利用百农AK58/碧蚂4号重组自交系定位小麦品质性状QTL

doi:10.7668/hbnxb.20193411

摘要/Abstract

摘要：

为发掘小麦品质性状相关数量性状位点(QTL),以小麦骨干亲本百农AK58和碧蚂4号衍生的包含248个株系的重组自交系群体为材料,利用4个环境获得的蛋白质含量、湿面筋含量、淀粉含量、沉降值和延伸性等品质参数及已构建的单核苷酸多态性(SNP)高密度遗传连锁图谱进行QTL定位。通过完备区间作图法共检测到60个品质性状相关QTL,分布于除6B以外的20对小麦染色体上。有21个QTL可以在2个或2个以上环境中被检测到,其中15个QTL的增效基因来自百农AK58,6个QTL的增效基因来自碧蚂4号。4A染色体上116.4~139.0 cM(629.36~701.53 Mb)是一个QTL簇,该区段同时定位了与蛋白质含量(QGpc.his-4A-2)、湿面筋含量(QWgc.his-4A-2)、沉降值(QSv.his-4A)和延伸性(QEx.his-4A)相关的QTL。分析重组自交系群体中1BL/1RS易位对品质性状的影响,发现1BL/1RS易位有助于提高籽粒蛋白质含量、湿面筋含量和沉降值,对淀粉含量有一定的负效应,推测1BL/1RS易位系对品质性状的作用可能和其遗传背景相关。

关键词: 小麦, 品质性状, QTL定位, 1BL/1RS

Abstract:

In order to find the loci associated with quality traits in wheat,the mapping of quantitative trait loci(QTL)was conducted using the high-density genetic linkage map based on single nucleotide polymorphism(SNP)and the quality parameters surveyed in four environments including grain protein content,wet gluten content,starch content,sedimentation value and extensibility with the recombinant inbred line population derived from Bainong AK58/Bima 4.A total of 60 QTLs were identified by inclusive composite interval mapping,which were distributed on 20 chromosomes except for 6B.21 QTLs were found in two or more environments,of which the positive alleles of 15 QTLs were derived from Bainong AK58 and the positive genes of the remaining 6 QTLs were derived from Bima 4. A QTL cluster was found on chromosome 4A in the interval of 116.4—139.0 cM(629.36—701.53 Mb),and several QTLs such as QGpc.his-4A-2,QWgc.his-4A-2,QSv.his-4A and QEx.his-4A were identified in this region.Furthermore,1BL/1RS translocation caused improvement of protein content,wet gluten content and sedimentation value,but showed a negative effect on starch content.It is supposed that the impact of 1BL/1RS translocation lines on quality may be associated with their genetic background.

Key words: Wheat, Quality traits, QTL mapping, 1BL/1RS

张德华, 徐鑫, 王玉洁, 张自阳, 李小军. 利用百农AK58/碧蚂4号重组自交系定位小麦品质性状QTL[J]. 华北农学报, 2023, 38(1): 23-31. doi: 10.7668/hbnxb.20193411.

ZHANG Dehua, XU Xin, WANG Yujie, ZHANG Ziyang, LI Xiaojun. Mapping of Quantitative Trait Loci for Quality Traits Using Recombinant Inbred Line Population Derived from Bainong AK58/Bima 4 in Wheat[J]. Acta Agriculturae Boreali-Sinica, 2023, 38(1): 23-31. doi: 10.7668/hbnxb.20193411.

