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作物学报

作物学报 ›› 2025, Vol. 51 ›› Issue (6): 1558-1568.doi: 10.3724/SP.J.1006.2025.44166

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

大豆开花时间和成熟期性状全基因组关联分析与候选基因预测

王琼1(), 邹丹霞1, 陈兴运1, 张威1, 张红梅1, 刘晓庆1, 贾倩茹1, 魏利斌2, 崔晓艳1, 陈新1, 王学军2,*(), 陈华涛1,*()   

  1. 1江苏省农业科学院经济作物研究所, 江苏南京 210014
    2江苏沿江地区农业科学研究所, 江苏南通 226541
  • 收稿日期:2024-09-29 接受日期:2025-03-26 出版日期:2025-06-12 网络出版日期:2025-04-07
  • 通讯作者: *陈华涛, E-mail: cht@jaas.ac.cn;王学军, E-mail: wangxj4002@sina.com
  • 作者简介:E-mail: wqwzw@163.com
  • 基金资助:
    本研究由江苏省自然科学基金项目(BK20220740);江苏省重点研发计划项目(BE2022328);江苏省种业振兴揭榜挂帅项目(JBGS [2021] 057)

Genome-wide association analysis and candidate genes prediction of flowering time and maturity date traits in soybean (Glycine max L.)

WANG Qiong1(), ZOU Dan-Xia1, CHEN Xing-Yun1, ZHANG Wei1, ZHANG Hong-Mei1, LIU Xiao-Qing1, JIA Qian-Ru1, WEI Li-Bin2, CUI Xiao-Yan1, CHEN Xin1, WANG Xue-Jun2,*(), CHEN Hua-Tao1,*()   

  1. 1Institute of Industrial Crops, Jiangsu Academy of Agricultural Sciences, Nanjing 210014, Jiangsu, China
    2Jiangsu Yanjiang Institute of Agricultural Sciences, Nantong 226541, Jiangsu, China
  • Received:2024-09-29 Accepted:2025-03-26 Published:2025-06-12 Published online:2025-04-07
  • Contact: *E-mail: cht@jaas.ac.cn;E-mail: wangxj4002@sina.com
  • Supported by:
    This study was supported by the Natural Science Foundation of Jiangsu Province(BK20220740);the Key Research and Development Program ofJiangsu Province(BE2022328);the Jiangsu Seed Industry Revitalization “Unveiling and Commanding” Project(JBGS [2021] 057)

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摘要：

大豆是典型的短日照作物, 对光周期极为敏感, 其栽培和产量均受到田间光周期条件的制约。本研究对264份大豆种质资源的开花时间和成熟期性状进行了考察, 分析了开花相关性状与蛋白质含量、含油量、百粒重和株高等农艺性状之间的关系。随后利用全基因组关联分析, 鉴定出235个与开花时间和成熟期相关的位点, 并预测了14个可能参与调控大豆开花时间和成熟期的候选基因, 其中与开花时间相关的候选基因有10个, 与成熟期相关的基因有5个, 且有1个候选基因同时与开花时间和成熟期相关。这些候选基因为进一步解析大豆开花相关性状的调控机制和大豆广适性高效遗传改良奠定了基础。

关键词: 大豆, 开花时间, 成熟期, 全基因组关联分析, SNP标记

Abstract:

Soybean is a typical short-day crop that is highly sensitive to photoperiod, with its cultivation and yield constrained by field photoperiodic conditions. In this study, we analyzed the flowering time and pod maturity date in 264 diverse soybean accessions. We examined the relationships between flowering-related traits and key agronomic traits, including protein content (PC), oil content (OC), 100-seed weight (HSW), and plant height (PH). A genome-wide association study (GWAS) identified 235 loci associated with flowering time and pod maturity date. Additionally, we predicted 14 candidate genes involved in the regulation of these traits, including 10 genes related to flowering time and 5 genes associated with pod maturity date. Notably, one gene exhibited pleiotropic effects on both traits. These findings provide valuable genomic insights into the regulatory pathways of flowering in soybean and offer a foundation for genetic improvement aimed at enhancing soybean adaptation across broader latitudinal regions.

Key words: soybean, flowering time, maturity date, GWAS, SNP markers

图1

大豆开花相关性状的频率分布及表型相关性分析 (a): 所有大豆材料开花时间的分布; (b): 所有大豆材料成熟期的分布; (c): 开花性状间的相关系数; (d): 开花性状与含油量(OC)、蛋白质含量(PC)、百粒重(HSW)和株高(PH)的相关系数。*、**和***分别表示在0.05、0.01和0.001水平上相关性显著。NJ19: 2019年南京; NJ21: 2021年南京; NJ22: 2022年南京。"

附图1

开花时间和成熟期性状关联分析曼哈顿图 (a)~(d): BLUP、NJ19、NJ21和NJ22环境下开花时间关联分析的曼哈顿图; (e)~(h): BLUP、NJ19、NJ21和NJ22环境下成熟期关联分析的曼哈顿图。箭头表示已知开花基因的位置。20条染色体用不同的颜色表示。水平实线表示?log10(P)≥5的阈值，高于该阈值的位点用橙色圆点表示，水平虚线表示GEC软件计算的显著性阈值，高于该阈值用红色圆点表示。"

