Research Advances on Rice Seed Vigor

doi:10.16819/j.1001-7216.2018.7140

Abstract

Abstract:

It has always been one of common concerns to improve grain yield of rice, one of the most important cereal crops. Seed vigor has been an increasingly important agricultural trait as direct-seeded rice technology is widely applied in some Asian countries and regions. The concepts of seed development, germination and seed vigor were introduced. And the influencing factors of seed vigor, physiological and biochemical factors and genetic mechanism were analyzed. The research progress of QTL/gene mapping in rice seed vigor was introduced and the prospect was predicted, which is of great significance to crop breeding aiming to improve the seed vigor.

Key words: rice, seed vigor, physiological and biochemical mechanism, QTL mapping, gene mapping

摘要：

水稻作为重要的禾谷类作物之一,如何提高其产量一直备受人们关注。随着直播技术在亚洲一些国家和地区的广泛应用,水稻种子活力这一农艺性状显得尤为重要。本文对种子的发育和萌发过程和种子活力的概念加以说明,同时分析了水稻种子活力的影响因素、生理生化机制和遗传机理,介绍了水稻种子活力QTL/基因定位的研究进展并提出了展望,对指导以改良种子活力为目标的作物育种具有重要意义。

关键词: 水稻, 种子活力, 生理生化机制, QTL定位, 基因定位

CLC Number:

S511.01

Anpeng ZHANG, Qian QIAN, Zhenyu GAO. Research Advances on Rice Seed Vigor[J]. Chinese Journal OF Rice Science, 2018, 32(3): 296-303.

张安鹏, 钱前, 高振宇. 水稻种子活力的研究进展[J]. 中国水稻科学, 2018, 32(3): 296-303.