http://www.hbnxb.net/CN/Y2023/V38/I1/23

图/表 6

参考文献 27

[1]	刘锐, 魏益民, 张波. 小麦蛋白质与面条品质关系的研究进展[J]. 麦类作物学报, 2011, 31(6):1183-1187.doi:10.7606/j.issn.1009-1041.2011.06.033. doi: 10.7606/j.issn.1009-1041.2011.06.033
	Liu R, Wei Y M, Zhang B. Review on relationship between wheat protein and noodle quality[J]. Journal of Triticeae Crops, 2011, 31(6):1183-1187.
[2]	Li J, Cui F, Ding A M, Zhao C H, Wang X Q, Wang L, Bao Y G, Qi X L, Li X F, Gao J R, Feng D S, Wang H G. QTL detection of seven quality traits in wheat using two related recombinant inbred line populations[J]. Euphytica, 2012, 183(2):207-226.doi:10.1007/s10681-011-0448-4. doi: 10.1007/s10681-011-0448-4 URL
[3]	Goel S, Singh K, Singh B, Grewal S, Dwivedi N, Alqarawi A A, Abd Allah E F, Ahmad P, Singh N K. Analysis of genetic control and QTL mapping of essential wheat grain quality traits in a recombinant inbred population[J]. PLoS One, 2019, 14(3):e0200669.doi:10.1371/journal.pone.0200669. doi: 10.1371/journal.pone.0200669 URL
[4]	Huang X Q, Cloutier S, Lycar L, Radovanovic N, Humphreys D G, Noll J S, Somers D J, Brown P D. Molecular detection of QTLs for agronomic and quality traits in a doubled haploid population derived from two Canadian wheats(Triticum aestivum L.)[J]. Theoretical and Applied Genetics, 2006, 113(4):753-766.doi:10.1007/s00122-006-0346-7. doi: 10.1007/s00122-006-0346-7 pmid: 16838135
[5]	Li Y L, Zhou R H, Wang J, Liao X Z, Branlard G, Jia J Z. Novel and favorable QTL allele clusters for end-use quality revealed by introgression lines derived from synthetic wheat[J]. Molecular Breeding, 2012, 29(3):627-643.doi:10.1007/s11032-011-9578-6. doi: 10.1007/s11032-011-9578-6 URL
[6]	Bajgain P, Sallam A H, Annor G, Conley E, Steffenson B J, Muehlbauer G J, Anderson J A. Genetic characterization of flour quality and bread-making traits in a spring wheat nested association mapping population[J]. Crop Science, 2021, 61(2):1168-1183.doi:10.1002/csc2.20432. doi: 10.1002/csc2.20432 URL
[7]	Xu X, Li X J, Zhang D H, Zhao J S, Jiang X L, Sun H L, Ru Z G. Identification and validation of QTLs for kernel number per spike and spike length in two founder genotypes of wheat[J]. BMC Plant Biology, 2022, 22(1):146.doi:10.1186/s12870-022-03544-6. doi: 10.1186/s12870-022-03544-6 pmid: 35346053
[8]	Chai J F, Zhou R H, Jia J Z, Liu X. Development and application of a new codominant PCR marker for detecting 1BL.1RS wheat-rye chromosome translocations[J]. Plant Breeding, 2006, 125(3):302-304.doi:10.1111/j.1439-0523.2006.01186.x. doi: 10.1111/j.1439-0523.2006.01186.x URL
[9]	丁明亮, 林丽萍, 李明菊, 胡欣, 何迟, 廖合发, 赵红, 李绍祥, 刘琨. 云南育成小麦品种(系)品质性状遗传多样性分析及综合评价[J]. 南方农业学报, 2020, 51(2):255-266.doi:10.3969/j.issn.2095-1191.2020.02.002. doi: 10.3969/j.issn.2095-1191.2020.02.002
	Ding M L, Lin L P, Li M J, Hu X, He C, Liao H F, Zhao H, Li S X, Liu K. Genetic diversity analysis and comprehensive evaluation of quality traits of wheat varieties(lines)bred in Yunnan[J]. Journal of Southern Agriculture, 2020, 51(2):255-266.
[10]	李桂萍, 张根生, 巴青松, 张改生. 杂种小麦品质性状的性状相关和主成分分析[J]. 浙江农业学报, 2016, 28(9):1447-1453.doi:10.3969/j.issn.1004-1524.2016.09.01. doi: 10.3969/j.issn.1004-1524.2016.09.01
