Fine Mapping of qGL1.1, a Minor QTL for Grain Length, Using Near Isogenic Lines Derived from Residual Heterozygotes in Rice

doi:10.16819/j.1001-7216.2020.9125

Abstract

Abstract:

【Objective】The objective of this study is to fine-map qTGW1.1b which was previously mapped in a 521.8-kb region on the long arm of rice chromosome 1.【Method】In this study, two BC₂F₉ plants carrying qTGW1.1 and qTGW1.2, respectively, was crossed and produced an F₄ population. Three plants carrying sequential heterozygous segments in the interval Wn28826-RM1231 were selected and selfed to develop three sets of near isogenic lines (NILs) comprising the two homozygous genotypes. The NILs in F_5:6 were planted in Hangzhou in 2017 and the 1000-grain weight, grain length and grain width were measured. The effect of qTGW1.1b was validated by two-way analysis of variance (ANOVA) using SAS program. Then six new sequential residual heterozygotes carrying smaller heterozygous segments overlapped in the interval Wn28826-RM1231 were identified. Six sets of pairwise NILs in F_8:9 were planted in Lingshui in 2018 and the three traits were measured. Phenotypic differences between the two homozygous genotypic groups were analyzed using ANOVA.【Results】The allelic direction of qTGW1.1b remained consistent and the genetic effects were stable in the two trials. The Milyang 46 allele could significantly increase grain length and 1000-grain weight by 0.027 mm and 0.17 g with the contribution up to 27.12% and 19.09%, respectively.【Conclusion】Considering that qTGW1.1b stably showed significant effect to grain length but not to grain width in both previous and this study, qTGW1.1b was renamed as qGL1.1. As a result, qGL1.1 was delimited into a 76.8-kb region franked by Wn29077 and Wn29154 based on comparing of the segregating regions among the six sets of NILs.

Key words: rice, near isogenic line, minor effect, grain length, fine mapping

摘要：

【目的】本研究旨在对前期在水稻第1染色体长臂521.8 kb的区间内定位到的qTGW1.1b进行精细定位。【方法】从qTGW1.1和qTGW1.2所在区间分别呈杂合的2个BC₂F₉单株配组衍生的F₄群体中,筛选到Wn28826- RM1231区间内杂合片段呈梯系排列的3个单株,构建了3套F_5:6近等基因系。2017年种植于浙江杭州,考查千粒重、粒长和粒宽。利用SAS软件的GLM程序进行双因素方差分析,对qTGW1.1b的效应进行了验证。在此基础上,筛选出杂合片段更小且呈交迭排列的6个剩余杂合体,发展了6套F_8:9近等基因系,2018年种植于海南陵水。对每套近等基因系中双亲基因型株系的表型差异进行双因素方差分析。【结果】qTGW1.1b在2个试验中对粒长和千粒重均呈极显著差异,效应方向一致且大小稳定。密阳46等位基因能分别增加粒长0.027 mm和提高千粒重0.17 g,贡献率分别达到27.12%和19.09%。【结论】鉴于qTGW1.1b在前后试验中对粒长影响最为显著,而对粒宽作用不显著,故将qTGW1.1b重新命名为qGL1.1。通过比较各套近等基因系的分离区间的基因组位置,最终将qGL1.1定位于Wn29077和Wn29154之间约76.8 kb的区间内。

关键词: 水稻, 近等基因系, 微效作用, 粒长, 精细定位

CLC Number:

Panpan LI, Yujun ZHU, Liang GUO, Jieyun ZHUANG, Yeyang FAN. Fine Mapping of qGL1.1, a Minor QTL for Grain Length, Using Near Isogenic Lines Derived from Residual Heterozygotes in Rice[J]. Chinese Journal OF Rice Science, 2020, 34(2): 125-134.

李盼盼, 朱玉君, 郭梁, 庄杰云, 樊叶杨. 利用剩余杂合体衍生的近等基因系精细定位水稻粒长微效QTL qGL1.1[J]. 中国水稻科学, 2020, 34(2): 125-134.

