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作物学报

作物学报 ›› 2021, Vol. 47 ›› Issue (10): 1874-1890.doi: 10.3724/SP.J.1006.2021.04216

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

花生荚果与种子相关性状QTL定位及与环境互作分析

孟鑫浩1(), 张靖男1, 崔顺立1, Charles Y.Chen2, 穆国俊1, 侯名语1, 杨鑫雷1,*(), 刘立峰1,*()   

  1. 1华北作物改良与调控国家重点实验室 / 河北省种质资源重点实验室 / 河北农业大学, 河北保定 071001
    2奥本大学作物、土壤与环境科学系, 美国奥本 36849
  • 收稿日期:2020-09-22 接受日期:2021-03-19 出版日期:2021-10-12 网络出版日期:2021-04-08
  • 通讯作者: 杨鑫雷,刘立峰
  • 作者简介:E-mail: mxinhao1994@126.com
  • 基金资助:
    国家现代农业产业技术体系建设专项(CARS-13);国家自然科学基金项目(31701459);国家自然科学基金项目(31771833);河北省科技计划项目(16226301D);河北省现代农业产业技术体系油料创新团队项目(HBCT2018090202);河北省青年拔尖人才项目(0602015)

QTL mapping and QTL × Environment interaction analysis of pod and seed related traits in cultivated peanut (Arachis hypogaea L.)

MENG Xin-Hao1(), ZHANG Jing-Nan1, CUI Shun-Li1, Charles Y. Chen2, MU Guo-Jun1, HOU Ming-Yu1, YANG Xin-Lei1,*(), LIU Li-Feng1,*()   

  1. 1State Key Laboratory of North China for Crop Improvement and Regulation / Key laboratory of Crop Germplasm Resources of Hebei Province / Hebei Agricultural University, Baoding 071001, Hebei, China
    2Department of Crop, Soil and Environmental Sciences, Auburn University, Auburn 36849, USA
  • Received:2020-09-22 Accepted:2021-03-19 Published:2021-10-12 Published online:2021-04-08
  • Contact: YANG Xin-Lei,LIU Li-Feng
  • Supported by:
    Special Fund for Modern Agro-industry Technology Research System of China(CARS-13);National Natural Science Foundation of China(31701459);National Natural Science Foundation of China(31771833);Science and Technology Research and Development Program of Hebei Province(16226301D);Earmarked Fund for Hebei Oil Crop Innovation Team of Modern Agro-industry Technology Research System(HBCT2018090202);Support Program for the Top Young Talents of Hebei Province(0602015)

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摘要：

花生荚果、种子性状与产量紧密相关, 是重要的农艺性状。为挖掘与荚果、种子性状紧密连锁的分子标记, 本研究以大果品种冀花5号和小果美国资源M130组配衍生的315个家系RIL8群体为材料, 利用SSR、AhTE、SRAP和TRAP等标记构建了一张包含363个多态性位点的遗传连锁图谱。该图谱共包含21个连锁群, 总长为1360.38 cM, 标记间平均距离为3.75 cM。利用完备区间作图法对2017—2018年5个环境的荚果、种子相关性状进行数量性状基因座(quantitative trait locus, QTL)分析, 共鉴定到97个与荚果、种子性状相关的QTL, 可解释的表型变异为2.36%~12.15%, 分布在A02、A05、A08、A09、B02、B03、B04、B08和B09等9条染色体上。其中, 9个与荚果长相关, 13个与荚果宽相关, 14个与荚果厚相关, 11个与种子长相关, 14个与种子宽相关, 13个与种子厚相关, 13个与百果重相关, 10个与百仁重相关; 4个主效QTL分别为qPWA08.1、qPTA08.3、qPTA08.4和qSWB08.5, 可解释的表型变异分别为10.02%、11.06%、12.15%和11.97%; 45个稳定表达的QTL在3个以上环境可被重复检测; 连锁群A02、A08、B02、B04和B08上存在QTL聚集区。另外, 检测到15对上位性QTL, 可解释的表型变异为10.23%~51.84%。研究结果将为花生荚果、种子性状的分子标记辅助育种提供重要的理论依据。

关键词: 花生, 荚果, 种子, QTL, QTL×E

Abstract:

Pod and seed traits are important agronomy traits, which are closely related to yield in cultivated peanut (Arachis hypogaea L.). In the present study, to identify molecular markers closely linked to pod and seed traits, a RIL8 population with 315 families was developed that derived from Jihua 5 with large pod and M130 with small pod of US germplasm. A genetic linkage map containing 363 polymorphic loci was constructed using SSR, AhTE, SRAP, and TRAP markers. All polymorphic loci were mapped on 21 linkage groups, which spanned 1360.38 cM with an average distance of 3.75 cM. Subsequently, a total of 97 QTLs for pod and seed traits were identified by ICIM method at five environments from 2017 to 2018, explaining the phenotypic variations of 2.36%-12.15%, and located on A02, A05, A08, A09, B02, B03, B04, B08, and B09 chromosomes. Among them, nine QTLs were detected for pod length, 13 QTLs for pod width, 14 QTLs for pod thickness, 11 QTLs for seed length, 13 QTLs for seed width, 13 QTLs for hundred-pod weight, 10 QTLs for hundred-seed weight. Four QTLs with major effect were detected, including qPWA08.1, qPTA08.3, qPTA08.4, and qSWB08.5, which explained the phenotypic variations of 10.02%-12.15%. Furthermore, 45 stable QTLs were repeatedly detected in more than three environments. QTL clusters were detected on A02, A08, B02, B04, and B08 chromosomes, respectively. In addition, 15 epistatic QTLs were identified that explaining phenotypic variation of 10.23%-51.84%. These results will provide an important theoretical basis for molecular marker-assisted breeding of pod and seed traits in peanut.