	Li G P, Zhang G S, Ba Q S, Zhang G S. Correlation analysis and principal component analysis on quality traits in hybrid wheat[J]. Acta Agriculturae Zhejiangensis, 2016, 28(9):1447-1453.
[11]	王掌军, 刘妍, 李阳, 杨乔乔, 田蕊, 亢玲, 张晓岗, 刘萍, 刘生祥. 宁春4号与河东乌麦杂交F₂品质性状及其分子标记分析[J]. 河南农业科学, 2019, 48(2):17-24.doi:10.15933/j.cnki.1004-3268.2019.02.003.
	Wang Z J, Liu Y, Li Y, Yang Q Q, Tian R, Kang L, Zhang X G, Liu P, Liu S X. Analysis of quality traits and their molecular markers of F₂ hybrids from ningchun No.4 and Hedong black wheat[J]. Journal of Henan Agricultural Sciences, 2019, 48(2):17-24.
[12]	Lou H Y, Zhang R Q, Liu Y T, Guo D D, Zhai S S, Chen A Y, Zhang Y F, Xie C J, You M S, Peng H R, Liang R Q, Ni Z F, Sun Q X, Li B Y. Genome-wide association study of six quality-related traits in common wheat(Triticum aestivum L.)under two sowing conditions[J]. Theoretical and Applied Genetics, 2021, 134(1):399-418.doi:10.1007/s00122-020-03704-y. doi: 10.1007/s00122-020-03704-y
[13]	Krystkowiak K, Langner M, Adamski T, Salmanowicz B P, Kaczmarek Z, Krajewski P, Surma M. Interactions between Glu-1 and Glu-3 loci and associations of selected molecular markers with quality traits in winter wheat(Triticum aestivum L.)DH lines[J]. Journal of Applied Genetics, 2017, 58(1):37-48.doi:10.1007/s13353-016-0362-5. doi: 10.1007/s13353-016-0362-5 pmid: 27502940
[14]	Fatiukha A, Filler N, Lupo I, Lidzbarsky G, Klymiuk V, Korol A B, Pozniak C, Fahima T, Krugman T. Grain protein content and thousand kernel weight QTLs identified in a durum×wild emmer wheat mapping population tested in five environments[J]. Theoretical and Applied Genetics, 2020, 133(1):119-131.doi:10.1007/s00122-019-03444-8. doi: 10.1007/s00122-019-03444-8 pmid: 31562566
[15]	Li Q, Pan Z F, Gao Y, Li T, Liang J J, Zhang Z J, Zhang H L, Deng G B, Long H, Yu M Q. Quantitative trait locus(QTLs)mapping for quality traits of wheat based on high density genetic map combined with bulked segregant analysis RNA-seq(BSR-seq)indicates that the Basic 7S globulin gene is related to falling number[J]. Frontiers in Plant Science, 2020, 11:600788.doi:10.3389/fpls.2020.600788. doi: 10.3389/fpls.2020.600788 URL
[16]	Conti V, Roncallo P F, Beaufort V, Cervigni G L, Miranda R, Jensen C A, Echenique V C. Mapping of main and epistatic effect QTLs associated to grain protein and gluten strength using a RIL population of durum wheat[J]. Journal of Applied Genetics, 2011, 52(3):287-298.doi:10.1007/s13353-011-0045-1. doi: 10.1007/s13353-011-0045-1 pmid: 21523429
[17]	Liu J, Feng B, Xu Z B, Fan X L, Jiang F, Jin X F, Cao J, Wang F, Liu Q, Yang L, Wang T. A genome-wide association study of wheat yield and quality-related traits in southwest China[J]. Molecular Breeding, 2018, 38(1):1.doi:10.1007/s11032-017-0759-9. doi: 10.1007/s11032-017-0759-9 URL
[18]	Cui F, Zhang N, Fan X L, Zhang W, Zhao C H, Yang L J, Pan R Q, Chen M, Han J, Zhao X Q, Ji J, Tong Y P, Zhang H X, Jia J Z, Zhao G Y, Li J M. Utilization of a Wheat660K SNP array-derived high-density genetic map for high-resolution mapping of a major QTL for kernel number[J]. Scientific Reports, 2017, 7(1):3788.doi:10.1038/s41598-017-04028-6. doi: 10.1038/s41598-017-04028-6 pmid: 28630475
