Seed Storability in Rice: Physiological Foundations, Molecular Mechanisms, and Applications in Breeding

doi:10.1016/j.rsci.2024.02.011

Abstract

Abstract:

Long-term storage of crop seeds is critical for the conservation of germplasm resources, ensuring food supply, and supporting sustainable production. Rice, as a major food staple, has a substantial stock for consumption and production worldwide. However, its food value and seed viability tend to decline during storage. Understanding the physiological responses and molecular mechanisms of aging tolerance forms the basis for enhancing seed storability in rice. This review outlines the latest progress in influential factors, evaluation methods, and identification indices of seed storability. It also discusses the physiological consequences, molecular mechanisms, and strategies for breeding aging-tolerant rice in detail. Finally, it highlights challenges in seed storability research that require future attention. This review offers a theoretical foundation and research direction for uncovering the mechanisms behind seed storability and breeding aging-tolerant rice.

Key words: rice, seed storability, physiological response, molecular mechanism, aging-tolerant breeding

Zhou Tianshun, Yu Dong, Wu Liubing, Xu Yusheng, Duan Meijuan, Yuan Dingyang. Seed Storability in Rice: Physiological Foundations, Molecular Mechanisms, and Applications in Breeding[J]. Rice Science, 2024, 31(4): 401-416.

Figures/Tables 4

Fig. 1. Factors influencing rice seed storability.

Fig. 2. Main QTLs associated with seed storability in rice. Chr., Chromosome. QTLs with LOD > 2.5 and phenotypic variation explained > 15% are listed. These QTLs have been identified using different phenotypes such as germination rate (in red), seedling length (in green), fatty acid content (in blue), and P50 (in brown). Different names for a single locus indicate that the locus has been identified in at least two studies, suggesting that the QTL may have large and stable effects.

Fig. 3. Molecular mechanism mediated by aging-tolerant genes in rice. ROS, Reactive oxygen species; TAG, Triacylglycerol; PUFAs, Polyunsaturated fatty acids; LPOs, Lipid peroxidation; GA, Gibberellin; ABA, Abscisic acid; IAA, Auxin.

Fig. 4. Schematic diagram of improvement of aging-tolerant rice varieties. GWAS, Genome-wide association study.