Key words: peanut, pod, seed, QTL, QTL×E

图1

荚果及种子表型图 白色指示条为标尺, 大小为1 cm。"

表1

亲本及RIL群体的描述性统计结果"

性状
Trait		 环境
Environment		 亲本 Parents RIL群体 RIL population
冀花5号
Jihua 5		 M130 最小值
Minimum		 最大值
Maximum		 平均值±标准误
Mean±SD		 变异系数
CV		 Shapiro-Wilk
(w-test)		 偏度
Skewness		 峰度
Kurtosis
荚果长 17QY 43.28±1.46** 30.25±1.24 24.09 47.94 33.08±3.80 0.11 0.9592*** 0.75 0.72
PL (mm) 17DM 43.57±0.96** 29.84±0.65 24.81 49.04 34.72±4.13 0.12 0.9606*** 0.69 0.52
18QY 45.33±0.99** 31.66±1.15 23.96 49.04 34.53±4.18 0.12 0.9685*** 0.58 0.20
18DM 41.25±2.08** 28.84±1.06 24.80 44.60 33.23±3.61 0.11 0.9685*** 0.46 -0.18
18QA 43.59±1.03** 30.18±1.17 23.35 46.95 33.01±3.79 0.11 0.9723** 0.53 0.13
荚果宽 17QY 15.17±0.39** 12.42±0.50 9.12 17.88 12.86±1.63 0.13 0.9649*** 0.54 0.24
PW (mm) 17DM 15.16±0.97** 12.81±0.51 9.32 18.04 12.94±1.72 0.13 0.9597*** 0.47 0.13
18QY 14.95±1.03** 11.90±0.49 10.16 19.09 12.77±1.65 0.13 0.9327*** 0.91 0.74
18DM 15.40±0.81** 12.95±1.12 9.21 19.79 13.50±1.73 0.13 0.9668*** 0.61 0.25
18QA 15.02±0.86** 12.51±0.79 9.79 18.20 12.68±1.57 0.12 0.9347*** 0.87 0.53
荚果厚 17QY 15.33±0.54** 12.98±0.43 8.51 17.17 12.30±1.66 0.13 0.9262*** 0.85 0.22
PT (mm) 17DM 15.53±0.90** 13.2±0.38 8.78 18.79 12.41±1.72 0.14 0.9254*** 0.98 0.77
18QY 15.28±0.65** 13.19±0.59 9.65 19.50 13.21±1.64 0.12 0.9647*** 0.66 0.72
18DM 15.39±0.85** 12.77±0.71 9.16 17.73 13.46±1.58 0.12 0.982* 0.04 -0.14
18QA 15.23±0.95** 12.86±0.35 9.32 19.65 12.98±1.62 0.12 0.9698** 0.6 0.79
种子长 17QY 19.56±0.64** 16.38±0.96 12.26 23.09 16.43±1.88 0.11 0.9691** 0.54 0.24
SL (mm) 17DM 19.58±0.38** 16.12±0.56 11.89 22.94 16.94±2.04 0.12 0.9728** 0.38 -0.15
18QY 20.32±0.79** 17.30±0.79 13.34 23.33 17.17±1.95 0.11 0.9594 *** 0.57 0.01
18DM 18.79±1.75** 15.47±0.91 12.98 22.44 16.46±1.66 0.10 0.9723** 0.47 0.10
18QA 18.18±0.69** 16.03±0.81 13.06 22.03 16.63±1.65 0.10 0.9703** 0.47 0.17
种子宽 17QY 9.89±0.24** 8.67±0.63 5.89 11.02 8.29±0.89 0.11 0.9852* 0.01 0.09
SW (mm) 17DM 9.78±0.53** 8.39±0.22 6.33 11.19 8.37±0.80 0.10 0.9849* 0.20 0.36
18QY 10.42±0.74** 8.91±0.62 7.14 10.91 8.82±0.65 0.07 0.9725** 0.44 0.17
18DM 9.37±0.78** 8.43±0.54 6.89 11.32 8.70±0.67 0.08 0.9855* 0.35 0.64
18QA 9.49±0.40* 8.57±0.89 6.39 10.93 8.32±0.68 0.08 0.9848* 0.34 0.52
种子厚 17QY 10.29±0.32** 9.46±0.45 6.00 10.04 7.78±0.76 0.10 0.9783* 0.20 -0.22
ST (mm) 17DM 10.28±0.38** 9.20±0.37 6.18 10.39 7.96±0.67 0.08 0.9763* 0.51 0.82
18QY 10.45±0.93* 9.74±0.61 7.29 12.42 9.41±0.77 0.08 0.9899* 0.09 0.40
18DM 10.13±0.45** 9.18±0.94 6.91 11.96 9.34±0.82 0.09 0.9853* -0.13 0.19
18QA 9.83±0.36* 9.07±0.65 6.95 11.99 9.33±0.90 0.10 0.9811* 0.17 0.06
百果重 17QY 254.03±6.84** 170.13±0.91 71.04 239.22 144.50±30.93 0.21 0.9812* 0.19 -0.12
HPW (g) 17DM 247.18±2.30** 164.64±3.78 79.86 245.97 157.07±33.15 0.21 0.9681*** 0.37 -0.23
18QY 271.28±8.80** 184.48±2.45 94.85 325.53 180.11±40.66 0.23 0.9678*** 0.54 0.19
18DM 236.77±6.46** 155.79±3.57 61.32 256.43 163.31±36.51 0.22 0.9825* 0.05 -0.17
18QA 233.38±1.89** 159.70±5.95 78.28 296.87 159.77±34.14 0.21 0.9788* 0.51 0.51
百仁重 17QY 105.22±2.17** 70.04±0.69 27.73 91.24 56.08±11.05 0.20 0.9859* 0.05 -0.11
HSW (g) 17DM 100.82±1.49** 68.83±1.14 36.93 101.23 62.80±12.11 0.19 0.9719** 0.42 0.07
18QY 114.32±2.73** 76.20±3.11 43.73 119.87 74.77±13.96 0.19 0.9775* 0.34 -0.03
18DM 96.12±2.48** 63.88±1.06 30.07 106.51 67.38±13.71 0.20 0.9826* 0.14 -0.21
18QA 100.24±1.44** 63.90±1.65 37.21 108.20 63.25±13.05 0.21 0.9666 *** 0.53 0.14