Figures/Tables 7

References 40

[1]	Xing Y Z, Zhang Q F.Genetic and molecular bases of rice yield[J]. Annual Review of Plant Biology, 2010, 61: 421-442.
[2]	Duan P G, Ni S, Wang J M, Zhang B L, Xu R, Wang Y X, Chen H Q, Zhu X D, Li Y H.Regulation of OsGRF4 by OsmiR396 controls grain size and yield in rice[J]. Nature Plants, 2016, 2: 15203.
[3]	Hu J, Wang Y X, Fang Y X, Zeng L J, Xu J, Yu H P, Shi Z Y, Pan J J, Zhang D, Kang S J, Zhu L, Dong G J, Guo L B, Zeng D L, Zhang G H, Xie L H, Xiong G S, Li J Y, Qian Q.A rare allele of GS2 enhances grain size and grain yield in rice[J]. Molecular Plant, 2015, 8: 1455-1465.
[4]	Yu J P, Xiong H Y, Zhu X Y, Zhang H L, Li H H, Miao J L, Wang W S, Tang Z S, Zhang Z Y, Yao G X, Zhang Q, Pan Y H, Wang X, Rashid M A R, Li J J, Gao Y M, Li Z K, Yang W C, Fu X D, Li Z C. OsLG3 contributing to rice grain length and yield was mined by Ho-LAMap[J]. BMC Biology, 2017, 15: 28.
[5]	Yu J P, Miao J L, Zhang Z Y, Xiong H Y, Zhu X Y, Sun X M, Pan Y H, Liang Y T, Zhang Q, Abdul Rehman R M, Li J J, Zhang H L, Li Z C. Alternative splicing of OsLG3b controls grain length and yield in japonica rice[J]. Plant Biotechnology Journal, 2018, 16: 1667-1678.
[6]	Liu Q, Han R X, Wu K, Zhang J Q, Ye Y F, Wang S S, Chen J F, Pan Y J, Li Q, Xu X P, Zhou J W, Tao D Y, Wu Y J, Fu X D.G-protein βγ subunits determine grain size through interaction with MADS-domain transcription factors in rice[J]. Nature Communications, 2018, 9: 852.
[7]	Fan C C, Xing Y Z, Mao H L, Lu T T, Han B, Xu C G, Li X H, Zhang Q F.GS3, a major QTL for grain length and weight and minor QTL for grain width and thickness in rice, encodes a putative transmembrane protein[J]. Theoretical and Applied Genetics, 2006, 112: 1164-1171.
[8]	Qi P, Lin Y S, Song X J, Shen J B, Huang W, Shan J X, Zhu M Z, Jiang L W, Gao P J, Lin H X.The novel quantitative trait locus GL3.1 controls rice grain size and yield by regulating Cyclin-T1;3[J]. Cell Research, 2012, 22: 1666-1680.
[9]	Zhang X J, Wang J F, Huang J, Lan H X, Wang C L, Yin C F, Wu Y Y, Tang H J, Qian Q, Li J Y, Zhang H S.Rare allele of OsPPKL1 associated with grain length causes extra-large grain and a significant yield increase in rice[J]. Proceedings of the National Academy of Sciences of the United States of America, 2012, 109(52): 21534-21539.
[10]	Hu Z J, Lu S J, Wang M J, He H H, Sun L, Wang H R, Liu X H, Jiang L, Sun J L, Xin X Y, Kong W, Chu C C, Xue H W, Yang J S, Luo X J, Liu J X.A novel QTL qTGW3 encodes the GSK3/SHAGGY-like kinase OsGSK5/OsSK41 that interacts with OsARF4 to negatively regulate grain size and weight in rice[J]. Molecular Plant, 2018, 11:736-749.
[11]	Xia D, Zhou H, Liu R j, Dan W H, Li P B, Wu B, Chen J X, Wang L Q, Gao G J, Zhang Q L, He Y Q. GL3.3, a novel QTL encoding a GSK3/SHAGGY-like kinase, epistatically interacts with GS3 to produce extra-long grains in rice[J]. Molecular Plant, 2018, 11: 754-756.
[12]	Ying J Z, Ma M, Bai C, Huang X H, Liu J L, Fan Y Y, Song X J.TGW3, a major QTL that negatively modulates grain length and weight in rice[J]. Molecular Plant, 2018, 11: 750-753.
[13]	Wu W G, Liu X Y, Wang M H, Meyer R S, Luo X J, Ndjiondjop M N, Tan L B, Zhang J W, Wu J Z, Cai H W, Sun C Q, Wang X K, Wing R A, Zhu Z F.A single-nucleotide polymorphism causes smaller grain size and loss of seed shattering during African rice domestication[J]. Nature Plants, 2017, 3: 17064.
[14]	Wang A H, Hou Q Q, Si L Z, Huang X H, Luo J H, Lu D F, Zhu J J, Shangguan Y Y, Miao J S, Xie Y F, Wang Y C, Zhao Q, Feng Q, Zhou C C, Li Y, Fan D L, Lu Y Q, Tian Q L, Wang Z X, Han B.The PLATZ transcription factor GL6 affects grain length and number in rice[J]. Plant Physiology, 2019, 180 : 2077-2090.
[15]	Ishimaru K, Hirotsu N, Madoka Y, Murakami N, Hara N, Onodera H, Kashiwagi T, Ujiie K, Shimizu B I, Onishi A, Miyagawa H, Katoh E.Loss of function of the IAA-glucose hydrolase gene TGW6 enhances rice grain weight and increases yield[J]. Nature Genetics, 2013, 45(6): 707-711.
[16]	Si L Z, Chen J Y, Huang X H, Gong H, Luo J H, Hou Q Q, Zhou T Y, Lu T T, Zhu J J, Shangguan Y Y, Chen E W, Gong C X, Zhao Q, Jing Y F, Zhao Y, Li Y, Cui L L, Fan D L, Lu Y Q, Weng Q J, Wang Y C, Zhan Q L, Liu K Y, Wei X H, An K, An G, Han B.OsSPL13 controls grain size in cultivated rice[J]. Nature Genetics, 2016, 48(4): 447-456.
[17]	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[J]. Nature Genetics, 2007, 39(5): 623-630.
[18]	Li Y B, Fan C C, Xing Y Z, Jiang Y H, Luo L J, Sun L, Shao D, Xu C J, Li X H, Xiao J H, He Y Q, Zhang Q F.Natural variation in GS5 plays an important role in regulating grain size and yield in rice[J]. Nature Genetics, 2011, 43(12): 1266-1269.
[19]	Duan P G, Xu J S, Zeng D L, Zhang B L, Geng M F, Zhang G Z, Huang K, Huang L J, Xu R, Ge S, Qian Q, Li Y H.Natural variation in the promoter of GSE5 contributes to grain size diversity in rice[J]. Molecular Plant, 2017, 10: 685-694.
[20]	Liu J F, Chen J, Zheng X M, Wu F Q, Lin Q B, Heng Y Q, Tian P, Cheng Z J, Yu X W, Zhou K N, Zhang X, Guo X P, Wang J L, Wang H Y, Wan J M.GW5 acts in the brassinosteroid signaling pathway to regulate grain width and weight in rice[J]. Nature Plants, 2017, 3: 17043.