Figures/Tables 1

References 75

[1]	Catusse J, Job C, Job D.Transcriptome- and proteome-wide analyses of seed germination.CompRendBiol, 2008, 331(10):815-822.
[2]	Bewley J D.Seed germination and dormancy.Plant Cell, 1997, 9(7):1055-1066.
[3]	Da S E, Toorop P E, van LammerenAA, Hilhorst H W. ABA inhibits embryo cell expansion and early cell division events during coffee ( Coffeaarabica Coffeaarabica.AnnBot, 2008, 102(3):425-433.
[4]	Nakashima K, Yamaguchi-Shinozaki K.ABA signaling in stress-response and seed development.Plant CellRep, 2013, 32(7):959-970.
[5]	Zhu G H, Ye N H, Zhang J H.Glucose-induced delay of seed germination in rice is mediated by the suppression of ABA catabolism rather than an enhancement of ABA biosynthesis.Plant &Cell Physiol, 2009, 50(3):644-651.
[6]	Yang P F, Li X J, Wang X Q, Chen H, Chen F, Shen S H.Proteomic analysis of rice ( Oryza sativa ) seeds during germination.Proteomics, 2007, 7(18):3358-3368.
[7]	孙群, 王建华, 孙宝启. 种子活力的生理和遗传机理研究进展. 中国农业科学, 2007, 40(1):48-53.
	Sun Q, Wang J H, Sun B Q.Advances on seed vigor physiological and genetic mechanisms.SciAgricSin, 2007, 40(1):48-53. (in Chinese with English abstract)
[8]	高厚玉, 景立权, 陈龙, 居静, 王云霞, 朱建国, 杨连新, 王余龙. 自由空气中CO2浓度和温度增高对水稻种子活力的影响. 中国水稻科学, 2016, 30(4):371-379.
	Gao H Y, Jing L Q, Chen L, Ju J, Wang Y X, Zhu J G, Yang L X, Wang Y L.Effects of elevated atmospheric CO2 and temperature on seed vigor of rice under open-air field conditions.Chin J Rice Sci, 2016, 30(4):371-379. (in Chinese with English abstract)
[9]	方玉梅, 宋明. 种子活力研究进展. 种子科技, 2006, 24(2): 33-36.
	Fang Y M, Song M.Research progress of seed vigor.Seed Sci&Technol, 2006, 24(2):33-36. (in Chinese)
[10]	刘毓侠, 王铁固. 种子活力研究进展. 玉米科学, 2012, 20(4): 90-94.
	Liu Y X, Wang T G.Research progress of seed vigor.JMaize Sci, 2012, 20(4):90-94. (in Chinese with English abstract)
[11]	张红生, 胡晋. 种子学. 北京: 科学出版社, 2010.
	Zhang H S, Hu J. Seed Science.Beijing: Science Press, 2010. (in Chinese)
[12]	Hodgkin T, Hegarty T W.Genetically determined variation in seed germination and field emergence of Brassica oleracea. AnnApplBiol, 2010, 88(3):407-413.
[13]	王洋, 王盈盈, 洪德林. 太湖流域水稻种子活力和耐缺氧能力遗传变异研究. 南京农业大学学报, 2009, 32(3):1-7.
	Wang Y, Wang Y Y, Hong D L.Genetic variation of seed vigor and tolerance to anoxia among rice (Oryza sativa L.) varieties in Taihu Lake region. JNanjing AgricUniv, 2009, 32(3):1-7. (in Chinese with English abstract)
[14]	佟汉文, 刘易科, 朱展望, 张宇庆, 陈泠, 高春保. 基因型和环境对小麦种子活力的影响. 麦类作物学报, 2012, 32(6):1167-1170.
	Tong H W, Liu Y K, Zhu Z W, Zhang Y Q, Chen L, Gao C B.Effects of genotype and wheat seed vigor.JTritCrops, 2012, 32(6):1167-1170. (in Chinese with English abstract)
[15]	成广雷, 张海娇, 赵久然, 刘春阁, 王元东, 王晓光, 王荣焕, 陈传永, 徐田军. 临界胁迫贮藏条件下不同基因型玉米种子活力及生理变化. 中国农业科学, 2015, 48(1):33-42.
	Cheng G L, Zhang H J, Zhao J R, Liu C G, Wang Y D, Wang X G, Wang R H, Chen C Y, Xu T J.Vigor andphysiological changes of different genotypes of maize seed ( Zea mays L.) under critical stress storageconditions.SciAgric Sin, 2015, 48(1):33-42. (in Chinese with English abstract)
[16]	Ellis R H, Jackson M T.Seed production environment, time of harvest, and the potential longevity of seeds of three cultivars of rice (Oryza sativa L.). AnnBot, 1993, 72(6):583-590.
[17]	周新国. 杂交水稻制种喷施穗萌抑制剂的效果初探. 杂交水稻, 2003, 18(4):37-38.
	Zhou X G.Effects of Suimengyizhiji on inhibiting germination of seeds on panicles in hybrid rice seed production.Hybrid Rice, 2003, 18(4):37-38. (in Chinese with English abstract)
[18]	Fussell L K, Pearson C J.Effects of grain development and thermal history on grain maturation and seed vigour of Pennisetumamericanum. JExpBot, 1980, 31(2):635-643.
[19]	毛培胜, 韩建国, 浦心春, 宋锦峰, 倪小琴. 高羊茅种子成熟过程中的活力变化. 中国草地学报, 1997, 5(2):36-41.
	Mao P S, Han J G, Pu X C, Song J F, Ni X Q.Change of seed vigor during maturation in tall fescue.ChinJGrassl, 1997, 5(2):36-41. (in Chinese with English abstract)
[20]	王显国, 韩建国, 陈志红. 新麦草种子成熟过程中活力变化的研究. 草地学报, 2000, 8(4): 306-311.