图2

不同环境下RIL群体各性状的频率分布图 缩写同表1。"

表2

不同环境下各性状的相关性分析结果"

环境
Environment		 性状
Trait		 荚果长
PL		 荚果宽
PW		 荚果厚
PT		 种子长
SL		 种子宽
SW		 种子厚
ST		 百果重
HPW
17QY 荚果宽PW 0.6536***
荚果厚PT 0.7286*** 0.9226***
种子长SL 0.8522*** 0.6346*** 0.7011***
种子宽SW 0.2752*** 0.5291*** 0.4692*** 0.5357***
种子厚ST 0.3724*** 0.4974*** 0.5333*** 0.5713*** 0.7453***
百果重HPW 0.6587*** 0.6515*** 0.6454*** 0.6549*** 0.5196*** 0.5614***
百仁重HSW 0.6242*** 0.6188*** 0.6282*** 0.6737*** 0.6041*** 0.5970*** 0.8280***
17DM 荚果宽PW 0.6667***
荚果厚PT 0.7353*** 0.9087***
种子长SL 0.8430*** 0.6208*** 0.6751***
种子宽SW 0.2599*** 0.4804*** 0.3924*** 0.4370***
种子厚ST 0.3991*** 0.5326*** 0.6052*** 0.5025*** 0.6033***
百果重HPW 0.6214*** 0.6927*** 0.6884*** 0.6196*** 0.4634*** 0.5350***
百仁重HSW 0.6037*** 0.6143*** 0.6171*** 0.6326*** 0.5256*** 0.5961*** 0.8651***
18QY 荚果宽PW 0.7442***
荚果厚PT 0.6400*** 0.9129***
种子长SL 0.8711*** 0.6054*** 0.4848***
种子宽SW 0.4281*** 0.6622*** 0.5653*** 0.4141***
种子厚ST 0.3921*** 0.5563*** 0.5647*** 0.4347*** 0.6530***
百果重HPW 0.6801*** 0.6522*** 0.5825*** 0.7041*** 0.5554*** 0.6193***
百仁重HSW 0.7373*** 0.7132*** 0.6362*** 0.7571*** 0.6475*** 0.6927*** 0.8726***
18DM 荚果宽PW 0.6995***
荚果厚PT 0.6364*** 0.9102***
种子长SL 0.8786*** 0.6328*** 0.5997***
种子宽SW 0.4231*** 0.5523*** 0.5578*** 0.4899***
种子厚ST 0.3408*** 0.4766*** 0.5764*** 0.4895*** 0.6946***
百果重HPW 0.6586*** 0.6285*** 0.6474*** 0.6724*** 0.6040*** 0.6302***
百仁重HSW 0.6676*** 0.6262*** 0.6347*** 0.7130*** 0.6717*** 0.7052*** 0.8606***
18QA 荚果宽PW 0.7231***
荚果厚PT 0.6760*** 0.9333***
种子长SL 0.8586*** 0.6254*** 0.5920***
种子宽SW 0.2765*** 0.4873*** 0.4558*** 0.3500***
种子厚ST 0.3117*** 0.4761*** 0.5240*** 0.4215*** 0.6740***
百果重HPW 0.5869*** 0.6571*** 0.6460*** 0.6029*** 0.4700*** 0.5189***
百仁重HSW 0.4829*** 0.5246*** 0.5032*** 0.5874*** 0.5972*** 0.6089*** 0.8070***