[19]	Gao F M, Wen W E, Liu J D, Rasheed A, Yin G H, Xia X C, Wu X X, He Z H. Genome-wide linkage mapping of QTL for yield components,plant height and yield-related physiological traits in the Chinese wheat cross Zhou 8425B/Chinese spring[J]. Frontiers in Plant Science, 2015, 6:1099.doi:10.3389/fpls.2015.01099. doi: 10.3389/fpls.2015.01099
[20]	Jin H, Wen W, Liu J D, Zhai S N, Zhang Y, Yan J, Liu Z Y, Xia X C, He Z H. Genome-wide QTL mapping for wheat processing quality parameters in a Gaocheng 8901/Zhoumai 16 recombinant inbred line population[J]. Frontiers in Plant Science, 2016, 7:1032.doi:10.3389/fpls.2016.01032. doi: 10.3389/fpls.2016.01032 pmid: 27486464
[21]	王美芳, 雷振生, 何中虎, 孙其信, 吴政卿, 何盛莲, 杨攀. 冬小麦品种的HMW-GS组成和1BL/1RS易位分布及其与品质性状的关系研究[J]. 华北农学报, 2008, 23(5):62-66.doi:10.7668/hbnxb.2008.05.014. doi: 10.7668/hbnxb.2008.05.014
	Wang M F, Lei Z S, He Z H, Sun Q X, Wu Z Q, He S L, Yang P. The composition of HMW-GS and the frequency of 1BL/1RS translocation and their relationship to quality in winter wheat varieties in China[J]. Acta Agriculturae Boreali-Sinica, 2008, 23(5):62-66.
[22]	程敦公, 李豪圣, 肖永贵, 宋健民, 刘爱峰, 刘建军. 快速准确鉴定小麦1BL/1RS易位系新方法的研究[J]. 华北农学报, 2009, 24(S1):34-37.doi:10.7668/hbnxb.2009.S1.009. doi: 10.7668/hbnxb.2009.S1.009
	Cheng D G, Li H S, Xiao Y G, Song J M, Liu A F, Liu J J. New method to identify 1BL/1RS translocation in common wheat[J]. Acta Agriculturae Boreali-Sinica, 2009, 24(S1):34-37.
[23]	Lee J H, Graybosch R A, Peterson C J. Quality and biochemical effects of a IBL/IRS wheat-rye translocation in wheat[J]. Theoretical and Applied Genetics, 1995, 90(1):105-112.doi:10.1007/BF00221002. doi: 10.1007/BF00221002 pmid: 24173790
[24]	Gobaa S, Brabant C, Kleijer G, Stamp P. Effect of the 1BL.1RS translocation and of the Glu-B3 variation on fifteen quality tests in a doubled haploid population of wheat(Triticum aestivum L.)[J]. Journal of Cereal Science, 2008, 48(3):598-603.doi:10.1016/j.jcs.2007.12.006. doi: 10.1016/j.jcs.2007.12.006 URL
[25]	Ji J, Wang Z G, Sun J Z, Li J M, Zhang X Q, Wang D W, Zhang A M. Identification of new T1BL/1RS translocation lines derived from wheat(Triticum aestivum L.cultivar Xiaoyan No.6)and rye hybridization[J]. Acta Physiologiae Plantarum, 2008, 30(5):689-695.doi:10.1007/s11738-008-0167-1. doi: 10.1007/s11738-008-0167-1 URL
[26]	肖永贵, 阎俊, 何中虎, 张勇, 张晓科, 刘丽, 李天富, 曲延英, 夏先春. 1BL/1RS易位对小麦产量性状和白粉病抗性的影响及其QTL分析[J]. 作物学报, 2006, 32(11):1636-1641.doi:10.3321/j.issn:0496-3490.2006.11.007.
	Xiao Y G, Yan J, He Z H, Zhang Y, Zhang X K, Liu L, Li T F, Qu Y Y, Xia X C. Effect of 1BL/1RS translocation on yield traits and powdery mildew resistance in common wheat and QTL analysis[J]. Acta Agronomica Sinica, 2006, 32(11):1636-1641.
[27]	Lelley T, Eder C, Grausgruber H. Influence of 1BL.1RS wheat-rye chromosome translocation on genotype by environment interaction[J]. Journal of Cereal Science, 2004, 39(3):313-320.doi:10.1016/j.jcs.2003.11.003. doi: 10.1016/j.jcs.2003.11.003 URL