[21]	Wang S K, Wu K, Yuan Q B, Liu X Y, Liu Z B, Lin X Y, Zeng R Z, Zhu H T, Dong G J, Qian Q, Zhang G Q, Fu X D.Control of grain size, shape and quality by OsSPL16 in rice[J]. Nature Genetics, 2012, 44(8): 950-954.
[22]	Song X J, Kuroha T, Ayano M, Furuta T, Nagai K, Komeda N, Segami S, Miura K, Ogawa D, Kamura T, Suzuki T, Higashiyama T, Yamasaki M, Mori H, Inukai Y, Wu J Z, Kitano H, Sakakibara H, Jacobsen S E, Ashikari M.Rare allele of a previously unidentified histone H4 acetyltransferase enhances grain weight, yield, and plant biomass in rice[J]. Proceedings of the National Academy of Sciences of the United States of America, 2015, 112(1): 76-81.
[23]	Wang Y X, Xiong G S, Hu J, Jiang L, Yu H, Xu J, 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[J]. Nature Genetics, 2015, 47(8): 944-948.
[24]	Wang S K, Li S, Liu Q, Wu K, Zhang J Q, Wang S S, Wang Y, Chen X B, Zhang Y, Gao C X, Wang F, Huang H X, Fu X D.The OsSPL16-GW7 regulatory module determines grain shape and simultaneously improves rice yield and grain quality[J]. Nature Genetics, 2015, 47(8): 949-954.
[25]	Zhao D S, Li Q F, Zhang C Q, Zhang C, Yang Q Q, Pan L X, Ren X Y, Lu J, Gu M H, Liu Q Q.GS9 acts as a transcriptional activator to regulate rice grain shape and appearance quality[J]. Nature Communications, 2018, 9: 1240.
[26]	Li N, Xu R, Duan P G, Li Y H.Control of grain size in rice[J]. Plant Reproduction, 2018, 31: 237-251.
[27]	刘喜,牟昌铃,周春雷,程治军,江玲,万建民. 水稻粒型基因克隆和调控机制研究进展[J]. 中国水稻科学,2018, 32(1): 1-11.
	Liu X, Mou C L, Zhou C L, Cheng Z J, Jiang L, Wang J M.Research progress on cloning and regulation mechanism of rice grain shape genes[J]. Chinese Journal of Rice Science, 2018, 32(1): 1-11. (in Chinese with English abstract)
[28]	Mackay T F C, Stone E A, Ayroles J F. The genetics of quantitative traits: challenges and prospects[J]. Nature Reviews Genetics, 2009, 10: 565-577.
[29]	Kumar J, Gupta D S, Gupta S, Dubey S, Gupta P, Kumar S.Quantitative trait loci from identification to exploitation for crop improvement[J]. Plant Cell Reports, 2017, 36: 1187-1213.
[30]	Yamamoto T, Yonemaru J, Yano M.Towards the understanding of complex traits in rice: substantially or superficially[J]. DNA Research, 2009, 16: 141-154.
[31]	Zhang H W, Fan Y Y, Zhu Y J, Chen J Y, Yu S B, Zhuang J Y.Dissection of the qTGW1.1 region into two tightly-linked minor QTLs having stable effects for grain weight in rice[J]. BMC Genetics, 2016, 17: 98.
[32]	Guo L, Wang K, Chen J Y, Huang D R, Fan Y Y, Zhuang J Y.Dissection of two quantitative trait loci for grain weight linked in repulsion on the long arm of chromosome 1 of rice (Oryza sativa L.)[J]. The Crop Journal, 2013, 1: 70-76.
[33]	Zheng K L, Huang N, Bennett J, Khush G S.PCR-based marker-assisted selection in rice breeding//IRRI Discussion Paper Series No. 12. Los Banos: International Rice Research Institute, 1995.
[34]	Chen X, Temnykh S, Xu Y, Cho Y G, McCouch S R. Development of a microsatellite framework map providing genome-wide coverage in rice (Oryza sativa L.)[J]. Theoretical and Applied Genetics, 1997, 95: 553-567.
[35]	SAS Institute Inc.SAS/STAT User’s Guide[M]. Cary, NC: SAS Institute, 1999.
[36]	Dai W M, Zhang K Q, Wu J R, Wang L, Duan B W, Zheng K L, Cai R, Zhuang J Y.Validating a segment on the short arm of chromosome 6 responsible for genetic variation in the hull silicon content and yield traits of rice[J]. Euphytica, 2008, 160: 317-324.
[37]	Abiola O, Angel J M, Avner P, Bachmanov A A, Belknap J K, Bennett B, Blankenhorn E P, Blizard D A, Bolivar V, Brockmann G A, Buck K J, Bureau J F, Casley W L, Chesler E J, Cheverud J M, Churchill G A, Cook M, Crabbe J C, Crusio W E, Darvasi A, Haan G D, Dermant P, Doerge R W, Elliot R W, Farber C R, Flaherty L, Flint J, Gershenfeld H, Gibson J P, Gu J, Gu W, Himmelbauer H, Hitzemann R, Hsu H C, Hunter K, Iraqi F F, Jansen R C, Johnson T E, Jones B C, Kempermann G, Lammert F, Lu L, Manly K F, Matthews D B, Medrano J F, Mehrabian M, Mittlemann G, Mock B A, Mogil J S, Montagutelli X, Morahan G, Mountz J D, Nagase H, Nowakowski R S, O'Hara B F, Osadchuk A V, Paigen B, Palmer A A, Peirce J L, Pomp D, Rosemann M, Rosen G D, Schalkwyk L C, Seltzer Z, Settle S, Shimomura K, Shou S, Sikela J M, Siracusa L D, Spearow J L, Teuscher C, Threadgill D W, Toth L A, Toye A A, Vadasz C, Van Zant G, Wakeland E, Williams R W, Zhang H G, Zou F. Complex Trait Consortium. The nature and identification of quantitative trait loci: A community’s view[J]. Nature Reviews Genetics, 2003, 4(11): 911-916.
[38]	Wang L L, Chen Y Y, Guo L, Zhang H W, Fan Y Y, Zhuang J Y.Dissection of qTGW1.2 to three QTLs for grain weight and grain size in rice (Oryza sativa L.)[J]. Euphytica, 2015, 202: 119-127.
[39]	Dong Q, Zhang Z H, Wang L L, Zhu Y J, Fan Y Y, Mou T M, Ma L Y, Zhuang J Y.Dissection and fine-mapping of two QTL for grain size linked in a 460-kb region on chromosome 1 of rice[J]. Rice, 2018, 11: 44.
[40]	Wang W H, Wang L L, Zhu Y J, Fan Y Y, Zhuang J Y.Fine-mapping of qTGW1.2a, a quantitative trait locus for 1000-grain weight in rice[J]. Rice Science, 2019, 26(4): 220-228.