表3

RIL群体各性状方差分析及广义遗传力"

变异来源
Source of variation		 自由度
DF		 荚果长
PL (mm)		 荚果宽
PW (mm)		 荚果厚
PT (mm)		 种子长
SL (mm)		 种子宽
SW (mm)		 种子厚
ST (mm)		 百果重
HPW (g)		 百仁重
HSW (g)
区组间Block/environment 5 2.05 ns 3.39 ns 5.87 ns 2.84 ns 3.23 ns 4.53 ns 29.36 ns 14.19 ns
基因型Genotype (G) 314 143.74** 101.27** 105.53** 46.78** 10.30** 17.39** 31.78** 77.12**
环境Environment (E) 4 502.85** 310.54** 915.58** 112.56** 154.97** 2337.02** 349.28** 1898.64**
基因型×环境G × E 1196 8.79** 7.36** 7.53** 3.82** 3.08** 4.54** 4.30** 10.18**
遗传变异系数GVC (%) — 12.87 15.34 14.21 13.96 16.39 15.68 17.54 17.28
广义遗传力h2B (%) — 64.61 60.76 61.21 65.88 61.71 67.50 54.09 57.83

图3

花生遗传连锁图谱 缩写同表1。"

图4

本研究遗传图谱与整合图谱的共线性"

表4

荚果、种子相关性状QTL定位结果"