	Wang X G, Han J G, Chen Z H.Research on vigor changes of new wheatgrass seeds during maturation.ActaAgrest Sin, 2000, 8(4):306-311.(in Chinese with English abstract)
[21]	张建成, 王辉. 不同成熟度花生种子发芽率及活力差异性研究. 种子, 2005, 24(1): 3-4.
	Zhang J C, Wang H.Studies on the seed germination and viability of different maturation peanut.Seed,2005, 24(1):3-4. (in Chinese with English abstract)
[22]	Adam N M, McDonnald M B, Henderlong P R. The influence of seed position, planting and harvesting dates on soybean seed quality.Seed SciTechnol, 1989, 17(1):143-152.
[23]	Guan Y J, Hu J, Wang Z F, Zhu S J, Wang J C, Knapp A.Time series regression analysis between changes in kernel size and seed vigor during developmental stage of sh2, sweet corn (Zea mays L.) seeds.SciHort, 2013, 154(2):25-30.
[24]	朱世杨, 郭媛, 洪德林. 水稻种子抗老化遗传分析. 遗传, 2008, 30(2):217-224.
	Zhu S Y, Guo Y, Hong D L.Genetic analysis on aging-resistant in rice seed.Hereditas(Beijing), 2008, 30(2):217-224. (in Chinese with English abstract)
[25]	李金华, 王丰, 廖亦龙, 刘武革. 水稻种子活力的生理生化及遗传研究. 分子植物育种, 2009, 7(4):772-777.
	Li J H, Wang F, Liao Y L, Liu W G.Advance on seed vigor physiological-biochemical and geneticmechanisms in rice.MolPlant Breed, 2009, 7(4):772-777. (in Chinese with English abstract)
[26]	Troyjack C, Pimentel J R, Ítala T D P, Ruddy A V E, Lanes B A J, Felipe K, Manoela A M, Gustavo H D, Vinícius J S, Ivan R C, Luis O B S, Tiago Z A, Tiago P. Nitrogen fertilization on maize sowing: plant growth and seed vigor.AmJPlant Sci, 2018, 9(1):83-97.
[27]	汪晓峰, 丛滋金. 种子活力的生物学基础及提高和保持种子活力的研究进展. 种子, 1997, 16(6): 36-39.
	Wang X F, Cong Z J.The biological basis of seed vigor and the research progress of improving andmaintaining seed vigor. Seed, 1997, 16(6):36-39. (in Chinese)
[28]	张玉兰, 汪晓峰, 景新明, 林坚. 水稻种子含水量及其对贮藏寿命的影响. 中国农业科学, 2005, 38(7):1480-1486.
	Zhang Y L, Wang X F, Jing X M, Lin J.The effect of moisture content on storage life of rice seeds.SciAgricSin, 2005, 38(7):1480-1486. (in Chinese with English abstract)
[29]	张兆英, 秦淑英, 王文全, 韩婧. 不同贮藏条件对3种药用植物种子活力的影响. 安徽农业科学, 2012, 40(9):5157-5159.
	Zhang Z Y, Qin S Y, Wang W Q, Han J.Effects of storage condition on seed vigor of three medicinalplants.JAnhui AgricSci, 2012, 40(9):5157-5159. (in Chinese with English abstract)
[30]	张凤, 刘美, 杨翠翠, 杨文思, 孙庆泉. 贮藏温度和种子含水量对大豆种子活力的影响. 山东农业科学, 2014(8):37-41.
	Zhang F, Liu M, Yang C C, Yang W S, Sun Q Q.Effects of storage temperature and seed moisture contenton soybean seed vigor.Shandong AgricSci, 2014(8):37-41. (in Chinese with Englishabstract)
[31]	杨永青, 汪晓峰. 种子活力与生物膜的研究现状. 植物学报, 2004, 21(6):641-648.
	Yang Y Q, Wang X F.Advances on relationship between biomembrane and seed vigor.ChinBullBot, 2004, 21(6):641-648. (in Chinese with English abstract)
[32]	Yamane K, Garcia R, Imayoshi K, Mabesa-Telosa R C, Banayo N P M C, Vergara G V, Yamauchi A, Cruz P S, Kato Y. Seed vigour contributes to yield improvement in dry direct-seeded rainfed lowland rice.AnnApplBiol, 2018, 172(1):100-110.
[33]	马书燕, 李吉跃, 彭祚登. 人工老化过程中柔枝松种子酶活性变化的研究. 种子, 2011, 30(5):9-14.
	Ma S Y, Li J Y, Peng Z D.Study on the activity changes of enzyme in the seeds of Pinusfiexilis Jamesduring artificial aging. Seed, 2011, 30(5):9-14. (in Chinese with English abstract)
[34]	Wang Y X, Xiong G S, Hu J, Jiang L, Yu H, Fang Y X, Zeng L J, Xu E B, Xu J, Ye W J, Meng X B, Liu R F, Chen H Q, Jing Y H, Wang Y H, Zhu X D, Li J Y, Qian Q.Copy number variation at the GL7 locus contributes to grain size diversity in rice.NatGenet, 2015, 47(8):944-948.
[35]	Catusse J, Strub J M, Job C, Dorsselaer A V, Job D.Proteome-wide characterization of sugarbeet seed vigor and its tissue specific expression.Proc Natl AcadSciUSA, 2008, 105(29):10262-10267.
[36]	Châtelain E, Satour P, Laugier E, Vu B L, Payet N, Rey P, Montrichard F.Evidence for participation of the methionine sulfoxide reductase repair system in plant seed longevity.Proc Natl AcadSciUSA, 2013, 110(9):3633-3638.
[37]	Qin M L, Luo F X, Liu L S, Zeng Z Y, Jiang X C.A Study on the relationship between the expression of miR164c and miR168b and seed vigor of rice.Acta Laser BiolSin, 2013, 22(2):166-173.