性状 Trait	环境 Environment	亲本Parent		重组自交系RIL
性状 Trait	环境 Environment	百农AK58 Bainong AK58	碧蚂4号 Bima 4	平均值 Average	标准差 s	变异范围 Range	变异系数/% CV
蛋白质含量/%	E1	14.39	14.79	15.18	1.13	12.13~19.04	7.42
GPC	E2	14.07	—	15.65	1.75	11.07~20.68	11.21
	E3	14.50	14.05	14.87	1.18	12.45~18.39	7.93
	E4	13.94	15.49	14.18	1.27	11.41~17.85	8.98
	EA	14.23	14.78	14.96	1.08	12.02~18.32	7.22
湿面筋含量/%	E1	31.50	32.35	32.87	2.59	25.95~41.86	7.87
WGC	E2	30.60	—	33.65	4.01	23.79~44.82	11.91
	E3	30.93	29.84	31.79	2.71	26.09~40.38	8.54
	E4	28.90	33.01	29.69	3.00	22.66~38.24	10.10
	EA	30.48	31.73	31.96	2.48	25.44~40.04	7.76
淀粉含量/%	E1	19.90	33.81	27.97	4.68	17.19~50.05	16.73
SC	E2	22.19	—	30.65	6.44	15.48~63.12	21.00
	E3	23.91	40.47	27.93	5.86	14.37~74.28	20.97
	E4	22.25	35.53	30.98	9.25	12.64~61.85	29.87
	EA	22.06	36.60	29.41	4.61	17.05~47.35	15.67
沉降值/mL	E1	27.20	31.83	31.09	3.85	21.97~45.41	12.37
SV	E2	27.23	—	32.79	5.91	17.36~50.62	18.03
	E3	28.57	31.82	29.64	4.42	20.72~51.69	14.90
	E4	26.54	34.78	30.53	5.87	15.30~48.19	19.22
	EA	27.38	32.81	31.02	3.79	20.74~42.71	12.23
延伸性/mm	E1	130.04	153.05	149.88	10.74	117.34~189.20	7.16
EX	E2	130.69	—	157.30	17.56	110.20~206.86	11.16
	E3	141.26	155.17	154.72	11.15	125.16~189.30	7.20
	E4	131.52	157.51	142.67	12.94	114.66~185.96	9.07
	EA	133.38	155.24	151.20	10.33	124.78~184.95	6.83