标记名称	类型	限制性内切酶	正向引物（5'-3'）	反向引物（5'-3'）
Name	Type	Restriction enzyme	Forward primer (5'-3')	Reverse primer (5'-3')
Wn28826	CAPS	BstN I	GACAAGTTGGGATAATTCTTCGAT	TAACGTGTCGATCTCTGACC
Wn28893	dCAPS	Hha I	GATCGCTCCCTTGTATACGCTGA	CCATTCCGCCCGGTTGATGAAACGC
Wn28944	InDel		CATTACAAGGTAAATTGTAGATTGG	TCATTTAGGGATTATGTTGGTC
Wn28990	InDel		AGTTTATAAATCCGAAGCCAT	AGCACAAATAAGTAATTATGCCTA
Wn29048	dCAPS	N1a III	GAATAAGTCCACTTTACGCATCTTTCTCA	GGATCAAGATTTTCCGTATTGCAG
Wn29077	dCAPS	Kpn I	CAGTTCACGGGATACGAAGC	CAGTTTGACCATCCTCTAAGCAAAGGGTA
Wn29125	dCAPS	Sty I	AAGTGTGTACGGTCAAATGTTTGCCA	ACGTCAGTCAAAACAAATACGG
Wn29154	CAPS	Xba I	TGGATTAATTAGGCTAGGTAGACA	TTCTCCCTCTCGTGATCGC