References 100

[1]	Abe K, Araki E, Suzuki Y, Toki S, Saika H. 2018. Production of high oleic/low linoleic rice by genome editing. Plant Physiol Biochem, 131: 58-62.
[2]	Amanda M, Marcela T N, Vanessa M, Paulo C C, Rosana S M, Samuel M, Andressa F L, Vladison F P, Cristielle K M. 2022. Rice drying, storage and processing: Effects of post-harvest operations on grain quality. Rice Sci, 29(1): 16-30.
[3]	An J Y, Liu Y H, Han J J, He C, Chen M, Zhu X B, Hu W M, Song W J, Hu J, Guan Y J. 2022. Transcriptional multiomics reveals the mechanism of seed deterioration in Nicotiana tabacum L. and Oryza sativa L. J Adv Res, 42: 163-176.
[4]	Bai S Y, He N Q, Zhou L, Shen B B, Wu W, Liu X, Jiang L, Wan J M. 2015. Knock-down of OsLOX by RNA interference leads to improved seed viability in rice. J Plant Biol, 58(5): 293-302.
[5]	Biesalski H K. 2002. Free radical theory of aging. Curr Opin Clin Nutr Metab Care, 5(1): 5-10.
[6]	Bollinedi H, Singh A K, Singh N, Gopala K S, Bhowmick P K, Vinod K K, Nagarajan M, Ellur R K. 2021. Genetic and genomic approaches to address rapid rancidity of rice bran. Crit Rev Food Sci Nutr, 61(1): 75-84.
[7]	Bollinedi H, Singh N, Gopala Krishnan S, Vinod K K, Bhowmick P K, Nagarajan M, Ellur R K, Singh A K. 2022. A novel LOX3-null allele (lox3-b) originated in the aromatic Basmati rice cultivars imparts storage stability to rice bran. Food Chem, 369: 130887.
[8]	Buitink J, Leprince O. 2008. Intracellular glasses and seed survival in the dry state. C R Biol, 331(10): 788-795.
[9]	Chen D F, Li Y L, Fang T, Shi X L, Chen X W. 2016. Specific roles of tocopherols and tocotrienols in seed longevity and germination tolerance to abiotic stress in transgenic rice. Plant Sci, 244: 31-39.
[10]	Chen H H, Chu P, Zhou Y L, Li Y, Liu J, Ding Y, Tsang E W T, Jiang L W, Wu K Q, Huang S Z. 2012. Overexpression of AtOGG1, a DNA glycosylase/AP lyase, enhances seed longevity and abiotic stress tolerance in Arabidopsis. J Exp Bot, 63(11): 4107-4121.
[11]	Choudhary P, Pramitha L, Aggarwal P R, Rana S, Vetriventhan M, Muthamilarasan M. 2023. Biotechnological interventions for improving the seed longevity in cereal crops: Progress and prospects. Crit Rev Biotechnol, 43(2): 309-325.
[12]	Dong X Y, Fan S X, Liu J, Wang Q, Li M R, Jiang X, Liu Z Y, Yin Y C, Wang J Y. 2017. Identification of QTLs for seed storability in rice under natural aging conditions using two RILs with the same parent Shennong 265. J Integr Agric, 16(5): 1084-1092.
[13]	Fleming M B, Richards C M, Walters C. 2017. Decline in RNA integrity of dry-stored soybean seeds correlates with loss of germination potential. J Exp Bot, 68(9): 2219-2230.
[14]	Gao J D, Fu H, Zhou X Q, Chen Z J, Luo Y, Cui B Y, Chen G H, Liu J. 2016. Comparative proteomic analysis of seed embryo proteins associated with seed storability in rice (Oryza sativa L) during natural aging. Plant Physiol Biochem, 103: 31-44.
[15]	Gayen D, Ali N, Ganguly M, Paul S, Datta K, Datta S K. 2014. RNAi mediated silencing of lipoxygenase gene to maintain rice grain quality and viability during storage. Plant Cell Tiss Org, 118(2): 229-243.
[16]	Gayen D, Ali N, Sarkar S N, Datta S K, Datta K. 2015. Down- regulation of lipoxygenase gene reduces degradation of carotenoids of golden rice during storage. Planta, 242(1): 353-363.
[17]	Groot S P C, Surki A A, de Vos R C H, Kodde J. 2012. Seed storage at elevated partial pressure of oxygen, a fast method for analysing seed ageing under dry conditions. Ann Bot, 110(6): 1149-1159.