[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. Regulation of seed vigor by manipulation of raffinose family oligosaccharides (RFOs) in maize and Arabidopsis.MolPlant, 2017, 10(12):1540-1555.
[39]	Kaur H, Petla B P, Kamble N U, Singh A, Rao V, Salvi P, Ghosh S, Majee M,.Differentially expressed seed aging responsive heat shock protein OsHSP18.2 implicates in seed vigor, longevity and improves germination and seedling establishment under abiotic stress.FrontPlant Sci, 2015, 6:713-725.
[40]	Cheng X Y, Wu Y, Guo J P, Du B, Chen R Z, Zhu L L, He G C.A rice lectin receptor-like kinase that is involved in innate immune responses also contributes to seed germination.Plant J, 2013, 76(4):687.
[41]	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.Protein repairL-isoaspartyl methyltransferase 1(PIMT1) in rice improves seed longevity by preserving embryo vigor and viability.Plant MolBiol,2015,89(4-5):475-492.
[42]	Petla B P, Kamble N U, Kumar M, Verma P, Ghosh S, Singh V, Rao V, Salvi P, Kaur H, Saxena S C,Majee M.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, 2016, 211(2):627-645.
[43]	Liu J, Zhou J, Xing D.Phosphatidylinositol 3-Kinase plays a vital role in regulation of rice seed vigor via altering NADPH oxidase activity.Plos One, 2012, 7(3):e33817-e33827.
[44]	Wang E T, Xu X, Zhang L, Zhang H, Lin L, Wang Q, Li Q, Ge S, Lu B R, Wang W, He Z H.Duplication and independent selection of cell-wall invertase genes GIF1 and OsCIN1 during rice evolution and domestication.BMCEvolBiol, 2010, 10(1):108-120.
[45]	孙亚莉, 刘红梅, 徐庆国. 镉胁迫对不同水稻品种种子萌发特性的影响. 中国水稻科学, 2017, 31(4):425-431.
	Sun Y L, Liu H M, Xu Q G.Effects of cadmium stress on rice seed germination characteristics. Chin J Rice Sci, 2017, 31(4):425-431. (in Chinese with English abstract)
[46]	Redoña E D, Mackill D J.Mapping quantitative trait loci for seeding vigor in rice using RFLPs.TheorAppl Genet, 1996, 92(3-4):395-402.
[47]	Regan K L, Siddique K H M, Turner N C, Whan B R. Potential for increasing early vigor and total biomass in spring wheat II characteristics associated with early vigor.CropPastSci, 1992, 43(3):541-553.
[48]	Huang X Z, Qian Q, Liu Z B, Sun H Y, He S Y, Luo D, Xia G M, Chu C C, Li J Y, Fu X D.Natural variation at the DEP1 locus enhances grain yield in rice.NatGenet, 2009, 41(4):494-497.
[49]	Song X J, Huang W, Shi M, Zhu M Z, Lin H X.A QTL for rice grain width and weight encodes a previously unknown RING-type E3 ubiquitin ligase.NatGenet, 2007, 39(5):623-630.
[50]	Wang W Q, Liu S J, Song S Q, Møller I M.Proteomics of seed development, desiccation tolerance, germination and vigor.Plant Physiol&Biochem, 2015, 86(86):1-15.
[51]	Han L Z, Zhang Y Y, Qiao Y L, Cao G L, Zhang S Y, Kim J H, Koh H J.Genetic and QTL analysis for low-temperature vigor of germination in rice.Acta GenetSin, 2006, 33(11):998-1006.
[52]	Dang X J, Thi T G, Dong G S, Wang H, Edzesi W M, Hong D L.Genetic diversity and association mapping of seed vigor in rice ( Oryza sativa L.).Planta, 2014, 239(6):1309-1319.
[53]	Liu L F, Lai Y Y, Cheng J P, Wang L, Du W L, Wang Z F, Zhang H S.Dynamic quantitative trait locus analysis of seed vigor at three maturity stages in rice.MolBreed, 2014, 34(2):501-510.
[54]	Cui K H, Peng S B, Xing Y Z, Xu C G, Yu S B, Zhang Q.Molecular dissection of seedling-vigor and associated physiological traits in rice.TheorAppl Genet, 2002, 105(5):745-753.
[55]	Xie L X, Tan Z W, Zhou Y, Xu R B, Feng L B, Xing Y Z, Qi X Q.Identification and fine mapping of quantitative trait loci for seed vigor in germination and seedling establishment in rice.JIntegrPlant Biol, 2014, 56(8):749-759.
[56]	Abe A, Takagi H, Fujibe T, Aya K, Kojima M, Sakakibara H, Uemura A, Matsuoka M, Terauchi R.OsGA20ox1, a candidate gene for a major QTL controlling seedling vigor in rice.TheorAppl Genet, 2012, 125(4):647-657.
[57]	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.QTL identification and fine mapping for seed storability in rice ( Oryza sativa L.).Euphytica, 2017, 213(6):127-138.