附图2

开花时间和成熟期性状关联分析QQ图 (a)~(d): BLUP、NJ19、NJ21和NJ22环境下开花时间关联分析的QQ图; (e)~(h): BLUP、NJ19、NJ21和NJ22环境下成熟期关联分析的QQ图。"

表S1

本研究鉴定的大豆开花时间和成熟期相关位点以及与已知基因的重叠信息"

关联信号
Signal		 最显著位点
Peak loci		 染色体
Chromosome		 位置
Position (bp)		 P值
P		 环境
Environment		 性状
Trait		 基因名称
Gene		 基因编号
Gene ID		 开始
Start (bp)		 结束
End (bp)
1_16.8 Chr.01: 16756203 01 16,756,203 2.82E-06 NJ19 开花时间Flowering time
1_19.8 Chr.01: 19797880 01 19,797,880 9.72E-06 NJ22 开花时间Flowering time
1_50.1 Chr.01: 50113489 01 50,113,489 6.79E-06 NJ22 开花时间 Flowering time
2_0.1 Chr.02: 90922 02 90,922 7.54E-06 NJ22 开花时间 Flowering time
2_0.2 Chr.02: 198739 02 198,739 9.87E-06 NJ21 开花时间 Flowering time
2_11.6 Chr.02: 11628488 02 11,628,488 5.55E-06 NJ19 开花时间 Flowering time AP1d Glyma.02G121600 12,084,714 120,890,43
2_13.6 Chr.02: 13554490 02 13,554,490 6.40E-06 NJ19 开花时间 Flowering time
2_30 Chr.02: 29989462 02 29,989,462 1.67E-06 NJ21 开花时间 Flowering time
2_39.4 Chr.02: 39385406 02 39,385,406 3.66E-06 NJ19 开花时间 Flowering time
3_1.1 Chr.03: 1131854 03 1,131,854 1.63E-06 NJ21 开花时间 Flowering time
3_7 Chr.03: 6956708 03 6,956,708 4.13E-06 NJ21 开花时间 Flowering time
3_11.4 Chr.03: 11386468 03 11,386,468 5.14E-06 NJ21 开花时间 Flowering time
3_17 Chr.03: 16963162 03 16,963,162 6.30E-06 NJ21 开花时间 Flowering time
3_33.2 Chr.03: 33154866 03 33,154,866 8.19E-06 NJ19 开花时间 Flowering time
3_33.6 Chr.03: 33551605 03 33,551,605 2.47E-07 NJ21 开花时间 Flowering time
3_40.6 Chr.03: 40635793 03 40,635,793 5.59E-06 NJ19 开花时间 Flowering time
3_40.6 Chr.03: 40635793 03 40,635,793 8.87E-06 NJ21 开花时间 Flowering time
3_43.3 Chr.03: 43294123 03 43,294,123 7.10E-07 NJ19 开花时间 Flowering time
4_24.1 Chr.04: 24085510 04 24,085,510 9.73E-06 NJ21 开花时间 Flowering time
5_0.1 Chr.05: 66789 05 66,789 5.31E-06 NJ21 开花时间 Flowering time
5_17.8 Chr.05: 17804685 05 17,804,685 8.57E-06 NJ22 开花时间 Flowering time
5_25.8 Chr.05: 25798559 05 25,798,559 7.67E-06 NJ21 开花时间 Flowering time
6_11.7 Chr.06: 11735349 06 11,735,349 2.02E-06 NJ21 开花时间 Flowering time
6_11.8 Chr.06: 11765729 06 11,765,729 6.82E-06 NJ21 开花时间 Flowering time
6_17.1 Chr.06: 17072269 06 17,072,269 1.11E-06 NJ22 开花时间 Flowering time
6_18.9 Chr.06: 18915994 06 18,915,994 5.23E-06 NJ22 开花时间 Flowering time
6_19 Chr.06: 19018252 06 19,018,252 5.71E-06 NJ22 开花时间 Flowering time QNE1 Glyma.06G204300 19,210,586 19,213,448
6_20.7 Chr.06: 20734601 06 20,734,601 4.66E-06 NJ21 开花时间 Flowering time E1 Glyma.06G207800 20,207,077 20,207,940
6_42.7 Chr.06: 42698406 06 42,698,406 2.05E-06 NJ21 开花时间 Flowering time
6_43 Chr.06: 43014590 06 43,014,590 9.48E-06 NJ21 开花时间 Flowering time
6_45.5 Chr.06: 45525901 06 45,525,901 7.14E-06 NJ22 开花时间 Flowering time
6_51.3 Chr.06: 51347716 06 51,347,716 6.05E-06 NJ21 开花时间 Flowering time
7_9.1 Chr.07: 9139724 07 9,139,724 2.93E-06 NJ21 开花时间 Flowering time
7_9.3 Chr.07: 9281382 07 9,281,382 1.16E-06 NJ21 开花时间 Flowering time
7_9.4 Chr.07: 9412365 07 9,412,365 6.75E-06 NJ21 开花时间 Flowering time