[18]	Hay F R, de Guzman F, Sackville Hamilton N R. 2015. Viability monitoring intervals for genebank samples of Oryza sativa. Seed Sci Technol, 43(2): 218-237.
[19]	Hang N T, Lin Q Y, Liu L L, Liu X, Liu S J, Wang W Y, Li L F, He N Q, Liu Z, Jiang L, Wan J M. 2015. Mapping QTLs related to rice seed storability under natural and artificial aging storage conditions. Euphytica, 203(3): 673-681.
[20]	Hazra A, Varshney V, Verma P, Kamble N U, Ghosh S, Achary R K, Gautam S, Majee M. 2022. Methionine sulfoxide reductase B5 plays a key role in preserving seed vigor and longevity in rice (Oryza sativa). New Phytol, 236(3): 1042-1060.
[21]	He Y Q, Cheng J P, He Y, Yang B, Cheng Y H, Yang C, Zhang H S, Wang Z F. 2019. Influence of isopropylmalate synthase OsIPMS1 on seed vigour associated with amino acid and energy metabolism in rice. Plant Biotechnol J, 17(2): 322-337.
[22]	Huang J X, Cai M H, Long Q Z, Liu L L, Lin Q Y, Jiang L, Chen S H, Wan J M. 2014. OsLOX2, a rice type I lipoxygenase, confers opposite effects on seed germination and longevity. Transgenic Res, 23(4): 643-655.
[23]	Ji J, Lin S S, Xin X, Li Y, He J J, Xu X Y, Zhao Y X, Su G F, Lu X X, Yin G K. 2023. Effects of OsAOX1a deficiency on mitochondrial metabolism at critical node of seed viability in rice. Plants, 12(12): 2284.
[24]	Jiang W Z, Lee J, Jin Y M, Qiao Y L, Piao R H, Jang S M, Woo M O, Kwon S W, Liu X H, Pan H Y, Du X L, Koh H J. 2011. Identification of QTLs for seed germination capability after various storage periods using two RIL populations in rice. Mol Cells, 31(4): 385-392.
[25]	Jin J, Long W X, Wang L T, Liu X D, Pan G J, Xiang W, Li N W, Li S Q. 2018. QTL mapping of seed vigor of backcross inbred lines derived from Oryza longistaminata under artificial aging. Front Plant Sci, 9: 1909.
[26]	Joosen R V L, Kodde J, Willems L A J, Ligterink W, van der Plas L H W, Hilhorst H W M. 2010. GERMINATOR: A software package for high-throughput scoring and curve fitting of Arabidopsis seed germination. Plant J, 62(1): 148-159.
[27]	Kamble N U, Majee M. 2022. ABI transcription factors and PROTEIN L-ISOASPARTYL METHYLTRANSFERASE module mediate seed desiccation tolerance and longevity in Oryza sativa. Development, 149(11): dev200600.
[28]	Kaur H, Petla B P, Kamble N U, Singh A, Rao V, Salvi P, Ghosh S, Majee M. 2015. Differentially expressed seed aging responsive heat shock protein OsHSP18.2 implicates in seed vigor, longevity and improves germination and seedling establishment under abiotic stress. Front Plant Sci, 6: 713.
[29]	Khan M S S, Basnet R, Islam S A, Shu Q Y. 2019. Mutational analysis of OsPLDα1 reveals its involvement in phytic acid biosynthesis in rice grains. J Agric Food Chem, 67(41): 11436-11443.
[30]	Khan M S S, Basnet R, Ahmed S, Bao J S, Shu Q Y. 2020. Mutations of OsPLDa1 increase lysophospholipid content and enhance cooking and eating quality in rice. Plants, 9(3): 390.
[31]	Kranner I, Chen H Y, Pritchard H W, Pearce S R, Birtić S. 2011. Inter-nucleosomal DNA fragmentation and loss of RNA integrity during seed ageing. Plant Growth Regul, 63(1): 63-72.
[32]	Kurek K, Plitta-Michalak B, Ratajczak E. 2019. Reactive oxygen species as potential drivers of the seed aging process. Plants, 8(6): 174.
[33]	Lee J S, Kwak J, Yoon M R, Lee J S, Hay F R. 2017. Contrasting tocol ratios associated with seed longevity in rice variety groups. Seed Sci Res, 27(4): 273-280.
[34]	Lee J S, Velasco-Punzalan M, Pacleb M, Valdez R, Kretzschmar T, McNally K L, Ismail A M, Cruz P C S, Sackville Hamilton N R, Hay F R. 2019. Variation in seed longevity among diverse Indica rice varieties. Ann Bot, 124(3): 447-460.