性状
Trait		 位点
QTL		 环境Environment 染色体
Chr.		 位置
Position		 范围
Range		 标记区间
Marker interval		 LOD 可解释的表型变异PVE (%) 加性效应
Additive		 来源
Variety
荚果长 qPLA08.5 17QY A08 19.317 1.395-24.581 me3em14-196-Ah4-4 3.0543 4.5412 1.0026 冀花5号Jihua 5
PL (mm) qPLB04.2 18QY B04 48.814 45.2-53.084 T3em5-340-me1em3-75 3.1524 4.4841 -1.052 M130
qPLA02.2 18QY A02 33 28.586-35.04 AHGS1163-AHGS1886 3.4276 4.1455 -1.009 M130
qPLA08.2 18QY A08 16.487 1.395-24.581 me3em14-196-Ah4-4 3.165 5.3035 1.142 冀花5号Jihua 5
qPLB04.1 18DM B04 31.814 29.825-32.253 PMc348-GM1641 2.7216 2.4116 -0.7788 M130
qPLA08.3 18DM A08 17.43 1.395-24.581 me3em14-196-Ah4-4 3.6112 5.5059 1.1219 冀花5号Jihua 5
qPLA02.1 18QA A02 31 28.586-35.04 AHGS1163-AHGS1886 3.1466 3.8886 -0.8444 M130
qPLA08.4 18QA A08 17.43 1.395-24.581 me3em14-196-Ah4-4 3.1664 6.2318 1.0698 冀花5号Jihua 5
qPLA08.1 18QA A08 8.403 2.674-9.42 HBAUAh177-AhTE0658 3.1246 3.7403 0.83 冀花5号Jihua 5
荚果宽 qPWA08.9 17QY A08 27 24.581-32.375 Ah4-4-TC9B08 3.0347 3.4027 0.3487 冀花5号Jihua 5
PW (mm) qPWA08.1 17QY A08 5.539 2.674-9.42 HBAUAh177-AhTE0658 8.3529 10.0181 0.6021 冀花5号Jihua 5
qPWA09.1 17DM A09 47.976 44.512-52.51 AHS1620-me4em15-230 3.4111 5.6928 -0.4961 M130
qPWA08.5 17DM A08 25 24.581-32.375 Ah4-4-TC9B08 2.6823 2.4856 0.3265 冀花5号Jihua 5
qPWB08.1 17DM B08 0 0-6.125 AHGS1286-TC20B05 4.5802 4.1738 -0.426 M130
qPWA08.2 17DM A08 6.971 2.674-9.42 HBAUAh177-AhTE0658 7.2488 8.0948 0.5908 冀花5号Jihua 5
qPWA08.6 18QY A08 26 24.581-32.375 Ah4-4-TC9B08 3.405 4.9727 0.3919 冀花5号Jihua 5
qPWA08.7 18DM A08 27 24.581-32.375 Ah4-4-TC9B08 5.1642 6.0632 0.4812 冀花5号Jihua 5
qPWB08.3 18DM B08 1 0-6.125 AHGS1286-TC20B05 4.8423 5.3595 -0.4547 M130
qPWA08.3 18DM A08 6.971 2.674-9.42 HBAUAh177-AhTE0658 6.2926 7.5419 0.5384 冀花5号Jihua 5
qPWA08.8 18QA A08 27 24.581-32.375 Ah4-4-TC9B08 4.0778 5.4765 0.3976 冀花5号Jihua 5
qPWB08.2 18QA B08 0 0-6.125 AHGS1286-TC20B05 3.2573 3.7144 -0.3288 M130
qPWA08.4 18QA A08 7.687 2.674-9.42 HBAUAh177-AhTE0658 5.0836 6.6321 0.439 冀花5号Jihua 5
荚果厚 qPTA08.6 17QY A08 22.146 1.395-24.581 me3em14-196-Ah4-4 4.2717 6.0053 0.4499 冀花5号Jihua 5
PT (mm) qPTA08.1 17QY A08 6.255 2.674-9.42 HBAUAh177-AhTE0658 3.8997 5.4416 0.4305 冀花5号Jihua 5
qPTA08.7 17DM A08 25 24.581-32.375 Ah4-4-TC9B08 4.683 5.2195 0.434 冀花5号Jihua 5
qPTB08.1 17DM B08 0 0-6.125 AHGS1286-TC20B05 3.5352 3.7738 -0.3716 M130
qPTA08.2 17DM A08 6.255 2.674-9.42 HBAUAh177-AhTE0658 4.1823 5.3897 0.4421 冀花5号Jihua 5
qPTA08.5 18QY A08 12 9.42-15.535 AhTE0658-TC6H03 3.3146 4.8431 0.3898 冀花5号Jihua 5
qPTA09.1 18DM A09 46.976 44.512-52.51 AHS1620-me4em15-230 3.9877 5.5482 -0.4349 M130
qPTA08.8 18DM A08 27 24.581-32.375 Ah4-4-TC9B08 3.8396 3.7738 0.3577 冀花5号Jihua 5
qPTA05.1 18DM A05 25.186 20.904-39.785 me13em5-112-GM1577 2.5575 2.6714 -0.3018 M130
qPTB08.3 18DM B08 2 0-6.125 AHGS1286-TC20B05 6.4241 6.1373 -0.4581 M130
qPTA08.3 18DM A08 7.688 2.674-9.42 HBAUAh177-AhTE0658 11.0128 11.0554 0.6137 冀花5号Jihua 5
qPTA08.9 18QA A08 27 24.581-32.375 Ah4-4-TC9B08 3.3743 4.1636 0.3621 冀花5号Jihua 5
qPTB08.2 18QA B08 0 0-6.125 AHGS1286-TC20B05 3.3221 3.5571 -0.3361 M130
qPTA08.4 18QA A08 7.688 2.674-9.42 HBAUAh177-AhTE0658 9.437 12.1489 0.6205 冀花5号Jihua 5
种子长 qSLB04.3 17QY B04 51.814 45.2-53.084 T3em5-340-me1em3-75 3.5098 4.5251 -0.4663 M130
SL (mm) qSLA02.4 17QY A02 34 28.586-35.04 AHGS1163-AHGS1886 4.5943 5.4414 -0.5079 M130
qSLA02.2 17DM A02 33 28.586-35.04 AHGS1163-AHGS1886 5.3238 7.4048 -0.6248 M130
qSLB04.2 18QY B04 49.814 45.2-53.084 T3em5-340-me1em3-75 4.3546 6.0388 -0.5601 M130
qSLA02.3 18QY A02 33 28.586-35.04 AHGS1163-AHGS1886 5.7194 6.4678 -0.5774 M130
qSLA08.2 18QY A08 18.373 1.395-24.581 me3em14-196-Ah4-4 4.392 6.4226 0.576 冀花5号Jihua 5
qSLB02.1 18QY B02 10 10-29.536 AHTE0398-CTW_NEW_38 2.5587 2.3638 -0.3495 M130
qSLB04.1 18DM B04 16.405 43.26-4.103 AHTE0001-AHM091 3.9425 5.5578 -0.43 M130
qSLA08.1 18DM A08 9.119 2.674-9.42 HBAUAh177-AhTE0658 2.898 3.8087 0.3432 冀花5号Jihua 5
qSLA02.1 18QA A02 32 28.586-35.04 AHGS1163-AHGS1886 3.3469 3.8192 -0.3879 M130
qSLA08.3 18QA A08 18.373 1.395-24.581 me3em14-196-Ah4-4 4.1987 6.4454 0.5045 冀花5号Jihua 5