9_16.7 Chr.09: 16690457 09 16,690,457 1.45E-06 NJ22 开花时间 Flowering time
9_33.5 Chr.09: 33543971 09 33,543,971 4.58E-06 BLUP 开花时间 Flowering time
10_13.9 Chr.10: 13877946 10 13,877,946 8.76E-06 NJ19 开花时间 Flowering time
10_14.1 Chr.10: 14063324 10 14,063,324 2.22E-06 NJ19 开花时间 Flowering time
10_14.4 Chr.10: 14393613 10 14,393,613 1.59E-06 NJ19 开花时间 Flowering time
10_18.5 Chr.10: 18549131 10 18,549,131 9.70E-06 NJ19 开花时间 Flowering time
10_29.1 Chr.10: 29145960 10 29,145,960 3.23E-06 NJ19 开花时间 Flowering time
10_44.9 Chr.10: 44948729 10 44,948,729 1.75E-06 NJ21 开花时间 Flowering time
10_45 Chr.10: 45045188 10 45,045,188 7.64E-07 NJ21 开花时间 Flowering time GI Glyma.10G221500 45,294,735 45,316,121
11_6 Chr.11: 6010522 11 6,010,522 7.50E-07 NJ22 开花时间 Flowering time
11_24.4 Chr.11: 24387887 11 24,387,887 7.46E-07 NJ21 开花时间 Flowering time
11_26.5 Chr.11: 26451428 11 26,451,428 2.39E-06 NJ21 开花时间 Flowering time
13_41.8 Chr.13: 41789036 13 41,789,036 8.54E-06 BLUP 开花时间 Flowering time
13_41.9 Chr.13: 41873651 13 41,873,651 9.06E-07 BLUP 开花时间 Flowering time
14_4.9 Chr.14: 4908776 14 4,908,776 1.69E-06 NJ21 开花时间 Flowering time
14_30.3 Chr.14: 30334422 14 30,334,422 9.52E-06 NJ21 开花时间 Flowering time
14_48.4 Chr.14: 48432986 14 48,432,986 6.47E-06 BLUP 开花时间 Flowering time
14_48.4 Chr.14: 48432986 14 48,432,986 3.56E-06 NJ19 开花时间 Flowering time
15_1.6 Chr.15: 1649953 15 1,649,953 2.83E-06 BLUP 开花时间 Flowering time
15_1.7 Chr.15: 1650236 15 1,650,236 9.40E-06 BLUP 开花时间 Flowering time
15_17.9 Chr.15: 17862246 15 17,862,246 3.73E-06 BLUP 开花时间 Flowering time
15_17.9 Chr.15: 17862246 15 17,862,246 4.67E-07 NJ22 开花时间 Flowering time
15_37.6 Chr.15: 37562128 15 37,562,128 7.23E-06 NJ21 开花时间 Flowering time
15_39.5 Chr.15: 39496561 15 39,496,561 1.71E-07 NJ19 开花时间 Flowering time
16_4.6 Chr.16: 4620056 16 4,620,056 2.95E-06 BLUP 开花时间 Flowering time FT5a Glyma.16G044100 4,135,885 4,137,742
16_5.3 Chr.16: 5275907 16 5,275,907 1.13E-06 BLUP 开花时间 Flowering time
16_5.3 Chr.16: 5275907 16 5,275,907 7.70E-06 NJ21 开花时间 Flowering time
16_5.4 Chr.16: 5357573 16 5,357,573 3.15E-06 BLUP 开花时间 Flowering time
16_5.4 Chr.16: 5363834 16 5,363,834 1.05E-06 NJ21 开花时间 Flowering time
16_5.7 Chr.16: 5719462 16 5,719,462 7.64E-06 NJ21 开花时间 Flowering time
16_5.7 Chr.16: 5748536 16 5,748,536 2.87E-06 BLUP 开花时间 Flowering time
16_22.2 Chr.16: 22180805 16 22,180,805 8.54E-06 BLUP 开花时间 Flowering time
16_24.9 Chr.16: 24918284 16 24,918,284 3.86E-06 NJ21 开花时间 Flowering time
16_29.8 Chr.16: 29817877 16 29,817,877 1.47E-06 NJ21 开花时间 Flowering time
16_32.5 Chr.16: 32485435 16 32,485,435 4.40E-06 BLUP 开花时间 Flowering time
17_6.6 Chr.17: 6600113 17 6,600,113 1.78E-06 NJ21 开花时间 Flowering time MDE17 Glyma.17G081200 6,312,563 6,321,583
17_15.3 Chr.17: 15293172 17 15,293,172 1.01E-07 NJ21 开花时间 Flowering time
17_15.4 Chr.17: 15392244 17 15,392,244 7.11E-07 NJ21 开花时间 Flowering time