[35]	Lee J S, Kwak J, Hay F R. 2020. Genetic markers associated with seed longevity and vitamin E in diverse Aus rice varieties. Seed Sci Res, 30(2): 133-141.
[36]	Li C S, Shao G S, Wang L, Wang Z F, Mao Y J, Wang X Q, Zhang X H, Liu S T, Zhang H S. 2017. QTL identification and fine mapping for seed storability in rice (Oryza sativa L.). Euphytica, 213(6): 127.
[37]	Li L F, Lin Q Y, Liu S J, Liu X, Wang W Y, Hang N T, Liu F, Zhao Z G, Jiang L, Wan J M. 2012. Identification of quantitative trait loci for seed storability in rice (Oryza sativa L.). Plant Breed, 131(6): 739-743.
[38]	Li T, Zhang Y M, Wang D, Liu Y, Dirk L M A, Goodman J, Downie A B, Wang J M, Wang G Y, Zhao T Y. 2017. Regulation of seed vigor by manipulation of raffinose family oligosaccharides in maize and Arabidopsis thaliana. Mol Plant, 10(12): 1540-1555.
[39]	Li Y, Wang Y, Xue H, Pritchard H W, Wang X F. 2017. Changes in the mitochondrial protein profile due to ROS eruption during ageing of elm (Ulmus pumila L.) seeds. Plant Physiol Biochem, 114: 72-87.
[40]	Lin Q Y, Jiang Y M, Sun A L, Cao P H, Li L F, Liu X, Tian Y L, He J, Liu S J, Chen L M, Jiang L. 2015a. Fine mapping of qSS-9, a major and stable quantitative trait locus, for seed storability in rice (Oryza sativa L.). Plant Breed, 134(3): 293-299.
[41]	Lin Q Y, Wang W Y, Ren Y K, Jiang Y M, Sun A L, Qian Y, Zhang Y F, He N Q, Hang N T, Liu Z, Li L F, Liu L L, Jiang L, Wan J M. 2015b. Genetic dissection of seed storability using two different populations with a same parent rice cultivar N22. Breed Sci, 65(5): 411-419.
[42]	Liu S J, Liu W H, Lai J Y, Liu Q J, Zhang W H, Chen Z J, Gao J D, Song S Q, Liu J, Xiao Y H. 2022. OsGLYI3, a glyoxalase gene expressed in rice seed, contributes to seed longevity and salt stress tolerance. Plant Physiol Biochem, 183: 85-95.
[43]	Long Q Z, Zhang W W, Wang P, Shen W B, Zhou T, Liu N N, Wang R, Jiang L, Huang J X, Wang Y H, Liu Y Q, Wan J M. 2013. Molecular genetic characterization of rice seed lipoxygenase 3 and assessment of its effects on seed longevity. J Plant Biol, 56(4): 232-242.
[44]	Ma L, Zhu F G, Li Z W, Zhang J F, Li X, Dong J L, Wang T. 2015. TALEN-based mutagenesis of lipoxygenase LOX3 enhances the storage tolerance of rice (Oryza sativa) seeds. PLoS One, 10(12): e0143877.
[45]	Ma L Y, Deng D Y, Su Y, Xiao L T. 2024. X-ray-μCT: Nondestructively identifying hidden microphenotypes inside living crop seeds. Trends Plant Sci, 29(1): 99-100.
[46]	Miao C B, Wang Z, Zhang L, Yao J J, Hua K, Liu X, Shi H Z, Zhu J K. 2019. The grain yield modulator miR156 regulates seed dormancy through the gibberellin pathway in rice. Nat Commun, 10(1): 3822.
[47]	Miura K, Lin S Y, Yano M, Nagamine T. 2002. Mapping quantitative trait loci controlling seed longevity in rice (Oryza sativa L.). Theor Appl Genet, 104(6/7): 981-986.
[48]	Müller A, Nunes M T, Maldaner V, Coradi P C, Santos de Moraes R, Martens S, Leal A F, Pereira V F, Marin C K. 2022. Rice drying, storage and processing: Effects of post-harvest operations on grain quality. Rice Sci, 29(1): 16-30.
[49]	National Grain and oil Standardization Technical Committee. 2006. Guidelines for evaluation of grain storage character: Paddy: GB/T 20569-2006. (2006-12-01) [2024-07-22]. China: Beijing.
[50]	Nisarga K N, Vemanna R S, Kodekallu Chandrashekar B, Rao H, Vennapusa A R, Narasimaha A, Makarla U, Basavaiah M R. 2017. Aldo-ketoreductase 1 (AKR1) improves seed longevity in tobacco and rice by detoxifying reactive cytotoxic compounds generated during ageing. Rice, 10(1): 11.