[58]	Zhang A P, Liu C L, Chen G, Hong K, Gao Y, Tian P, Peng Y L, Zhang B, Ruan B P, Jiang H Z, Guo L B, Qian Q, Gao Z Y.Genetic analysis for rice seedling vigor and fine mapping of a major QTL qSSL1b for seedling shoot length.BreedSci, 2017, 67(3):307-315.
[59]	Li M, Sun P L, Zhou H J, Yu S B.Identification of quantitative trait loci associated with germination using chromosome segment substitution lines of rice ( Oryza sativa L.).TheorApplGenet, 2011, 123(3):411-420.
[60]	Song S Y, Dai X Y, Zhang W H.A rice F-box gene,OsFbx352, is involved in glucose-delayed seed germination in rice. JExpBot, 2012, 63(15):5559-5568.
[61]	Chang P F, Jinn T L, Huang W K, Chen Y, Chang H M, Wang C W.Induction of a cDNA clone from rice encoding a class II small heat shock protein by heat stress, mechanical injury, and salicylic acid.Plant Sci, 2007, 172(1):64-75.
[62]	Yuan J, Chen D, Ren Y J, Zhang X L, Zhao J.Characteristic and expression analysis of a Metallothionein gene,OsMT2b, down-regulated by cytokinin suggests functions in root development and seed embryo germination of rice. Plant Physiol, 2008, 146(4):1637-1650.
[63]	Challam C, Kharshing G A, Yumnam J S, Tyagi W.Association of qLTG3-1 with germination stage cold tolerance in diverse rice germplasm from the Indian subcontinent.Plant GenetResour, 2013, 11(3):206-211.
[64]	Wang Z F, Wang J F, Bao Y M, Wang F H, Zhang H S.Quantitative trait loci analysis for rice seed vigor during the germination stage. JZhejiang Univ: SciB, 2010, 11(12):958-964.
[65]	Gu X Y, Foley M E, Horvath D P, Anderson J V, Feng J H, Zhang L H, Mowry C R, Ye H, Suttle J C, Kadowaki K I, Chen Z X.Association between seed dormancy and pericarp color is controlled by a pleiotropic gene that regulates abscisic acid and flavonoid synthesis in weedy red rice.Genetics, 2011, 189(4):1515-1524.
[66]	Sugimoto K, Takeuchi Y, Ebana K, Miyao A, Hirochika H, Hara N, Ishiyama K, Kobayashi M, Ban Y, Hattori T, Yano M.Molecular cloning of Sdr4 , a regulator involved in seed dormancy and domestication of rice.Proc Natl AcadSci USA, 2010, 107(13):5792-5797.
[67]	Han L Z, Qiao Y L, Zhang S Y, Zhang Y Y, Cao G L, Kim J, Lee K, Koh H.Identification of quantitative trait loci for cold response of seedling vigor traits in rice.Acta GenetSin, 2007, 34(3):239-246.
[68]	Cao L Y, Zhu J, Ren L F, Zhao S T, Yan Q C.Mapping QTLs and epistasis for seeding vigor in rice ( Oryza sativa L.).ActaAgronoSin, 2012, 28(6):809-815.
[69]	Anandan A, Anumalla M, Pradhan S K, Ali J.Population structure, diversity and trait association analysis in rice ( Oryza sativa L.) germplasm for early seedling vigor (ESV) using trait linked SSR markers.PLoS One, 2016, 11(3):e0152406-e0152427.
[70]	陈利华, 万杉. 不同温度条件下水稻种子活力QTL的定位分析. 植物科学学报, 2005, 23(2):125-130.
	Chen L H, Wan S.Mapping of QTL controlling seed vitality in rice under different temperature conditions.JPlant Sci, 2005, 23(2):125-130. (in Chinese with English abstract)
[71]	Sabouri A, Sabouri H, Ocampo M D.Genetic analysis seedling vigour under osmotic stress in rice by QTL mapping.RussAgricSci, 2012, 38(5-6):423-429.
[72]	Huang Z, Yu T, Su L, Yu S B, Zhang Z H, Zhu Y G.Identification of chromosome regions associated with seedling vigor in rice.Acta GenetSin, 2004, 31(6):596-603.
[73]	林英莉, 刘国戈. 浅析种子活力的影响因素. 种子世界, 2012, 12(6):17.
	Lin Y L, Liu G G.Influencing factors of seed vigor.Seed World, 2012, 12(6):17. (in Chinese)
[74]	李俊周, 李梦琪, 刘磊, 刘娟, 杜彦修, 赵全志. 水稻种子H2O2流速和种子活力的关系研究. 华北农学报, 2017, 32(4):189-194.
	Li J Z, Li M Q, Liu L, Liu J, Du Y X, Zhao Q Z.Study on the relationship between H2O2 velocity and seed vigor of rice seeds.ActaAgricBorSin, 2017, 32(4):189-194. (in Chinese with English abstract)
[75]	王铁固, 张怀胜, 马娟, 佘宁安, 陈士林. 玉米种子活力与产量的相关分析. 安徽农业科学, 2012, 40(10):5848-5849.
	Wang T G, Zhang H S, Ma J, Ren N A, Chen S L.Correlation analysis between seed vigor and maize yield.JAnhui AgricSci, 2012, 40(10):5848-5849. (in Chinese with English abstract)