标记名称	类型	限制性内切酶	正向引物（5'-3'）	反向引物（5'-3'）
Name	Type	Restriction enzyme	Forward primer (5'-3')	Reverse primer (5'-3')
Wn28826	CAPS	BstN I	GACAAGTTGGGATAATTCTTCGAT	TAACGTGTCGATCTCTGACC
Wn28893	dCAPS	Hha I	GATCGCTCCCTTGTATACGCTGA	CCATTCCGCCCGGTTGATGAAACGC
Wn28944	InDel		CATTACAAGGTAAATTGTAGATTGG	TCATTTAGGGATTATGTTGGTC
Wn28990	InDel		AGTTTATAAATCCGAAGCCAT	AGCACAAATAAGTAATTATGCCTA
Wn29048	dCAPS	N1a III	GAATAAGTCCACTTTACGCATCTTTCTCA	GGATCAAGATTTTCCGTATTGCAG
Wn29077	dCAPS	Kpn I	CAGTTCACGGGATACGAAGC	CAGTTTGACCATCCTCTAAGCAAAGGGTA
Wn29125	dCAPS	Sty I	AAGTGTGTACGGTCAAATGTTTGCCA	ACGTCAGTCAAAACAAATACGG
Wn29154	CAPS	Xba I	TGGATTAATTAGGCTAGGTAGACA	TTCTCCCTCTCGTGATCGC