17_15.5 Chr.17: 15476791 17 15,476,791 2.01E-06 NJ21 开花时间 Flowering time
17_15.6 Chr.17: 15628996 17 15,628,996 3.97E-06 NJ21 开花时间 Flowering time
17_15.7 Chr.17: 15716574 17 15,716,574 1.18E-07 NJ21 开花时间 Flowering time
17_15.8 Chr.17: 15819838 17 15,819,838 8.27E-08 NJ21 开花时间 Flowering time
17_15.9 Chr.17: 15859190 17 15,859,190 7.81E-08 NJ21 开花时间 Flowering time
17_16.1 Chr.17: 16053819 17 16,053,819 6.26E-08 NJ21 开花时间 Flowering time
17_16.2 Chr.17: 16182668 17 16,182,668 1.67E-06 NJ21 开花时间 Flowering time
17_16.3 Chr.17: 16341492 17 16,341,492 1.51E-06 NJ21 开花时间 Flowering time
17_16.4 Chr.17: 16444519 17 16,444,519 1.28E-06 NJ21 开花时间 Flowering time
17_16.5 Chr.17: 16526522 17 16,526,522 6.02E-07 NJ21 开花时间 Flowering time
17_16.6 Chr.17: 16631273 17 16,631,273 6.19E-07 NJ21 开花时间 Flowering time
17_16.7 Chr.17: 16680555 17 16,680,555 1.28E-06 NJ21 开花时间 Flowering time
17_16.8 Chr.17: 16814325 17 16,814,325 4.77E-06 NJ21 开花时间 Flowering time
17_16.9 Chr.17: 16931162 17 16,931,162 1.38E-06 NJ21 开花时间 Flowering time
17_17 Chr.17: 17028485 17 17,028,485 1.28E-06 NJ21 开花时间 Flowering time
17_17.2 Chr.17: 17158996 17 17,158,996 4.90E-06 NJ21 开花时间 Flowering time
17_17.3 Chr.17: 17292001 17 17,292,001 1.28E-06 NJ21 开花时间 Flowering time
17_17.4 Chr.17: 17352058 17 17,352,058 1.28E-06 NJ21 开花时间 Flowering time
17_17.5 Chr.17: 17494501 17 17,494,501 1.38E-06 NJ21 开花时间 Flowering time
17_17.6 Chr.17: 17584756 17 17,584,756 1.38E-06 NJ21 开花时间 Flowering time
17_17.7 Chr.17: 17683919 17 17,683,919 1.28E-06 NJ21 开花时间 Flowering time
17_18 Chr.17: 17963083 17 17,963,083 1.67E-06 NJ21 开花时间 Flowering time
18_0.7 Chr.18: 687088 18 687,088 1.23E-06 NJ21 开花时间 Flowering time
18_0.8 Chr.18: 841400 18 841,400 5.18E-07 NJ21 开花时间 Flowering time
18_0.8 Chr.18: 843464 18 843,464 4.83E-07 NJ19 开花时间 Flowering time
18_0.9 Chr.18: 861675 18 861,675 4.13E-07 NJ19 开花时间 Flowering time
18_0.9 Chr.18: 879311 18 879,311 2.63E-06 NJ21 开花时间 Flowering time
18_1.6 Chr.18: 1628084 18 1,628,084 1.65E-06 NJ21 开花时间 Flowering time
18_1.7 Chr.18: 1707431 18 1,707,431 3.47E-06 NJ21 开花时间 Flowering time
18_1.8 Chr.18: 1844386 18 1,844,386 8.14E-06 NJ21 开花时间 Flowering time
18_1.9 Chr.18: 1925015 18 1,925,015 4.75E-07 NJ21 开花时间 Flowering time
18_2 Chr.18: 2029511 18 2,029,511 9.73E-07 NJ21 开花时间 Flowering time
18_36.4 Chr.18: 36385829 18 36,385,829 4.91E-06 NJ21 开花时间 Flowering time
18_50.3 Chr.18: 50275974 18 50,275,974 5.27E-06 NJ19 开花时间 Flowering time
18_51.6 Chr.18: 51622787 18 51,622,787 4.53E-06 NJ21 开花时间 Flowering time SOC1a Glyma.18G224500 51,203,136 51,222,128
18_52.7 Chr.18: 52659743 18 52,659,743 4.41E-07 NJ21 开花时间 Flowering time
18_52.9 Chr.18: 52944567 18 52,944,567 9.39E-06 BLUP 开花时间 Flowering time
18_53.7 Chr.18: 53676491 18 53,676,491 3.36E-06 NJ21 开花时间 Flowering time
18_53.8 Chr.18: 53808169 18 53,808,169 9.33E-06 NJ21 开花时间 Flowering time
