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作物学报

作物学报 ›› 2019, Vol. 45 ›› Issue (6): 856-871.doi: 10.3724/SP.J.1006.2019.83059

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

8种水旱环境下2个玉米群体穗部性状QTL间的上位性及环境互作效应分析

赵小强,任斌,彭云玲(),徐明霞,方鹏,庄泽龙,张金文,曾文静,高巧红,丁永福,陈奋奇   

  1. 甘肃省干旱生境作物学重点实验室 / 甘肃农业大学农学院, 甘肃兰州 730070
  • 收稿日期:2018-08-16 接受日期:2018-12-24 出版日期:2019-06-12 网络出版日期:2019-06-12
  • 通讯作者: 赵小强,任斌,彭云玲
  • 作者简介:赵小强, E-mail: zhaoxq3324@163.com
  • 基金资助:
    本研究由国家重点研发计划项目(2018YFD0100203-4);中国科学院“西部之光”项目(20180504);甘肃省重点研发计划(18YF1NA071);甘肃省自然科学基金项目(18JR3RA189);甘肃省重大科技专项(17ZD2NA016)

Epistatic and QTL × environment interaction effects for ear related traits in two maize (Zea mays) populations under eight watering environments

Xiao-Qiang ZHAO,Bin REN,Yun-Ling PENG(),Ming-Xia XU,Peng FANG,Ze-Long ZHUANG,Jin-Wen ZHANG,Wen-Jing ZENG,Qiao-Hong GAO,Yong-Fu DING,Fen-Qi CHEN   

  1. Gansu Provincial Key Laboratory of Aridland Crop Science / College of Agronomy, Gansu Agricultural University, Lanzhou 730070, Gansu, China
  • Received:2018-08-16 Accepted:2018-12-24 Published:2019-06-12 Published online:2019-06-12
  • Contact: Xiao-Qiang ZHAO,Bin REN,Yun-Ling PENG
  • Supported by:
    This study was supported by the National Key R&D Project(2018YFD0100203-4);Chinese Academy of Sciences “Light of West China” Program(20180504);Lanzhou Sci & Technol Project (2018-1-103), the Key R&D Program of Gansu, China(18YF1NA071);the National Science Foundation of Gansu Province(18JR3RA189);the Key Science and Technology Projects in Gansu Province(17ZD2NA016)

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摘要：

深入剖析干旱胁迫条件下玉米穗部性状的遗传机制可为玉米抗旱高产分子育种提供参考依据。以大穗型旱敏感自交系TS141为共同亲本, 分别与小穗型强抗旱自交系廊黄和昌7-2杂交, 构建了含有202个(LTPOP)和218个(CTPOP)家系的F2:3群体, 在8种水旱环境下进行单穗重、穗轴重、穗粒重、百粒重、出籽率及穗长等6个穗部性状的表型鉴定, 并采用复合区间作图法(CIM)和基于混合线性模型的复合区间作图法(MCIM)对其进行单环境和多环境联合数量性状位点(QTL)分析。结果表明, 采用CIM法, 单环境下在2套F2:3群体间检测到62个穗部性状QTL, 其中干旱胁迫环境下检测到38个QTL, 进一步在2套F2:3群体多个干旱胁迫环境下检测到10个稳定表达的QTL (sQTL), 分别位于Bin 1.01-1.03、Bin 1.03-1.04、Bin 1.05、Bin 1.07、Bin 1.07-1.08、Bin 2.04、Bin 4.08、Bin 5.06-5.07、Bin 6.05和Bin 9.04-9.06。采用MCIM法, 联合分析定位到54个穗部性状联合QTL, 其中24个表现显著的QTL与环境互作(QTL×E), 17对参与了显著的加性与加性/显性(AA/AD)上位性互作, 其表型贡献率较低。这些研究结果可为系统地剖析玉米穗部性状的分子遗传机制提供理论依据; 且这2套F2:3群体多个环境下检测到的sQTL可作为穗部性状改良的重要候选染色体区段, 用于图位克隆或抗旱高产分子育种, 但要注重环境及上位性互作效应的影响。

关键词: 玉米, 干旱, 穗部性状, QTL, QTL与环境互作, 上位性

Abstract:

Exploring genetic mechanisms of ear related traits in maize (Zea mays) under drought stress is important in maize molecular breeding for drought tolerance and high yield. Two F2:3 populations, namely 202 F2:3 families (LTPOP) and 218 F2:3 families (CTPOP) derived from the common male parent TS141 with drought-sensitive and larger ear and female parents Langhuang/Chang 7-2 with higher drought tolerance and small ear were used to investigate ear weight (EW), cob weight (CW), grain weight (GW), 100-kernel weight (KW), kernel ratio (KR), and ear length (EL) under eight watering environments, and then to analyze quantitative trait locus (QTL) in a single environment by composite interval mapping (CIM) and in the eight environments by mixed linear model based on composite interval mapping (MCIM). Sixty-two QTLs for ear related traits were detected in two F2:3 populations in a single environment by CIM, among than 38 QTLs were mapped under water-stressed environments, and further analysis showed that ten stable QTLs (sQTLs) were simultaneously identified in two F2:3 populations under multiple water-stressed environments, these sQTLs were located in Bin 1.01-1.03, Bin 1.03-1.04, Bin 1.05, Bin 1.07, Bin 1.07-1.08, Bin 2.04, Bin 4.08, Bin 5.06-5.07, Bin 6.05, and Bin 9.04-9.06. Fifty-four joint QTLs for ear related traits were identified in the eight environments by joint analysis with MCIM, among than 24 had significant QTL by environment interaction (QTL×E), and 17 significant epistatic interactions with additive by additive/dominance (AA/AD) effects, with less phenotypic variation. These results lay a foundation for systematically revealing molecular genetic mechanism of ear related traits, these sQTLs detected in two F2:3 populations under multiple environments are important genomic regions that could be used in positional cloning and molecular breeding for drought tolerance and high yield, however, more attention should be paid to the effects of environment or epistatic interaction.

Key words: maize (Zea mays), drought, ear correlative traits, QTL, QTL by environment interaction (QTL×E), epistasis

图1

4个试验点(武威、张掖、古浪和景泰)气象数据"

表1

8种水分环境下F2:3群体(LTPOP/CTPOP) 6个穗部性状的表型值"