种子宽 qSWB08.1 17QY B08 0 0-6.125 AHGS1286-TC20B05 3.3062 5.0411 -0.197 M130
SW (mm) qSWB08.2 17DM B08 0 0-6.125 AHGS1286-TC20B05 3.2831 3.6766 -0.1703 M130
qSWA08.2 17DM A08 26 15.535-26.911 TC6H03-AhTE0477 3.9168 4.8359 0.1949 冀花5号Jihua 5
qSWA08.4 18QY A08 27 24.581-32.375 Ah4-4-TC9B08 2.7094 3.7658 0.1335 JH5
qSWB08.4 18QY B08 2 0-6.125 AHGS1286-TC20B05 5.52 7.3282 -0.188 M130
qSWB02.1 18QY B02 12 10-29.536 AHTE0398-CTW_NEW_38 2.5171 3.4186 -0.1275 M130
qSWA09.1 18DM A09 23.976 23.052-24.982 RN27A10-AHTE0122 2.6959 3.1535 0.1305 冀花5号Jihua 5
qSWA02.1 18DM A02 30 28.586-35.04 AHGS1163-AHGS1886 3.4558 3.8727 0.1449 冀花5号Jihua 5
qSWB08.5 18DM B08 3 0-6.125 AHGS1286-TC20B05 9.9761 11.9698 -0.2551 M130
qSWA08.1 18DM A08 12 9.42-15.535 AhTE0658-TC6H03 4.1572 4.7351 0.1598 冀花5号Jihua 5
qSWA02.2 18QA A02 32 28.586-35.04 AHGS1163-AHGS1886 3.2739 4.3875 0.1518 冀花5号Jihua 5
qSWA08.3 18QA A08 26 24.581-32.375 Ah4-4-TC9B08 3.511 4.5457 0.1544 冀花5号Jihua 5
qSWB08.3 18QA B08 0 0-6.125 AHGS1286-TC20B05 4.8277 5.6235 -0.1723 M130
qSWB02.2 18QA B02 13 10-29.536 AHTE0398-CTW_NEW_38 3.055 4.4285 -0.1526 M130
种子厚 qSTA09.1 17QY A09 22.976 21.827-23.052 T1me13-75-RN27A10 2.9792 4.3897 0.1586 冀花5号Jihua 5
ST (mm) qSTB08.1 17QY B08 0 0-6.125 AHGS1286-TC20B05 4.1523 5.9184 -0.1856 M130
qSTB08.5 17DM B08 1 0-6.125 AHGS1286-TC20B05 3.511 5.2695 -0.1578 M130
qSTA08.3 18QY A08 19.317 1.395-24.581 me3em14-196-Ah4-4 4.3155 6.9533 0.2296 冀花5号Jihua 5
qSTB08.2 18QY B08 0 0-6.125 AHGS1286-TC20B05 4.6686 4.8687 -0.1935 M130
qSTA08.4 18QY A08 21 15.535-26.911 TC6H03-AhTE0477 2.8698 3.8907 0.1716 冀花5号Jihua 5
qSTA09.2 18DM A09 24.976 23.052-24.982 RN27A10-AHTE0122 3.8787 4.14 0.1812 冀花5号Jihua 5
qSTB08.3 18DM B08 0 0-6.125 AHGS1286-TC20B05 5.4215 5.8448 -0.2147 M130
qSTB03.1 18DM B03 59.043 27.96-64.668 GM1954-IPAHM103 2.8689 3.5976 0.1682 冀花5号Jihua 5
qSTA08.1 18DM A08 10 9.42-15.535 AhTE0658-TC6H03 6.5305 7.3459 0.24 冀花5号Jihua 5
qSTA08.5 18QA A08 29 24.581-32.375 Ah4-4-TC9B08 5.1955 7.3853 0.2579 冀花5号Jihua 5
qSTB08.4 18QA B08 0 0-6.125 AHGS1286-TC20B05 2.608 3.1638 -0.1697 M130
qSTA08.2 18QA A08 18 15.535-26.911 TC6H03-AhTE0477 3.5829 5.2767 0.2178 冀花5号Jihua 5
百果重 qHPWA08.8 17QY A08 31 24.581-32.375 Ah4-4-TC9B08 2.612 2.884 6.1294 J冀花5号ihua 5
HPW (g) qHPWB08.3 17QY B08 1 0-6.125 AHGS1286-TC20B05 4.2913 4.8043 -7.9731 M130
qHPWA08.1 17QY A08 6.971 2.674-9.42 HBAUAh177-AhTE0658 3.942 5.0068 8.1239 冀花5号Jihua 5
qHPWB08.1 17DM B08 0 0-6.125 AHGS1286-TC20B05 5.4732 6.1589 -9.2979 M130
qHPWA08.2 17DM A08 6.971 2.674-9.42 HBAUAh177-AhTE0658 3.361 4.5239 7.9317 冀花5号Jihua 5
qHPWA08.5 18QY A08 26 24.581-32.375 Ah4-4-TC9B08 3.7742 5.4674 10.1994 冀花5号Jihua 5
qHPWA09.1 18DM A09 1.574 1.243-1.881 me10em6-244-T3me4-50 2.9275 4.4348 9.1722 冀花5号Jihua 5
qHPWA08.6 18DM A08 27 24.581-32.375 Ah4-4-TC9B08 4.3701 4.589 9.3388 冀花5号Jihua 5
qHPWB08.4 18DM B08 1 0-6.125 AHGS1286-TC20B05 6.3447 6.2963 -10.993 M130
qHPWA08.3 18DM A08 7.687 2.674-9.42 HBAUAh177-AhTE0658 7.3085 7.9283 12.3093 冀花5号Jihua 5
qHPWA08.7 18QA A08 27 24.581-32.375 Ah4-4-TC9B08 3.3108 4.3355 7.727 冀花5号Jihua 5
qHPWB08.2 18QA B08 0 0-6.125 AHGS1286-TC20B05 3.1924 3.6078 -7.0792 M130
qHPWA08.4 18QA A08 8.403 2.674-9.42 HBAUAh177-AhTE0658 6.215 7.9641 10.5143 冀花5号Jihua 5
百仁重 qHSWA09.2 17QY A09 19.976 19.851-20.044 AHGS0362-AHS1950 2.7386 3.9653 2.281 冀花5号Jihua 5
HSW (g) qHSWA08.6 17QY A08 28 24.581-32.375 Ah4-4-TC9B08 3.2048 4.984 2.5511 冀花5号Jihua 5
qHSWA08.4 18QY A08 26 24.581-32.375 Ah4-4-TC9B08 6.1813 8.9514 4.4051 冀花5号Jihua 5
qHSWB02.1 18QY B02 10 10-29.536 AHTE0398-CTW_NEW_38 3.4265 4.1876 -3.0213 M130
qHSWA08.5 18DM A08 27 24.581-32.375 Ah4-4-TC9B08 3.116 4.3944 2.9937 冀花5号Jihua 5
qHSWB08.1 18DM B08 0 0-6.125 AHGS1286-TC20B05 4.3546 5.07 -3.2325 M130
qHSWA08.1 18DM A08 9.119 2.674-9.42 HBAUAh177-AhTE0658 4.7443 5.7178 3.4257 冀花5号Jihua 5
qHSWA09.1 18QA A09 1.574 1.243-1.881 me10em6-244-T3me4-50 2.6987 4.1254 3.2042 冀花5号Jihua 5
qHSWA08.2 18QA A08 18.373 1.395-24.581 me3em14-196-Ah4-4 2.6696 4.3569 3.306 冀花5号Jihua 5
qHSWA08.3 18QA A08 20 15.535-26.911 TC6H03-AhTE0477 4.2105 5.6344 3.7483 冀花5号Jihua 5