性状 Trait	环境 Environment	亲本Parent		重组自交系RIL
性状 Trait	环境 Environment	百农AK58 Bainong AK58	碧蚂4号 Bima 4	平均值 Average	标准差 s	变异范围 Range	变异系数/% CV
蛋白质含量/%	E1	14.39	14.79	15.18	1.13	12.13~19.04	7.42
GPC	E2	14.07	—	15.65	1.75	11.07~20.68	11.21
	E3	14.50	14.05	14.87	1.18	12.45~18.39	7.93
	E4	13.94	15.49	14.18	1.27	11.41~17.85	8.98
	EA	14.23	14.78	14.96	1.08	12.02~18.32	7.22
湿面筋含量/%	E1	31.50	32.35	32.87	2.59	25.95~41.86	7.87
WGC	E2	30.60	—	33.65	4.01	23.79~44.82	11.91
	E3	30.93	29.84	31.79	2.71	26.09~40.38	8.54
	E4	28.90	33.01	29.69	3.00	22.66~38.24	10.10
	EA	30.48	31.73	31.96	2.48	25.44~40.04	7.76
淀粉含量/%	E1	19.90	33.81	27.97	4.68	17.19~50.05	16.73
SC	E2	22.19	—	30.65	6.44	15.48~63.12	21.00
	E3	23.91	40.47	27.93	5.86	14.37~74.28	20.97
	E4	22.25	35.53	30.98	9.25	12.64~61.85	29.87
	EA	22.06	36.60	29.41	4.61	17.05~47.35	15.67
沉降值/mL	E1	27.20	31.83	31.09	3.85	21.97~45.41	12.37
SV	E2	27.23	—	32.79	5.91	17.36~50.62	18.03
	E3	28.57	31.82	29.64	4.42	20.72~51.69	14.90
	E4	26.54	34.78	30.53	5.87	15.30~48.19	19.22
	EA	27.38	32.81	31.02	3.79	20.74~42.71	12.23
延伸性/mm	E1	130.04	153.05	149.88	10.74	117.34~189.20	7.16
EX	E2	130.69	—	157.30	17.56	110.20~206.86	11.16
	E3	141.26	155.17	154.72	11.15	125.16~189.30	7.20
	E4	131.52	157.51	142.67	12.94	114.66~185.96	9.07
	EA	133.38	155.24	151.20	10.33	124.78~184.95	6.83