性状 Character		染色体 Chr.	参考文献 Reference	性状 Character		染色体 Chr.	参考文献 Reference
发芽指数Germinationindex	qGI1	1	[53]	种子休眠性	OslecRK	4	[40]
	qGI-1	1	[64]	Seed dormancy	qSD7-1/qPC7	7	[65]
	qGI3	3	[53]		Sdr4	8	[66]
	qGI7	7	[53]		OsFbx352	10	[60]
	qGI-7	7	[64]	根长 Root length	qSV1	1	[55]
	qGI8	8	[53]		qSRL2	2	[58]
	qGI10	10	[53]		unnamed	2	Gramene
	qGI-11	11	[64]		qSRL3	3	[58]
发芽势	qGP3	3	[53]		qSRL5	5	[58]
Germination potentiality	qGP5	5	[53]		qSV5c	5	[55]
	qGP10	10	[53]		unnamed	5	Gramene
发芽率 Germination rate	qGR1	1	[53]		qSV6a	6	[55]
	qGR-1	1	[64]		qSV6b	6	[55]
	qGR-2	2	[64]		qSRL7	7	[58]
	qGR3-1	3	[54]		unnamed	7	Gramene
	qGR3-2	3	[54]		qSV8-2	8	[70]
	qGR3-3	3	[54]		qRL8c	8	[71]
	qGR5	5	[53]		qSRL10	10	[58]
	qGR5-1	5	[54]		qRL-10	10	[71]
	qGR5-2	5	[54]	鲜质量	unnamed	1	Gramene
	PI3K	5	[43]	Seedling wet weight	unnamed	1	Gramene
	OsMT2b	5	[62]		unnamed	1	Gramene
	qGR6-1	6	[54]		unnamed	2	Gramene
	qGR6-2	6	[54]		qSWW4	4	[58]
	qGR7-1	7	[54]		unnamed	4	Gramene
	qGR7	7	[53]		qSWW5	5	[58]
	qGR8	8	[53]		unnamed	6	Gramene
	OsPIMT1	8	[41]		qSWW7	7	[58]
	qGP9	9	[57]		unnamed	7	Gramene
	qGR10	10	[53]		qSWW8	8	[58]
	qGR-11	11	[64]		qSWW12	12	[58]
苗长 Seedling length	unnamed	1	Gramene	干质量	unnamed	1	Gramene
	qSSL1b	1	[58]	Seedling dry weight	qTDW1-1	1	[54]
	qCSH1	1	[67]		qTDW1-2	1	[54]
	qCSH2	2	[67]		unnamed	2	Gramene
	qSL3-2	3	[68]		qTDW3-1	3	[54]
	qPHS4	4	[56]		qSDW3	3	[58]
	qSV5a	5	[55]		qSDW4	4	[58]
	qSV-5	5	Gramene		qTDW5-1	5	[54]
	unnamed	5	[52]		qTDW5-2	5	[54]
	unnamed	7	[69]		unnamed	5	[72]
	qSV8	8	[55]		qTDW6-1	6	[54]
	qSV11	11	[55]		qTDW6-2	6	[54]
	qSL11	11	[68]		qTDW9-1	6	[54]
	qCSH12	12	[67]		qSDW9	9	[58]
低温发芽势Germination vigor	qLTG3-1	3	[63]	耐热性Heat tolerance	Oshsp18.2	1	[61]
at low-temperature