19_18.1 Chr.19: 18069101 19 18,069,101 4.91E-06 NJ22 开花时间 Flowering time
19_44.9 Chr.19: 44925371 19 44,925,371 8.80E-06 NJ22 开花时间 Flowering time
19_45.4 Chr.19: 45368289 19 45,368,289 1.86E-06 NJ22 开花时间 Flowering time
19_45.5 Chr.19: 45456935 19 45,456,935 2.71E-06 NJ22 开花时间 Flowering time
20_15.1 Chr.20: 15061974 20 15,061,974 2.70E-06 NJ21 开花时间 Flowering time
20_34.9 Chr.20: 34852481 20 34,852,481 1.14E-06 BLUP 开花时间 Flowering time
20_34.9 Chr.20: 34852481 20 34,852,481 5.46E-06 NJ21 开花时间 Flowering time
1_53.5 Chr.01: 53455419 1 53,455,419 9.52E-06 NJ22 成熟期 Maturity date
3_24.9 Chr.03: 24877913 3 24,877,913 3.16E-06 NJ21 成熟期 Maturity date
3_38.2 Chr.03: 38239475 3 38,239,475 7.23E-06 NJ22 成熟期 Maturity date
3_40.6 Chr.03: 40648275 3 40,648,275 6.21E-06 NJ21 成熟期 Maturity date
4_13.3 Chr.04: 13315724 4 13,315,724 4.66E-06 BLUP 成熟期 Maturity date
5_7.8 Chr.05: 7788997 5 7,788,997 4.09E-06 BLUP 成熟期 Maturity date
5_39.9 Chr.05: 39904246 5 39,904,246 2.86E-06 NJ21 成熟期 Maturity date
6_15.2 Chr.06: 15190257 6 15,190,257 7.35E-06 BLUP 成熟期 Maturity date
6_31.6 Chr.06: 31617314 6 31,617,314 7.32E-06 NJ19 成熟期 Maturity date
6_31.6 Chr.06: 31628655 6 31,628,655 7.33E-06 NJ22 成熟期 Maturity date
6_32.7 Chr.06: 32672780 6 32,672,780 7.58E-06 NJ19 成熟期 Maturity date
6_51.3 Chr.06: 51320491 6 51,320,491 6.32E-08 NJ21 成熟期 Maturity date
7_20.3 Chr.07: 20259235 7 20,259,235 5.12E-06 BLUP 成熟期 Maturity date
8_24 Chr.08: 23981243 8 23,981,243 1.96E-06 BLUP 成熟期 Maturity date
8_38.8 Chr.08: 38794063 8 38,794,063 4.41E-06 BLUP 成熟期 Maturity date
8_38.9 Chr.08: 38878988 8 38,878,988 3.73E-06 NJ21 成熟期 Maturity date
8_38.9 Chr.08: 38884153 8 38,884,153 1.09E-06 BLUP 成熟期 Maturity date
9_32.5 Chr.09: 32534471 9 32,534,471 8.01E-06 NJ22 成熟期 Maturity date
10_4 Chr.10: 3977824 10 3,977,824 7.76E-06 BLUP 成熟期 Maturity date
10_4 Chr.10: 4023746 10 4,023,746 4.71E-06 NJ22 成熟期 Maturity date
10_43.4 Chr.10: 43371195 10 43,371,195 4.43E-08 BLUP 成熟期 Maturity date
11_24.6 Chr.11: 24616338 11 24,616,338 5.72E-06 BLUP 成熟期 Maturity date
11_25.4 Chr.11: 25447626 11 25,447,626 3.71E-06 NJ22 成熟期 Maturity date
11_25.4 Chr.11: 25448017 11 25,448,017 4.33E-07 BLUP 成熟期 Maturity date
11_25.5 Chr.11: 25452041 11 25,452,041 1.93E-06 BLUP 成熟期 Maturity date
11_25.5 Chr.11: 25452041 11 25,452,041 7.79E-06 NJ22 成熟期 Maturity date
11_25.9 Chr.11: 25916809 11 25,916,809 2.96E-06 NJ22 成熟期 Maturity date
11_25.9 Chr.11: 25937388 11 25,937,388 1.75E-06 BLUP 成熟期 Maturity date
11_26 Chr.11: 25950712 11 25,950,712 7.50E-06 BLUP 成熟期 Maturity date
11_26 Chr.11: 25971580 11 25,971,580 1.41E-06 NJ22 成熟期 Maturity date
11_26 Chr.11: 25992150 11 25,992,150 4.75E-06 NJ21 成熟期 Maturity date
12_20.7 Chr.12: 20727547 12 20,727,547 6.38E-06 NJ19 成熟期 Maturity date
12_28.5 Chr.12: 28523263 12 28,523,263 7.51E-06 NJ22 成熟期 Maturity date
12_35.6 Chr.12: 35626973 12 35,626,973 2.72E-06 NJ19 成熟期 Maturity date
12_35.7 Chr.12: 35655153 12 35,655,153 2.10E-06 BLUP 成熟期 Maturity date