性状	试验		名称	平均	标准差	变异系数	范围	偏斜度	峰度
Trait	Trial		Name	Mean	SD	CV	Range	Skewness	Kurtosis
千粒重 1000-grain weight /g	杭州		Y1	25.55	0.34	0.013	24.92~26.27	0.30	-0.59
	Hangzhou		Y2	26.74	0.36	0.013	25.84~27.92	0.45	1.15
			Y3	25.67	0.26	0.010	25.05~26.17	-0.27	-0.38
	陵水		LP1	30.16	0.36	0.012	29.42~30.89	0.07	-0.34
	Lingshui		LP2	29.25	0.61	0.021	27.67~30.36	-0.50	-0.18
			LP3	29.93	0.29	0.010	29.26~30.57	0.07	-0.43
			LP4	30.74	0.31	0.010	30.04~31.37	0.24	-0.52
			LP5	29.81	0.48	0.016	28.35~30.79	-0.31	0.28
			LP6	29.73	0.52	0.017	28.69~30.68	-0.27	-0.71
粒长 Grain length / mm	杭州		Y1	8.120	0.057	0.007	7.999~8.261	-0.08	-0.06
	Hangzhou		Y2	8.216	0.057	0.007	8.106~8.362	0.58	0.28
			Y3	8.136	0.050	0.006	8.033~8.264	0.02	-0.34
	陵水		LP1	8.106	0.037	0.005	8.034~8.191	0.30	-0.39
	Lingshui		LP2	8.114	0.040	0.005	8.040~8.201	0.07	-0.87
			LP3	8.097	0.041	0.005	8.003~8.199	0.16	-0.10
			LP4	8.268	0.047	0.006	8.138~8.358	-0.44	0.02
			LP5	8.094	0.054	0.007	7.978~8.217	0.13	-0.47
			LP6	8.164	0.045	0.005	8.051~8.245	-0.15	-0.29
粒宽 Grain width / mm	杭州		Y1	2.958	0.022	0.008	2.899~3.024	0.13	0.81
	Hangzhou		Y2	3.053	0.018	0.006	3.014~3.103	0.35	0.43
			Y3	2.924	0.024	0.008	2.875~2.972	0.14	-0.71
	陵水		LP1	3.288	0.014	0.004	3.244~3.312	-0.67	0.82
	Lingshui		LP2	3.218	0.044	0.014	3.091~3.290	-0.78	0.48
			LP3	3.273	0.022	0.007	3.217~3.306	-0.98	0.49
			LP4	3.296	0.014	0.004	3.267~3.324	-0.26	-0.63
			LP5	3.265	0.031	0.009	3.156~3.313	-1.81	4.27
		LP6		3.236	0.040	0.012	3.129~3.306	-0.66	-0.14

性状	试验		名称	平均	标准差	变异系数	范围	偏斜度	峰度
Trait	Trial		Name	Mean	SD	CV	Range	Skewness	Kurtosis
千粒重 1000-grain weight /g	杭州		Y1	25.55	0.34	0.013	24.92~26.27	0.30	-0.59
	Hangzhou		Y2	26.74	0.36	0.013	25.84~27.92	0.45	1.15
			Y3	25.67	0.26	0.010	25.05~26.17	-0.27	-0.38
	陵水		LP1	30.16	0.36	0.012	29.42~30.89	0.07	-0.34
	Lingshui		LP2	29.25	0.61	0.021	27.67~30.36	-0.50	-0.18
			LP3	29.93	0.29	0.010	29.26~30.57	0.07	-0.43
			LP4	30.74	0.31	0.010	30.04~31.37	0.24	-0.52
			LP5	29.81	0.48	0.016	28.35~30.79	-0.31	0.28
			LP6	29.73	0.52	0.017	28.69~30.68	-0.27	-0.71
粒长 Grain length / mm	杭州		Y1	8.120	0.057	0.007	7.999~8.261	-0.08	-0.06
	Hangzhou		Y2	8.216	0.057	0.007	8.106~8.362	0.58	0.28
			Y3	8.136	0.050	0.006	8.033~8.264	0.02	-0.34
	陵水		LP1	8.106	0.037	0.005	8.034~8.191	0.30	-0.39
	Lingshui		LP2	8.114	0.040	0.005	8.040~8.201	0.07	-0.87
			LP3	8.097	0.041	0.005	8.003~8.199	0.16	-0.10
			LP4	8.268	0.047	0.006	8.138~8.358	-0.44	0.02
			LP5	8.094	0.054	0.007	7.978~8.217	0.13	-0.47
			LP6	8.164	0.045	0.005	8.051~8.245	-0.15	-0.29
粒宽 Grain width / mm	杭州		Y1	2.958	0.022	0.008	2.899~3.024	0.13	0.81
	Hangzhou		Y2	3.053	0.018	0.006	3.014~3.103	0.35	0.43
			Y3	2.924	0.024	0.008	2.875~2.972	0.14	-0.71
	陵水		LP1	3.288	0.014	0.004	3.244~3.312	-0.67	0.82
	Lingshui		LP2	3.218	0.044	0.014	3.091~3.290	-0.78	0.48
			LP3	3.273	0.022	0.007	3.217~3.306	-0.98	0.49
			LP4	3.296	0.014	0.004	3.267~3.324	-0.26	-0.63
			LP5	3.265	0.031	0.009	3.156~3.313	-1.81	4.27
		LP6		3.236	0.040	0.012	3.129~3.306	-0.66	-0.14

