under low-temperature stress.

Genome-Wide Association Mapping Unravels the Genetic Control of Seed Vigor under Low-Temperature Conditions in Rapeseed (Brassica napus L.).

Screening and breeding elite varieties with rapid germination and uniform seedling emergence under low temperature is an effective strategy to deal with the cold climate occurring under late sowing conditions in the Yangtze River basin. This study focused on the performance of seven functional traits, including percentage of germination, percentage of emergence, mean germination time, mean emergence time, total seedling length, total dry weight, and seedling vigor index of 436 natural rapeseed populations under normal-temperature (25/20 °C) and low-temperature (15/10 °C) conditions. Furthermore, ten genotypes were screened to verify their low-temperature tolerance based on cultivar traits in a pot experiment. The results show that the germination- and emergence-related functional traits of rapeseed genotypes exhibit rich genotypic diversity in response to low-temperature stress; the variation among these traits ranged from 1–25% under normal-temperature and 10–49% under low-temperature conditions. Variation in seed characteristics also affected the capacity for low-temperature tolerance in the process of seed germination and seedling emergence, and could explain 22% of the total variance for low-temperature stress tolerance indices. There existed high correlations between the stress tolerance index of total dry weight (STI_TDW) and thousand-seed weight, and between the stress tolerance index of emergence percentage (STI_PE) and oil content. The contents of erucic acid, glucosinolate, and eicosenoic acid were positively correlated with the stress tolerance index of mean germination time (STI_MGT) and mean emergence time (STI_MET). The D-CRITIC (distance-based intercriteria correlation) weight method was selected in this experiment to calculate each variety’s comprehensive low-temperature stress tolerance index by integrating the standard deviation and distance correlation coefficient of each index. The genotypes with large comprehensive low-temperature stress tolerance index also had higher low-temperature stress tolerance index of biomass and yield in the pot experiment, indicating that the comprehensive low-temperature stress tolerance index has high reliability and applicability. This study could provide a theoretical basis for the utilization of low-temperature-tolerant germplasm resources, as well as a reference for the cold resistance and yield stability under late- and direct-sowing conditions of rapeseed in the Yangtze River basin and other similar environments around the world.

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Seed Characteristics Affect Low-Temperature Stress Tolerance Performance of Rapeseed (Brassica napus L.) during Seed Germination and Seedling Emergence Stages

Chunni Zhang

Papers

Associating transcriptional regulation for rapid germination of rapeseed (Brassica napus L.) under low temperature stress through weighted gene co-expression network analysis.

High Yield Achieved by Early-Maturing Rapeseed with High Light Energy and Temperature Production Efficiencies under Ideal Planting Density

Evaluation of the Low‐Temperature Tolerance of Rapeseed Genotypes at the Germination and Seedling Emergence Stages

Genome-Wide Association Mapping Unravels the Genetic Control of Seed Vigor under Low-Temperature Conditions in Rapeseed (Brassica napus L.).

Seed Characteristics Affect Low-Temperature Stress Tolerance Performance of Rapeseed (Brassica napus L.) during Seed Germination and Seedling Emergence Stages