[51]	Oenel A, Fekete A, Krischke M, Faul S C, Gresser G, Havaux M, Mueller M J, Berger S. 2017. Enzymatic and non-enzymatic mechanisms contribute to lipid oxidation during seed aging. Plant Cell Physiol, 58(5): 925-933.
[52]	Parkhey S, Naithani S C, Keshavkant S. 2012. ROS production and lipid catabolism in desiccating Shorea robusta seeds during aging. Plant Physiol Biochem, 57: 261-267.
[53]	Peng B, He L L, Tan J, Zheng L T, Zhang J T, Qiao Q W, Wang Y, Gao Y, Tian X Y, Liu Z Y, Song X H, Sun Y Y, Pang R H, Li J T, Yuan H Y. 2019. Effects of rice aging on its main nutrients and quality characters. J Agric Sci, 11(17): 44.
[54]	Petla B P, Kamble N U, Kumar M, Verma P, Ghosh S, Singh A, Rao V, Salvi P, Kaur H, Saxena S C, Majee M. 2016. Rice PROTEIN l-ISOASPARTYL METHYLTRANSFERASE isoforms differentially accumulate during seed maturation to restrict deleterious isoAsp and reactive oxygen species accumulation and are implicated in seed vigor and longevity. New Phytol, 211(2): 627-645.
[55]	Powell A A, Yule L J, Jing H C, Groot S P, Bino R J, Pritchard H W. 2000. The influence of aerated hydration seed treatment on seed longevity as assessed by the viability equations. J Exp Bot, 51: 2031-2043.
[56]	Prasad C T M, Kodde J, Angenent G C, Hay F R, McNally K L, Groot S P C. 2023. Identification of the rice Rc gene as a main regulator of seed survival under dry storage conditions. Plant Cell Environ, 46(6): 1962-1980.
[57]	Rajjou L, Debeaujon I. 2008. Seed longevity: Survival and maintenance of high germination ability of dry seeds. C R Biol, 331(10): 796-805.
[58]	Raquid R, Kohli A, Reinke R, Dionisio-Sese M, Kwak J, Chebotarov D, Mo Y, Lee J S. 2021. Genetic factors enhancing seed longevity in tropical japonica rice. Curr Plant Biol, 26: 100196.
[59]	Rehmani M S, Xian B S, Wei S W, He J, Feng Z X, Huang H, Shu K. 2023. Seedling establishment: The neglected trait in the seed longevity field. Plant Physiol Biochem, 200: 107765.
[60]	Ren R J, Wang P, Wang L N, Su J P, Sun L J, Sun Y, Chen D F, Chen X W. 2020. Os4BGlu14, a monolignol β-glucosidase, negatively affects seed longevity by influencing primary metabolism in rice. Plant Mol Biol, 104(4/5): 513-527.
[61]	Sano N, Rajjou L, North H M, Debeaujon I, Marion-Poll A, Seo M. 2016. Staying alive: Molecular aspects of seed longevity. Plant Cell Physiol, 57(4): 660-674.
[62]	Sano N, Takebayashi Y, To A, Mhiri C, Rajjou L C, Nakagami H, Kanekatsu M. 2019. Shotgun proteomic analysis highlights the roles of long-lived mRNAs and de novo transcribed mRNAs in rice seeds upon imbibition. Plant Cell Physiol, 60(11): 2584-2596.
[63]	Sasaki K, Fukuta Y, Sato T. 2005. Mapping of quantitative trait loci controlling seed longevity of rice (Oryza sativa L.) after various periods of seed storage. Plant Breed, 124(4): 361-366.
[64]	Sendin K, Williams P J, Manley M. 2018. Near infrared hyperspectral imaging in quality and safety evaluation of cereals. Crit Rev Food Sci Nutr, 58(4): 575-590.
[65]	Shin J H, Kim S R, An G. 2009. Rice aldehyde dehydrogenase 7 is needed for seed maturation and viability. Plant Physiol, 149(2): 905-915.
[66]	Shirasawa K, Takeuchi Y, Ebitani T, Suzuki Y. 2008. Identification of gene for rice (Oryza sativa) seed lipoxygenase-3 involved in the generation of stale flavor and development of SNP markers for lipoxygenase-3 deficiency. Breed Sci, 58(2): 169-176.
[67]	Suzuki Y, Higo K, Hagiwara K, Ohtsubo K, Kobayashi A, Nozu Y. 1992. Production and use of monoclonal antibodies against rice embryo lipoxygenase-3. Biosci Biotechnol Biochem, 56(4): 678-679.