试验 Trial	名称 Name	分离区间 Segregating region	性状 Trait	表型值(平均值±标准差)		P	A	R² / %
				Phenotype (Mean±SD)
				NIL^ZS97	NIL^MY46
杭州	Y1	Wn28826-RM1231	TGW/g	25.42±0.30	25.69±0.32	0.0017	0.13	8.99
Hangzhou			GL/mm	8.098±0.062	8.143±0.041	0.0018	0.023	8.13
			GW/mm	2.950±0.019	2.966±0.023	0.0048	0.008	8.19
	Y2	Wn28990-RM1231	TGW/g	26.68±0.32	26.78±0.38	0.2372
			GL/mm	8.201±0.050	8.231±0.061	0.0343	0.015	5.33
			GW/mm	3.049±0.017	3.056±0.019	0.1553
	Y3	Wn29154-RM1231	TGW/g	25.65±0.25	25.68±0.28	0.6181
			GL/mm	8.143±0.053	8.129±0.046	0.2623
			GW/mm	2.922±0.023	2.927±0.025	0.3944
陵水	LP1	Wn28826-Wn28893	TGW/g	30.19±0.39	30.12±0.34	0.4894
Lingshui			GL/mm	8.110±0.034	8.104±0.040	0.5480
			GW/mm	3.291±0.013	3.286±0.014	0.1920
	LP2	Wn28826-Wn28990	TGW/g	29.32±0.58	29.19±0.64	0.4124
			GL/mm	8.123±0.042	8.105±0.037	0.1011
			GW/mm	3.224±0.041	3.212±0.046	0.3026
	LP3	Wn28990-Wn29048	TGW/g	30.00±0.27	29.85±0.29	0.0394	-0.08	4.30
			GL/mm	8.109±0.043	8.086±0.035	0.0256	-0.012	5.25
			GW/mm	3.275±0.022	3.272±0.023	0.6138
	LP4	Wn29048-Wn29125	TGW/g	30.56±0.25	30.91±0.27	<0.0001	0.17	19.09
			GL/mm	8.241±0.043	8.296±0.033	<0.0001	0.027	27.12
			GW/mm	3.297±0.015	3.295±0.014	0.7289
	LP5	Wn29125-RM1231	TGW/g	29.84±0.44	29.78±0.52	0.6563
			GL/mm	8.083±0.049	8.105±0.057	0.1154
			GW/mm	3.268±0.024	3.262±0.036	0.4580
	LP6	Wn29154-RM1231	TGW/g	29.67±0.49	29.78±0.55	0.4381
			GL/mm	8.177±0.042	8.151±0.043	0.0206	-0.013	6.03
			GW/mm	3.227±0.043	3.245±0.034	0.0711
A-密阳46等位基因取代珍汕97等位基因所产生的遗传效应。R²-效应对表型方差的贡献率。 A, Additive effect of replacing a Zhenshan 97 allele with a Milyang 46 allele. R², Proportion of phenotypic variance explained by the QTL effect.