表5

荚果和种子相关性状的上位性QTL定位结果"

性状
Trait		 上位性QTL名称
Epi-QTL name		 环境
Environment		 染色体
Chr.		 位置1
Position 1		 标记区间1
Marker interval 1		 上位性QTL名称
Epi-QTL name		 染色体
Chr.		 位置2
Position 2		 标记区间2
Marker interval 2		 LOD 可解释的遗传变异
PVE (%)
PL (mm) Epi-qPLA09.1 18BD A09 50.976 AHS1620-me4em15-230 Epi-qPLA05.1 A05 50 T2me4-75-seq18C2 5.09 10.23
PW (mm) Epi-qPWB08.1 17BD B08 5 AHGS1286-TC20B05 Epi-qPWA08.2 A08 25 TC6H03-AhTE0477 5.33 11.95
Epi-qPWA08.1 18BD A08 15.544 me3em14-196-Ah4-4 Epi-qPWA08.1 A08 20.26 me3em14-196-Ah4-4 5.54 42.46
PT (mm) Epi-qPTB09.1 17BD B09 5 T3me2-388-AHGS1576 Epi-qPTB09.1 B09 10 T3me2-388-AHGS1576 6.05 39.57
Epi-qPTA02.3 17BD A02 50 AHGS1886-AHGS1159 Epi-qPTA02.3 A02 55 AHGS1886-AHGS1159 5.05 40.63
Epi-qPTA08.1 17BD A08 10.827 me3em14-196-Ah4-4 Epi-qPTA08.1 A08 15.544 me3em14-196-Ah4-4 7.66 51.84
Epi-qPTA03.1 17BD A03 40 AhTE0570-TC4G02 Epi-qPTA03.1 A03 45 AhTE0570-TC4G02 6.42 37.16
Epi-qPTA10.1 17BD A10 20 AhTE0586-AHGS1192 Epi-qPTA10.1 A10 25 AHGS1192-seq3e10 6.56 38.94
Epi-qPTA02.2 17DM A02 45 AHGS1886-AHGS1159 Epi-qPTA02.2 A02 50 AHGS1886-AHGS1159 6.05 40.39
Epi-qPTA08.2 17DM A08 15.544 me3em14-196-Ah4-4 Epi-qPTA08.2 A08 20.26 me3em14-196-Ah4-4 5.01 39.62
Epi-qPTA02.1 18QA A02 10 pPGSseq14F4-Ah3TC13E05 Epi-qPTA02.1 A02 15 Ah3TC13E05-AHGS1463 5.37 26.30
SL (mm) Epi-qSLB08.1 17BD B08 0 AHGS1286-TC20B05 Epi-qSLA08.1 A08 25 TC6H03-AhTE0477 7.60 11.61
HPW (g) Epi-qHPWB08.1 17BD B08 5 AHGS1286-TC20B05 Epi-qHPWA08.1 A08 25 TC6H03-AhTE0477 8.05 17.75
Epi-qHPWB08.2 18DM B08 5 AHGS1286-TC20B05 Epi-qHPWA08.2 A08 25 TC6H03-AhTE0477 5.04 15.36
HSW (g) Epi-qHSWB08.1 17BD B08 5 AHGS1286-TC20B05 Epi-qHSWA08.1 A08 25 TC6H03-AhTE0477 7.30 13.99

表6

荚果和种子相关性状的加性QTL与环境互作结果"