[68]	Suzuki Y, Nagamine T, Okuno K. 1996. Genetic analysis of a null- allele for lipoxygenase-3 in rice seeds. Euphytica, 91(1): 99-101.
[69]	Suzuki Y, Takeuchi Y, Shirasawa K. 2011. ) Identification of a seed phospholipase D null allele in rice (Oryza sativa L. and development of SNP markers for phospholipase D deficiency. Crop Sci, 51(5): 2113-2118.
[70]	Suzuki Y, Miura K, Shigemune A, Sasahara H, Ohta H, Uehara Y, Ishikawa T, Hamada S, Shirasawa K. 2015. Marker-assisted breeding of a LOX-3-null rice line with improved storability and resistance to preharvest sprouting. Theor Appl Genet, 128(7): 1421-1430.
[71]	Tesnier K, Strookman-Donkers H M, van Pijlen J G, van der Geest A H M, Bino R J, Groot S P C. 2002. A controlled deterioration test for Arabidopsis thaliana reveals genetic variation in seed quality. Seed Sci Technol, 30(1): 149-165.
[72]	Tiwari G J, Liu Q, Shreshtha P, Li Z Y, Rahman S. 2016. RNAi-mediated down-regulation of the expression of OsFAD2-1: Effect on lipid accumulation and expression of lipid biosynthetic genes in the rice grain. BMC Plant Biol, 16(1): 189.
[73]	Verdier J, Lalanne D, Pelletier S, Torres-Jerez I, Righetti K, Bandyopadhyay K, Leprince O, Chatelain E, Vu B L, Gouzy J, Gamas P, Udvardi M K, Buitink J. 2013. A regulatory network- based approach dissects late maturation processes related to the acquisition of desiccation tolerance and longevity of Medicago truncatula seeds. Plant Physiol, 163(2): 757-774.
[74]	Wang B Q, Wang S Y, Tang Y Q, Jiang L L, He W, Lin Q L, Yu F, Wang L. 2022. Transcriptome-wide characterization of seed aging in rice: Identification of specific long-lived mRNAs for seed longevity. Front Plant Sci, 13: 857390.
[75]	Wang F X, Xu H B, Zhang L, Shi Y R, Song Y, Wang X Y, Cai Q H, He W, Xie H A, Zhang J F. 2023. The lipoxygenase OsLOX10 affects seed longevity and resistance to saline-alkaline stress during rice seedlings. Plant Mol Biol, 111(4/5): 415-428.
[76]	Wang W Q, Xu D Y, Sui Y P, Ding X H, Song X J. 2022. A multiomic study uncovers a bZIP23-PER1A-mediated detoxification pathway to enhance seed vigor in rice. Proc Natl Acad Sci USA, 119(9): e2026355119.
[77]	Wang Y, Branicky R, Noë A, Hekimi S. 2018. Superoxide dismutases: Dual roles in controlling ROS damage and regulating ROS signaling. J Cell Biol, 217(6): 1915-1928.
[78]	Waterworth W M, Bray C M, West C E. 2015. The importance of safeguarding genome integrity in germination and seed longevity. J Exp Bot, 66(12): 3549-3558.
[79]	Waterworth W M, Bray C M, West C E. 2019. Seeds and the art of genome maintenance. Front Plant Sci, 10: 706.
[80]	Wei Y D, Xu H B, Diao L R, Zhu Y S, Xie H G, Cai Q H, Wu F X, Wang Z H, Zhang J F, Xie H A. 2015. Protein repair L-isoaspartyl methyltransferase 1 (PIMT1) in rice improves seed longevity by preserving embryo vigor and viability. Plant Mol Biol, 89(4/5): 475-492.
[81]	Wiebach J, Nagel M, Börner A, Altmann T, Riewe D. 2020. Age- dependent loss of seed viability is associated with increased lipid oxidation and hydrolysis. Plant Cell Environ, 43(2): 303-314.
[82]	Wu F X, Luo X, Wang L Q, Wei Y D, Li J G, Xie H A, Zhang J F, Xie G S. 2021. Genome-wide association study reveals the QTLs for seed storability in world rice core collections. Plants, 10(4): 812.
[83]	Wu R, Ding Y Q, Li C Y, Wu B K, Huang Z J, Li Z N, Wang X M, Zhao G W. 2022. An R2R3-type transcription factor OsMYBAS1 regulates seed germination under artificial accelerated aging in transgenic rice (Oryza sativa L.). Agronomy, 12(8): 1955.