性状 Trait	蛋白质含量 GPC	湿面筋含量 WGC	淀粉含量 SC	沉降值 SV	延伸性 EX
蛋白质含量 GPC	1.00
湿面筋含量 WGC	0.99^***	1.00
淀粉含量 SC	0.45^***	0.42^***	1.00
沉降值 SV	0.90^***	0.87^***	0.76^***	1.00
延伸性 EX	0.89^***	0.87^***	0.67^***	0.88^***	1.00

性状 Trait	蛋白质含量 GPC	湿面筋含量 WGC	淀粉含量 SC	沉降值 SV	延伸性 EX
蛋白质含量 GPC	1.00
湿面筋含量 WGC	0.99^***	1.00
淀粉含量 SC	0.45^***	0.42^***	1.00
沉降值 SV	0.90^***	0.87^***	0.76^***	1.00
延伸性 EX	0.89^***	0.87^***	0.67^***	0.88^***	1.00

性状 Trait		QTL	环境 Environment	染色体 Chromosome	位置/cM Position	侧翼标记 Flanking markers	LOD值 LOD value	表型贡献率/% PVE	加性效应 Additive effect
蛋白质含量		QGpc.his-1B	E4	1B	6	AX-110000934~AX-109892648	2.67	3.21	0.25
GPC		QGpc.his-1D	EA	1D	400	AX-111589460~AX-111515122	3.75	5.16	-0.25
		Qgpc.his-2B	E3	2B	98	AX-111040705~AX-110120266	3.00	3.07	0.22
			EA	2B	105	AX-109352812~AX-111639840	3.88	5.06	0.25
		QGpc.his-2D	E1	2D	63	AX-109414589~AX-110051842	3.08	4.97	0.24
		QGpc.his-3A	E1	3A	14	AX-111129296~AX-110587728	2.61	4.07	0.22
			E3	3A	12	AX-108787734~AX-111070416	4.02	4.28	0.26
		QGpc.his-3B	E1	3B	56	AX-111239152~AX-109857556	7.29	12.21	0.39
		QGpc.his-3D	E1	3D	141	AX-109306840~AX-110827948	2.73	4.37	-0.23
		QGpc.his-4A-1	E1	4A	61	AX-111170340~AX-109931552	2.86	4.46	-0.24
			E4	4A	54	AX-109968652~AX-109821994	3.56	4.11	-0.28
		QGpc.his-4A-2	E1	4A	121	AX-111170720~AX-110675502	4.88	7.79	0.32
			E3	4A	120	AX-111170720~AX-110675502	10.93	12.56	0.45
			E4	4A	117	AX-110649064~AX-111170720	3.33	3.80	0.27
			EA	4A	120	AX-111170720~AX-110675502	6.58	9.21	0.35
		QGpc.his-4B	E3	4B	98	AX-89708030~AX-108793833	3.19	3.78	-0.24
		QGpc.his-4D	EA	4D	66	AX-110476142~AX-111002463	2.99	3.87	-0.22
		QGpc.his-5A	E4	5A	119	AX-110954181~AX-111253877	5.19	6.15	-0.34
		QGpc.his-6D	E2	6D	4	AX-108745027~AX-109374059	3.33	6.99	0.45
		QGpc.his-7A-1	E3	7A	26	AX-108947190~AX-111659056	3.99	4.25	-0.26
		QGpc.his-7A-2	E4	7A	201	AX-111488311~AX-110973510	3.17	3.71	0.26
		QGpc.his-7B	E4	7B	150	AX-110470708~AX-108877192	2.86	3.33	0.25
		QGpc.his-7D	E3	7D	616	AX-89713293~AX-110611417	6.30	6.71	-0.32
湿面筋含量		QWgc.his-1A	E2	1A	76	AX-110379520~AX-109338114	4.47	5.72	1.03
WGC		QWgc.his-1B	E2	1B	35	AX-111548801~AX-109509313	3.68	4.67	0.94
			E4	1B	29	AX-109397003~AX-111056129	2.68	5.13	0.66
		QWgc.his-1D	EA	1D	400	AX-111589460~AX-111515122	4.19	6.11	-0.60
		QWgc.his-2B	E2	2B	128	AX-108945173~AX-108796024	3.32	4.30	0.91
			EA	2B	105	AX-109352812~AX-111639840	4.59	6.37	0.61
		QWgc.his-2D	E1	2D	64	AX-109414589~AX-110051842	2.88	4.62	0.54
			E3	2D	81	AX-94506151~AX-110068988	3.57	3.79	0.53
		QWgc.his-3A	E1	3A	13	AX-111070416~AX-111129296	2.65	4.32	0.52
			E3	3A	12	AX-108787734~AX-111070416	3.49	3.81	0.54
性状 Trait	QTL		环境 Environment	染色体 Chromosome	位置/cM Position	侧翼标记 Flanking markers	LOD值 LOD value	表型贡献率/% PVE	加性效应 Additive effect
	QWgc.his-3B		E1	3B	56	AX-111239152~AX-109857556	6.51	10.94	0.84
	QWgc.his-3D		E1	3D	141	AX-109306840~AX-110827948	2.66	4.33	-0.52
			E4	3D	141	AX-109306840~AX-110827948	2.64	5.11	-0.66
	QWgc.his-4A-1		E1	4A	62	AX-109474732~AX-111584564	3.07	4.89	-0.57
	QWgc.his-4A-2		E1	4A	121	AX-111170720~AX-110675502	4.78	7.71	0.72
			E3	4A	122	AX-110675502~AX-109332913	12.82	15.94	1.13