12_35.7 Chr.12: 35666432 12 35,666,432 2.60E-06 NJ19 成熟期 Maturity date
13_11 Chr.13: 10977075 13 10,977,075 9.32E-06 BLUP 成熟期 Maturity date
13_16.3 Chr.13: 16348650 13 16,348,650 7.87E-07 NJ19 成熟期 Maturity date
13_23.9 Chr.13: 23897160 13 23,897,160 7.14E-06 NJ21 成熟期 Maturity date
13_33 Chr.13: 33044518 13 33,044,518 3.72E-06 NJ21 成熟期 Maturity date
13_44.5 Chr.13: 44471214 13 44,471,214 9.06E-06 NJ21 成熟期 Maturity date
13_44.9 Chr.13: 44921511 13 44,921,511 6.89E-07 NJ21 成熟期 Maturity date
13_45 Chr.13: 45026819 13 45,026,819 3.37E-07 NJ21 成熟期 Maturity date
13_45.1 Chr.13: 45057706 13 45,057,706 1.31E-06 NJ21 成熟期 Maturity date
14_36.2 Chr.14: 36158402 14 36,158,402 2.59E-06 NJ21 成熟期 Maturity date
15_16.2 Chr.15: 16178266 15 16,178,266 5.94E-06 NJ21 成熟期 Maturity date
15_48.5 Chr.15: 48503783 15 48,503,783 4.93E-06 NJ21 成熟期 Maturity date
15_51.7 Chr.15: 51674778 15 51,674,778 3.43E-06 NJ22 成熟期 Maturity date
16_29.3 Chr.16: 29264471 16 29,264,471 3.64E-06 BLUP 成熟期 Maturity date
17_15.3 Chr.17: 15293172 17 15,293,172 8.12E-06 NJ21 成熟期 Maturity date
17_15.4 Chr.17: 15392244 17 15,392,244 2.69E-06 NJ21 成熟期 Maturity date
17_15.7 Chr.17: 15716574 17 15,716,574 8.85E-07 NJ21 成熟期 Maturity date
17_15.8 Chr.17: 15819838 17 15,819,838 5.76E-07 NJ21 成熟期 Maturity date
17_15.9 Chr.17: 15859190 17 15,859,190 9.86E-07 NJ21 成熟期 Maturity date
17_16.1 Chr.17: 16053819 17 16,053,819 1.32E-06 NJ21 成熟期 Maturity date
17_16.5 Chr.17: 16526522 17 16,526,522 8.52E-06 NJ21 成熟期 Maturity date
17_18 Chr.17: 18037446 17 18,037,446 7.41E-06 NJ21 成熟期 Maturity date
17_19.4 Chr.17: 19389681 17 19,389,681 8.03E-06 NJ21 成熟期 Maturity date
17_21.3 Chr.17: 21305063 17 21,305,063 1.56E-06 NJ21 成熟期 Maturity date
17_21.7 Chr.17: 21677960 17 21,677,960 7.10E-06 NJ21 成熟期 Maturity date
17_24.1 Chr.17: 24091653 17 24,091,653 3.33E-07 NJ21 成熟期 Maturity date
17_24.3 Chr.17: 24307894 17 24,307,894 3.43E-08 NJ21 成熟期 Maturity date
17_24.4 Chr.17: 24437170 17 24,437,170 2.72E-07 NJ21 成熟期 Maturity date
17_24.5 Chr.17: 24517740 17 24,517,740 6.14E-07 NJ21 成熟期 Maturity date
17_24.6 Chr.17: 24596822 17 24,596,822 2.97E-06 NJ21 成熟期 Maturity date
17_24.7 Chr.17: 24728155 17 24,728,155 2.90E-06 NJ21 成熟期 Maturity date
17_24.9 Chr.17: 24941611 17 24,941,611 1.80E-06 NJ21 成熟期 Maturity date
17_25 Chr.17: 25022937 17 25,022,937 1.06E-07 NJ21 成熟期 Maturity date
17_25.1 Chr.17: 25122017 17 25,122,017 9.61E-08 NJ21 成熟期 Maturity date
17_25.3 Chr.17: 25315919 17 25,315,919 9.81E-08 NJ21 成熟期 Maturity date
17_25.4 Chr.17: 25364731 17 25,364,731 8.51E-08 NJ21 成熟期 Maturity date
17_25.6 Chr.17: 25604042 17 25,604,042 2.97E-06 NJ21 成熟期 Maturity date
17_25.7 Chr.17: 25729099 17 25,729,099 6.43E-08 NJ21 成熟期 Maturity date
17_26.1 Chr.17: 26132875 17 26,132,875 1.50E-07 NJ21 成熟期 Maturity date
17_26.3 Chr.17: 26343874 17 26,343,874 3.43E-06 NJ21 成熟期 Maturity date