[84]	Wu X L, Ning F, Hu X L, Wang W. 2017. Genetic modification for improving seed vigor is transitioning from model plants to crop plants. Front Plant Sci, 8: 8.
[85]	Xu H B, Wei Y D, Zhu Y S, Lian L, Xie H G, Cai Q H, Chen Q S, Lin Z P, Wang Z H, Xie H A, Zhang J F. 2015. Antisense suppression of LOX3 gene expression in rice endosperm enhances seed longevity. Plant Biotechnol J, 13(4): 526-539.
[86]	Xu N, Chu Y L, Chen H L, Li X X, Wu Q, Jin L, Wang G X, Huang J L. 2018. Rice transcription factor OsMADS25 modulates root growth and confers salinity tolerance via the ABA-mediated regulatory pathway and ROS scavenging. PLoS Genet, 14(10): e1007662.
[87]	Xue Y, Zhang S Q, Yao Q H, Peng R H, Xiong A S, Li X, Zhu W M, Zhu Y Y, Zha D S. 2008. Identification of quantitative trait loci for seed storability in rice (Oryza sativa L.). Euphytica, 164(3): 739-744.
[88]	Yin G K, Whelan J, Wu S H, Zhou J, Chen B Y, Chen X L, Zhang J M, He J J, Xin X, Lu X X. 2016. Comprehensive mitochondrial metabolic shift during the critical node of seed ageing in rice. PLoS One, 11(4): e0148013.
[89]	Yin G K, Xin X, Fu S Z, An M N, Wu S H, Chen X L, Zhang J M, He J J, Whelan J, Lu X X. 2017. Proteomic and carbonylation profile analysis at the critical node of seed ageing in Oryza sativa. Sci Rep, 7: 40611.
[90]	Yuan Z Y, Fan K, Xia L F, Ding X L, Tian L, Sun W Q, He H Z, Yu S B. 2019. Genetic dissection of seed storability and validation of candidate gene associated with antioxidant capability in rice (Oryza sativa L.). Int J Mol Sci, 20(18): 4442.
[91]	Yuan Z Y, Fan K, Wang Y T, Tian L, Zhang C P, Sun W Q, He H Z, Yu S B. 2021. OsGRETCHENHAGEN3-2 modulates rice seed storability via accumulation of abscisic acid and protective substances. Plant Physiol, 186(1): 469-482.
[92]	Zaplin E S, Liu Q, Li Z Y, Butardo V M, Blanchard C L, Rahman S. 2013. Production of high oleic rice grains by suppressing the expression of the OsFAD2-1 gene. Funct Plant Biol, 40(10): 996-1004.
[93]	Zeng D L, Guo L B, Xu Y B, Yasukumi K, Zhu L H, Qian Q. 2006. QTL analysis of seed storability in rice. Plant Breed, 125(1): 57-60.
[94]	Zhang Y, Yu Z L, Lu Y X, Wang Y, She D H, Song M, Wu Y J. 2007. Effect of the absence of lipoxygenase isoenzymes on the storage characteristics of rice grains. J Stored Prod Res, 43(1): 87-91.
[95]	Zhao J, He Y Q, Huang S L, Wang Z F. 2021. Advances in the identification of quantitative trait loci and genes involved in seed vigor in rice. Front Plant Sci, 12: 659307.
[96]	Zhao M M, Hu B L, Fan Y W, Ding G M, Yang W L, Chen Y, Chen Y H, Xie J K, Zhang F T. 2021. Identification, analysis, and confirmation of seed storability-related loci in Dongxiang wild rice (Oryza rufipogon griff.). Genes, 12(11): 1831.
[97]	Zhou S Q, Huang K R, Zhou Y, Hu Y Q, Xiao Y C, Chen T, Yin M Q, Liu Y, Xu M L, Jiang X C. 2022. Degradome sequencing reveals an integrative miRNA-mediated gene interaction network regulating rice seed vigor. BMC Plant Biol, 22(1): 269.
[98]	Zhou W G, Chen F, Luo X F, Dai Y J, Yang Y Z, Zheng C, Yang W Y, Shu K. 2020. A matter of life and death: Molecular, physiological, and environmental regulation of seed longevity. Plant Cell Environ, 43(2): 293-302.
[99]	Zhou Y, Zhou S Q, Wang L P, Wu D, Cheng H L, Du X, Mao D D, Zhang C L, Jiang X C. 2020. miR164c and miR168a regulate seed vigor in rice. J Integr Plant Biol, 62(4): 470-486.
[100]	Zinsmeister J, Leprince O, Buitink J. 2020. Molecular and environmental factors regulating seed longevity. Biochem J, 477(2): 305-323.