性状
Trait		 位置
Position		 标记区间
Marker interval		 LOD LOD(A) LOD
(AbyE)		 可解释的遗传变异PVE 可解释的遗传变异PVE(A) 可解释的遗传变异PVE
(AbyE)		 加性效应
Add		 AbyE_01 AbyE_02 AbyE_03 AbyE_04 AbyE_05
HPW 1.976 T3me4-50-me10em13-84 3.2854 3.162 0.1234 0.6316 0.6176 0.014 0.3661 -0.006 -0.0408 0.0953 0.016 -0.0645
SL 16.405 AHM091-AHTE0001 6.4625 6.2415 0.2211 1.1913 1.145 0.0462 -0.5154 0.1206 -0.1441 -0.0132 0.1118 -0.0751
PL 32.814 GM1641-Ah3TC39B04 7.7019 7.1093 0.5927 1.3629 1.2801 0.0828 -0.5506 0.1878 0.0587 -0.178 -0.1467 0.0782
SL 49.814 T3em5-340-me1em3-75 6.2931 5.6064 0.6866 1.2594 1.1008 0.1586 -0.4907 0.0103 -0.1087 -0.282 0.2596 0.1208
SL 33 AHGS1163-AHGS1886 10.3475 9.9861 0.3615 1.9775 1.9001 0.0773 -0.6416 0.027 0.0248 -0.1949 0.2023 -0.0592
ST 37.706 GM1954-seq2H08 3.4137 3.1998 0.2139 0.683 0.6243 0.0588 -0.3667 -0.011 -0.2127 0.0458 0.0907 0.0872

表7

荚果和种子相关性状的上位性QTL与环境互作效应"

Chr.1 位置1
Position 1		 性状
Trait		 标记区间1
Marker interval 1		 Chr.2 位置2
Position 2		 标记区间2
Marker interval 2		 LOD LOD
(AAbyE)		 PVE PVE
(AAbyE)		 Add1 Add2 AddbyAdd AAbyE_01 AAbyE_02 AAbyE_03 AAbyE_04 AAbyE_05
A09 45.976 PL AHS1620-me4em15-230 B07 0 TC1A08-TC9H09 7.0121 0.0728 1.9524 0.0197 -0.1218 0.0794 -0.5448 -0.0493 -0.0654 0.0668 0.0148 0.0331
A09 50.976 PL AHS1620-me4em15-230 A06 35 me7em1-83-me8em16-92 7.8477 0.0188 2.2588 0.0066 0.0036 0.2423 -0.5959 0.0149 -0.0405 -0.0235 0.0533 -0.0043
A09 45.976 PL AHS1620-me4em15-230 B03 31.236 GM1954-IPAHM103 10.0123 0.4698 2.6465 0.0887 -0.183 -0.3668 0.6229 -0.1048 0.1381 0.139 -0.1102 -0.0622
A02 0 PT pPGSseq14F4-Ah3TC13E05 A05 5 me7em9-96-me13em5-112 8.6941 0.3066 2.3716 0.0727 -0.0142 -0.1429 -0.5926 -0.0159 -0.1916 0.0964 0.0257 0.0854
A02 60 PT AHGS1886-AHGS1159 A03 10 RM17H09-me8em1-286 7.1732 0.1598 1.9488 0.0372 0.1981 0.0409 -0.5484 0.0342 0.011 -0.1155 -0.0432 0.1136
A02 45 PT AHGS1886-AHGS1159 A06 36.412 TC7C06-AHTE0372 8.959 0.1938 2.44 0.0354 -0.2324 0.4976 -0.608 -0.0709 -0.0324 -0.069 0.11 0.0623
A08 6.111 PT me3em14-196-Ah4-4 B03 2.834 AHBGSC1003E10-GM1996 10.1986 0.0214 2.8149 0.0074 0.5756 0.21 0.6596 0.0296 0.0139 0.0111 -0.0598 0.0053
A08 1.395 PT me3em14-196-Ah4-4 A08 25 TC6H03-AhTE0477 5.4087 0.2251 1.4752 0.0536 0.4861 0.1886 -0.4783 -0.1029 0.175 -0.0222 -0.0274 -0.0225
A08 15.544 PT me3em14-196-Ah4-4 B02 10 AHTE0398-CTW_NEW_38 6.5397 0.3284 1.6356 0.0709 0.7735 -0.5558 -0.4935 0.0526 -0.1013 -0.0825 -0.0451 0.1763
B08 0 SL AHGS1286-TC20B05 B03 59.043 GM1954-IPAHM103 5.0287 0.5997 1.2864 0.1371 -0.0958 -0.1712 0.422 -0.1143 0.2612 -0.0654 -0.1314 0.05
B08 0 SL AHGS1286-TC20B05 A08 25 TC6H03-AhTE0477 15.7588 0.1295 4.009 0.0244 -0.1851 0.4017 0.7983 0.106 0.0345 -0.072 -0.022 -0.0465
A03 35 HSW AhTE0570-TC4G02 B03 6.453 AHGS1940a-AHGS1940b 9.1572 0.2059 2.4169 0.0351 -0.2737 -0.1534 -0.6022 0.0048 -0.1046 0.0963 -0.0615 0.065
A10 20 HSW AhTE0586-AHGS1192 B02 25 AHTE0398-CTW_NEW_38 5.4877 0.1016 1.4894 0.0194 0.0107 -0.2795 -0.4755 0.0571 -0.0731 -0.0229 0.0769 -0.038
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