			EA	4A	120	AX-111170720~AX-110675502	6.05	8.83	0.74
	QWgc.his-4D		EA	4D	66	AX-110476142~AX-111002463	2.84	3.89	-0.49
	QWgc.his-5A		E2	5A	98	AX-110437938~AX-109817972	4.58	6.00	-1.09
	QWgc.his-5D		E3	5D	646	AX-108974123~AX-89331886	2.52	3.19	-0.49
	QWgc.his-6D		E2	6D	4	AX-108745027~AX-109374059	3.18	4.09	0.88
	QWgc.his-7A		E3	7A	26	AX-108947190~AX-111659056	2.83	3.09	-0.48
	QWgc.his-7D		E3	7D	616	AX-89713293~AX-110611417	5.82	6.43	-0.70
			EA	7D	616	AX-89713293~AX-110611417	2.71	3.74	-0.46
淀粉含量	QSc.his-1B		E3	1B	86	AX-109822203~AX-111760595	2.54	4.79	-1.30
SC	QSc.his-2A-1		E2	2A	39	AX-109321357~AX-111030654	3.51	7.55	-1.66
	QSc.his-2A-2		E4	2A	91	AX-109303703~AX-86179766	3.31	4.38	-2.13
	QSc.his-2D		E1	2D	415	AX-111193092~AX-109464511	2.83	11.14	-1.90
	QSc.his-3D		E4	3D	89	AX-110543919~AX-109417352	2.67	4.77	-2.22
	QSc.his-5A		E1	5A	61	AX-109524688~AX-110950060	3.34	3.65	1.09
	QSc.his-6A		E4	6A	42	AX-111134280~AX-109370898	2.84	3.63	1.94
	QSc.his-6D-1		E2	6D	4	AX-108745027~AX-109374059	3.37	7.30	1.63
	QSc.his-6D-2		E1	6D	119	AX-108837043~AX-108793828	2.86	3.24	1.03
沉降值	QSv.his-1D		EA	1D	400	AX-111589460~AX-111515122	3.34	4.24	-0.87
SV	QSv.his-2D		E3	2D	554	AX-109417243~AX-108976867	2.84	4.45	-0.90
	QSv.his-2B		E1	3B	56	AX-111239152~AX-109857556	8.07	14.01	1.44
	QSv.his-3D		E3	3D	143	AX-111155780~AX-110365829	2.78	4.35	-0.88
			E4	3D	104	AX-110495749~AX-109799345	3.36	4.69	-1.39
			EA	3D	130	AX-108738283~AX-111027124	2.64	4.37	-0.87
	QSv.his-4A		E1	4A	121	AX-111170720~AX-110675502	3.39	5.51	0.92
			E3	4A	121	AX-111170720~AX-110675502	4.74	7.60	1.20
			EA	4A	118	AX-111170720~AX-110675502	4.93	6.25	1.07
	QSv.his-5A-1		E2	5A	56	AX-110641373~AX-109994159	27.31	14.53	4.56
			EA	5A	56	AX-110641373~AX-109994159	6.17	7.65	1.16
	QSv.his-5A-2		E2	5A	98	AX-110437938~AX-109817972	2.82	1.19	-1.34
			EA	5A	114	AX-111514440~AX-109494596	2.86	3.42	-0.78
	QSv.his-5A-3		E4	5A	194	AX-111038387~AX-110449012	3.29	4.61	-1.38
	QSv.his-6D-1		E2	6D	4	AX-108745027~AX-109374059	4.25	1.78	1.59
			EA	6D	4	AX-108745027~AX-109374059	2.93	3.51	0.78
	QSv.his-6D-2		E1	6D	117	AX-108837043~AX-108793828	3.89	6.46	0.97
延伸性	QEx.his-3A		E1	3A	10	AX-110038636~AX-108787734	3.56	5.65	2.72
EX	QEx.his-3B		E1	3B	45	AX-109914961~AX-108776238	2.88	4.51	2.46
	QEx.his-3D		E4	3D	138	AX-108795505~AX-110489977	2.69	4.58	-2.84
			EA	3D	141	AX-109306840~AX-110827948	2.86	3.62	-2.08
	QEx.his-4A		E1	4A	123	AX-110675502~AX-109332913	4.00	7.47	3.22
			E3	4A	121	AX-111170720~AX-110675502	8.60	11.57	4.29
			EA	4A	120	AX-111170720~AX-110675502	8.33	11.66	3.85
性状 Trait	QTL		环境 Environment	染色体 Chromosome	位置/cM Position	侧翼标记 Flanking markers	LOD值 LOD value	表型贡献率/% PVE	加性效应 Additive effect
	QEx.his-5A		E2	5A	56	AX-110641373~AX-109994159	23.62	12.78	12.60
			EA	5A	56	AX-110641373~AX-109994159	4.26	5.41	2.55
	QEx.his-5B		E3	5B	41	AX-109856154~AX-108769780	3.85	4.93	2.71
			EA	5B	41	AX-109856154~AX-108769780	3.91	4.89	2.42
	QEx.his-5D		E1	5D	643	AX-108974123~AX-89331886	5.04	8.67	-3.37
	QEx.his-6D		E2	6D	4	AX-108745027~AX-109374059	4.43	1.96	4.92
			EA	6D	4	AX-108745027~AX-109374059	2.83	3.57	2.06

选择文件类型/文献管理软件名称

选择包含的内容