17_26.4 Chr.17: 26414971 17 26,414,971 8.51E-08 NJ21 成熟期 Maturity date
17_26.5 Chr.17: 26500711 17 26,500,711 2.97E-06 NJ21 成熟期 Maturity date
17_26.6 Chr.17: 26570104 17 26,570,104 7.37E-06 NJ21 成熟期 Maturity date
17_26.8 Chr.17: 26766840 17 26,766,840 1.61E-06 NJ21 成熟期 Maturity date
17_27.1 Chr.17: 27112154 17 27,112,154 9.17E-07 NJ21 成熟期 Maturity date
17_27.3 Chr.17: 27303504 17 27,303,504 8.51E-08 NJ21 成熟期 Maturity date
17_27.4 Chr.17: 27422112 17 27,422,112 2.97E-06 NJ21 成熟期 Maturity date
17_27.6 Chr.17: 27598440 17 27,598,440 2.97E-06 NJ21 成熟期 Maturity date
17_27.7 Chr.17: 27658349 17 27,658,349 3.20E-06 NJ21 成熟期 Maturity date
17_28 Chr.17: 27974644 17 27,974,644 5.78E-06 NJ21 成熟期 Maturity date
17_28.1 Chr.17: 28115569 17 28,115,569 2.84E-06 NJ21 成熟期 Maturity date
17_28.5 Chr.17: 28544029 17 28,544,029 4.05E-06 NJ21 成熟期 Maturity date
17_28.6 Chr.17: 28596114 17 28,596,114 3.51E-06 NJ21 成熟期 Maturity date
17_28.7 Chr.17: 28709764 17 28,709,764 2.97E-06 NJ21 成熟期 Maturity date
17_28.8 Chr.17: 28799844 17 28,799,844 2.97E-06 NJ21 成熟期 Maturity date
17_28.9 Chr.17: 28859690 17 28,859,690 2.97E-06 NJ21 成熟期 Maturity date
17_29 Chr.17: 28959749 17 28,959,749 8.57E-08 NJ21 成熟期 Maturity date
17_29.1 Chr.17: 29098100 17 29,098,100 1.28E-06 NJ21 成熟期 Maturity date
17_29.4 Chr.17: 29392667 17 29,392,667 8.57E-08 NJ21 成熟期 Maturity date
17_29.8 Chr.17: 29823861 17 29,823,861 4.17E-06 NJ21 成熟期 Maturity date
17_30.3 Chr.17: 30343066 17 30,343,066 2.97E-06 NJ21 成熟期 Maturity date
17_30.4 Chr.17: 30361157 17 30,361,157 8.57E-08 NJ21 成熟期 Maturity date
17_30.5 Chr.17: 30472789 17 30,472,789 7.80E-08 NJ21 成熟期 Maturity date
17_30.6 Chr.17: 30610259 17 30,610,259 1.71E-08 NJ21 成熟期 Maturity date
17_30.7 Chr.17: 30654151 17 30,654,151 7.48E-07 NJ21 成熟期 Maturity date
17_31 Chr.17: 31004770 17 31,004,770 5.47E-07 NJ21 成熟期 Maturity date
17_31.2 Chr.17: 31226723 17 31,226,723 1.71E-08 NJ21 成熟期 Maturity date
17_31.3 Chr.17: 31348809 17 31,348,809 7.48E-07 NJ21 成熟期 Maturity date
17_31.4 Chr.17: 31369154 17 31,369,154 7.48E-07 NJ21 成熟期 Maturity date
17_32.9 Chr.17: 32923007 17 32,923,007 2.01E-06 NJ21 成熟期 Maturity date
17_35.5 Chr.17: 35478288 17 35,478,288 3.01E-06 NJ21 成熟期 Maturity date
18_11.9 Chr.18: 11946154 18 11,946,154 7.23E-06 NJ22 成熟期 Maturity date
19_5.8 Chr.19: 5769650 19 5,769,650 7.46E-06 NJ19 成熟期 Maturity date
19_35.6 Chr.19: 35579170 19 35,579,170 6.67E-06 NJ21 成熟期 Maturity date
19_44.7 Chr.19: 44659389 19 44,659,389 5.30E-06 NJ21 成熟期 Maturity date
19_45.1 Chr.19: 45078893 19 45,078,893 3.16E-06 NJ21 成熟期 Maturity date
19_45.2 Chr.19: 45241319 19 45,241,319 7.93E-06 NJ21 成熟期 Maturity date
19_45.3 Chr.19: 45271120 19 45,271,120 2.93E-06 NJ21 成熟期 Maturity date
20_6.7 Chr.20: 6685140 20 6,685,140 9.82E-06 BLUP 成熟期 Maturity date
20_6.7 Chr.20: 6685140 20 6,685,140 4.44E-06 NJ19 成熟期 Maturity date
20_14.3 Chr.20: 14255074 20 14,255,074 8.23E-06 NJ22 成熟期 Maturity date