性状 Character		染色体 Chr.	参考文献 Reference	性状 Character		染色体 Chr.	参考文献 Reference
发芽指数Germinationindex	qGI1	1	[53]	种子休眠性	OslecRK	4	[40]
	qGI-1	1	[64]	Seed dormancy	qSD7-1/qPC7	7	[65]
	qGI3	3	[53]		Sdr4	8	[66]
	qGI7	7	[53]		OsFbx352	10	[60]
	qGI-7	7	[64]	根长 Root length	qSV1	1	[55]
	qGI8	8	[53]		qSRL2	2	[58]
	qGI10	10	[53]		unnamed	2	Gramene
	qGI-11	11	[64]		qSRL3	3	[58]
发芽势	qGP3	3	[53]		qSRL5	5	[58]
Germination potentiality	qGP5	5	[53]		qSV5c	5	[55]
	qGP10	10	[53]		unnamed	5	Gramene
发芽率 Germination rate	qGR1	1	[53]		qSV6a	6	[55]
	qGR-1	1	[64]		qSV6b	6	[55]
	qGR-2	2	[64]		qSRL7	7	[58]
	qGR3-1	3	[54]		unnamed	7	Gramene
	qGR3-2	3	[54]		qSV8-2	8	[70]
	qGR3-3	3	[54]		qRL8c	8	[71]
	qGR5	5	[53]		qSRL10	10	[58]
	qGR5-1	5	[54]		qRL-10	10	[71]
	qGR5-2	5	[54]	鲜质量	unnamed	1	Gramene
	PI3K	5	[43]	Seedling wet weight	unnamed	1	Gramene
	OsMT2b	5	[62]		unnamed	1	Gramene
	qGR6-1	6	[54]		unnamed	2	Gramene
	qGR6-2	6	[54]		qSWW4	4	[58]
	qGR7-1	7	[54]		unnamed	4	Gramene
	qGR7	7	[53]		qSWW5	5	[58]
	qGR8	8	[53]		unnamed	6	Gramene
	OsPIMT1	8	[41]		qSWW7	7	[58]
	qGP9	9	[57]		unnamed	7	Gramene
	qGR10	10	[53]		qSWW8	8	[58]
	qGR-11	11	[64]		qSWW12	12	[58]
苗长 Seedling length	unnamed	1	Gramene	干质量	unnamed	1	Gramene
	qSSL1b	1	[58]	Seedling dry weight	qTDW1-1	1	[54]
	qCSH1	1	[67]		qTDW1-2	1	[54]
	qCSH2	2	[67]		unnamed	2	Gramene
	qSL3-2	3	[68]		qTDW3-1	3	[54]
	qPHS4	4	[56]		qSDW3	3	[58]
	qSV5a	5	[55]		qSDW4	4	[58]
	qSV-5	5	Gramene		qTDW5-1	5	[54]
	unnamed	5	[52]		qTDW5-2	5	[54]
	unnamed	7	[69]		unnamed	5	[72]
	qSV8	8	[55]		qTDW6-1	6	[54]
	qSV11	11	[55]		qTDW6-2	6	[54]
	qSL11	11	[68]		qTDW9-1	6	[54]
	qCSH12	12	[67]		qSDW9	9	[58]
低温发芽势Germination vigor	qLTG3-1	3	[63]	耐热性Heat tolerance	Oshsp18.2	1	[61]
at low-temperature