表1

在多个环境或性状中检测到的开花相关性状关联信号"

关联信号
Signal		 最显著位点
Peak loci		 染色体
Chr.		 位置
Position (bp)		 P值
P-value		 环境
Environment		 性状
Trait
2_0.2 Chr.02: 198739 02 198,739 9.87E-06 NJ22, NJ21 开花时间 Flowering time
3_40.6 Chr.03: 40635793 03 40,635,793 5.59E-06 NJ19, NJ21 开花时间 Flowering time
6_31.6 Chr.06: 31617314 06 31,617,314 7.32E-06 NJ19, NJ22 成熟期 Maturity date
6_51.3 Chr.06: 51320491 06 51,320,491 6.32E-08 NJ21 开花时间, 成熟期
Flowering time, maturity date
8_38.9 Chr.08: 38878988 08 38,878,988 3.73E-06 NJ21, BLUP 成熟期 Maturity date
10_4 Chr.10: 3977824 10 3,977,824 7.76E-06 NJ22, BLUP 成熟期 Maturity date
11_25.4 Chr.11: 25447626 11 25,447,626 3.71E-06 NJ22, BLUP 成熟期 Maturity date
11_26 Chr.11: 25950712 11 25,950,712 7.50E-06 NJ22, NJ21, BLUP 成熟期 Maturity date
12_35.7 Chr.12: 35655153 12 35,655,153 2.10E-06 NJ19, BLUP 成熟期 Maturity date
14_48.4 Chr.14: 48432986 14 48,432,986 6.47E-06 NJ19, BLUP 开花时间 Flowering time
15_17.9 Chr.15: 17862246 15 17,862,246 3.73E-06 NJ22, BLUP 开花时间 Flowering time
16_5.3 Chr.16: 5275907 16 5,275,907 1.13E-06 NJ21, BLUP 开花时间 Flowering time
16_5.7 Chr.16: 5719462 16 5,719,462 7.64E-06 NJ21, BLUP 开花时间 Flowering time
18_0.8 Chr.18: 841400 18 841,400 5.18E-07 NJ19, NJ21 开花时间 Flowering time
20_6.7 Chr.20: 6685140 20 6,685,140 9.82E-06 NJ19, BLUP 成熟期 Maturity date
20_34.9 Chr.20: 34852481 20 34,852,481 1.14E-06 NJ21, BLUP 开花时间 Flowering time

表2

参与开花时间和成熟期调控的候选基因"

图2

相关位点Chr18: 841400的候选基因鉴定 (a): Chr.18:841400的局部曼哈顿图和连锁不平衡热图; (b): 候选基因的表达谱; (c): Chr.18: 743337、Chr.18: 850114和Chr.18: 895954三个相关位点在大豆自然群体中的单倍型分析。SAM: 茎尖分生组织。"

图3

相关位点Chr.03: 40635793的候选基因鉴定 (a): Chr.03: 40635793的局部曼哈顿图和连锁不平衡热图; (b): 候选基因的表达谱; (c): 相关位点在自然群体中的单倍型分析。SAM: 茎尖分生组织。"

图4

相关位点Chr.16: 5275907的候选基因鉴定 (a): Chr.16: 5275907的局部曼哈顿图和连锁不平衡热图; (b): 候选基因的表达谱; (c): 相关位点在自然群体中的单倍型分析。"

表S2

成熟期相关候选基因的表达模式"

基因编号
Gene ID		 根
Root		 根毛
Root hairs		 根瘤
Nodules		 茎
Stem		 叶
Leaves		 茎尖分生组织
SAM		 花
Flower		 荚
Pod		 种子
Seed
Glyma.10G043300 7.66 7.77 8.50 11.15 28.58 9.85 27.42 12.39 12.55
Glyma.10G043500 0 0 0 0.08 0.04 0.06 0.09 0.05 0.02
Glyma.10G043600 9.71 11.05 10.49 8.77 6.97 8.27 11.12 9.09 9.43
Glyma.10G043700 18.14 2.15 1.65 13.31 14.26 12.31 10.84 4.27 16.52
Glyma.10G043900 0.34 1.35 1.53 1.71 2.45 2.30 1.73 2.37 2.44
Glyma.10G044400 26.50 8.28 9.43 6.40 21.92 3.95 9.31 5.20 2.39
Glyma.10G044800 0.10 0.89 4.63 5.42 0 0.05 21.12 1.24 17.61
Glyma.10G044900 0 0.05 0.09 0.63 0.03 0 1.20 0.02 0.43
Glyma.10G045000 12.59 4.32 3.34 2.29 0.11 3.05 6.19 1.48 9.05
Glyma.10G045100 1.37 1.59 2.83 0.79 0.08 2.31 0.67 0.08 0.07
Glyma.10G045200 1.51 0.33 0.20 0 0 0 0 0 0
Glyma.10G045600 1.82 2.80 5.81 5.92 2.75 4.98 1.64 7.31 10.45
Glyma.10G045700 0.81 0.44 0.15 0.41 0.98 15.00 2.44 7.34 2.52
Glyma.12G194400 1.67 0.73 0.01 1.48 8.26 16.99 10.74 9.34 11.65
Glyma.12G196100 0.35 0.43 0.58 0.12 0 0.59 0.52 0.15 0.82
Glyma.12G196200 1.16 2.74 3.21 1.25 64.15 1.61 84.01 1.84 2.34
Glyma.06G324300 122.59 213.10 98.41 129.56 7.69 68.99 5.38 27.69 89.66
Glyma.06G324500 10.04 8.88 8.39 8.93 9.40 6.93 14.55 6.63 8.30
Glyma.06G324700 6.12 9.86 9.39 8.83 21.79 34.41 6.63 12.94 20.16
Glyma.06G324800 2.60 4.08 3.33 6.50 3.69 5.32 3.93 6.68 4.53
Glyma.06G325000 0 0 0 0 0 0 0 0 0
Glyma.06G325100 0.35 1.16 0.28 1.59 19.50 7.62 2.49 4.93 4.54
Glyma.06G325300 103.78 82.57 48.05 104.96 33.73 67.34 25.18 60.08 16.79
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