性状Trait 环境
Env.		 双亲
Parents		 F1杂交种
F1 hybrid		 LTPOP群体
LTPOP population
廊黄
Langhuang		 TS141 F1LT 均值
Mean		 变幅
Range		 变异系数
CV (%)		 偏度Skewness 峰度Kurtosis
EW E1 71.33±6.89 111.30±9.72 184.27±7.03 149.81±43.25 52.85-285.91 28.87 0.315 0.313
(g) E2 66.58±7.95 100.67±8.44 163.51±6.79 133.19±51.83 43.27-265.43 38.92 0.453 -0.327
E3 88.29±10.43 128.85±6.45 197.80±6.74 163.19±41.49 70.21-281.87 25.42 0.086 -0.338
E4 84.14±6.77 110.56±8.01 169.14±7.83 152.76±51.41 52.27-276.94 33.65 0.382 -0.577
CW E1 12.53±1.65 29.90±2.15 33.71±2.81 27.65±9.43 9.47-53.31 34.11 0.326 -0.490
(g) E2 9.85±1.23 21.81±1.89 26.58±2.44 25.11±8.23 7.91-50.34 32.77 0.230 -0.051
E3 15.75±2.05 33.78±3.13 37.83±3.15 30.79±10.85 10.87-60.03 35.22 0.636 -0.141
E4 11.66±2.78 23.04±2.61 31.09±3.07 28.17±9.56 8.50-51.23 33.92 0.294 -0.410
GW E1 59.81±4.70 82.40±4.82 150.56±5.11 123.16±33.98 65.07-208.15 27.59 0.815 0.206
(g) E2 56.73±3.96 71.92±3.17 136.93±5.09 108.22±24.72 61.14-186.27 22.84 -0.192 -0.887
E3 64.54±4.51 85.07±4.66 159.97±4.85 132.46±25.05 68.94-207.92 18.91 -0.620 0.753
E4 60.48±5.03 73.52±4.49 138.05±4.23 125.01±23.36 65.31-194.58 18.68 0.904 0.727
KW E1 19.70±2.51 25.15±1.67 40.06±3.40 27.05±7.52 18.40-50.15 27.80 0.853 0.728
(g) E2 15.80±1.38 20.05±1.41 33.95±4.18 25.65±6.39 12.00-45.50 24.91 0.308 -0.376
E3 20.55±1.62 32.70±1.72 43.97±3.79 33.80±7.20 13.75-60.25 21.30 0.617 -0.407
E4 25.70±1.70 24.80±2.03 31.20±3.73 25.90±6.45 13.35-45.50 24.90 0.467 -0.339
性状Trait 环境
Env.		 双亲
Parents		 F1杂交种
F1 hybrid		 LTPOP群体
LTPOP population
廊黄
Langhuang		 TS141 F1LT 均值
Mean		 变幅
Range		 变异系数
CV (%)		 偏度Skewness 峰度Kurtosis
KR E1 83.43±1.84 77.34±1.67 83.93±2.01 81.59±4.94 70.63-91.08 6.05 0.711 -0.095
E2 81.02±1.35 73.08±1.55 82.15±1.84 78.80±5.79 65.97-88.84 7.35 -0.093 0.268
E3 85.14±1.42 79.62±1.73 85.92±1.85 83.01±5.05 71.89-92.51 6.08 -0.216 0.670
E4 82.26±1.61 73.98±1.89 83.93±1.79 80.56±5.93 68.05-89.93 7.36 -0.326 -0.431
EL E1 9.23±1.68 15.33±1.74 20.17±1.40 15.74±2.72 9.18-22.78 17.29 -0.069 0.000
(cm) E2 8.05±1.21 13.10±1.46 18.83±2.05 14.82±3.08 8.20-21.30 20.74 -0.203 -0.855
E3 9.55±2.02 15.62±1.31 20.84±1.87 15.81±2.92 9.10-23.20 18.46 0.073 -0.089
E4 8.72±1.04 12.60±1.76 17.68±1.92 15.07±2.52 8.21-19.80 16.69 -0.497 0.047
双亲
Parents		 F1杂交种
F1 hybrid		 CTPOP群体
CTPOP population
昌7-2
Chang 7-2		 TS141 F1CT 均值
Mean		 变幅
Range		 变异系数
CV (%)		 偏度Skewness 峰度Kurtosis
EW E5 63.68±4.57 107.64±7.99 235.32±7.15 121.00±39.89 27.07-224.56 32.97 0.528 0.041
(g) E6 59.05±6.22 96.47±6.80 215.76±6.98 111.20±38.62 22.34-211.91 34.73 0.596 0.152
E7 54.79±3.16 101.26±5.43 228.43±7.68 105.30±38.29 13.20-200.00 36.37 -0.033 -0.191
E8 50.00±2.69 90.43±5.90 205.37±8.03 90.03±41.93 12.07-198.34 46.26 0.238 -0.220
CW E5 6.86±3.30 21.41±3.88 30.28±1.89 17.51±6.71 4.24-41.21 38.33 0.577 0.619
(g) E6 5.90±5.28 17.68±4.72 26.77±2.01 15.81±5.84 4.16-35.99 36.94 0.701 1.037
E7 6.22±2.19 18.64±3.61 28.95±1.68 16.81±5.74 5.09-32.56 34.14 0.258 -0.119
E8 5.03±3.00 15.11±2.45 23.72±1.74 14.49±6.12 4.03-32.08 42.26 0.425 -0.368
GW E5 56.82±4.01 86.25±3.77 205.07±7.56 104.49±33.19 25.17-184.38 31.76 -0.911 0.150
(g) E6 52.12±3.65 73.79±4.09 189.99±6.70 95.39±28.05 19.78-180.21 29.41 0.228 1.006
E7 49.57±4.22 82.63±4.73 199.48±5.79 89.87±30.82 10.92-174.89 34.29 0.858 0.417
E8 43.97±3.99 70.32±4.36 180.55±6.04 77.54±31.21 9.95-173.24 40.25 0.325 0.271
KW E5 15.15±1.24 23.40±2.14 34.77±2.13 25.20±6.27 6.15-46.14 24.88 0.275 0.761
(g) E6 13.95±1.90 20.10±1.00 29.94±1.78 22.00±7.50 5.05-45.60 34.09 0.435 1.069
E7 14.05±1.05 21.21±1.12 31.26±1.69 22.80±6.29 8.40-42.71 27.58 0.483 0.407
E8 10.80±1.76 14.97±1.33 28.78±1.55 21.60±5.37 6.15-44.43 24.86 0.438 0.696
KR E5 87.22±2.16 80.19±2.32 88.36±1.69 85.50±4.89 73.47-90.80 5.72 0.916 0.188
E6 85.81±2.02 73.73±1.85 86.03±1.57 83.17±5.63 65.13-87.96 6.77 -0.470 0.735
E7 84.64±1.90 79.42±2.13 84.75±1.96 84.04±5.30 71.90-91.83 6.31 0.438 -0.307
E8 80.09±2.11 70.68±2.26 80.77±2.01 79.99±6.37 62.38-90.48 7.96 0.691 -0.740
EL E5 8.15±2.36 13.15±1.77 21.86±2.04 13.55±2.50 7.30-21.20 18.47 0.522 0.903
(cm) E6 7.02±2.69 10.44±3.10 17.99±1.21 12.49±2.32 6.80-20.60 18.59 0.164 0.455
E7 7.08±1.57 12.53±2.85 19.78±11.36 12.91±2.14 7.30-20.30 16.57 0.357 0.723
E8 6.00±2.31 10.48±2.08 16.04±1.02 12.01±2.59 5.70-18.30 21.52 -0.283 -0.218

图2

6个穗部性状RC和杂种优势分析缩写见表1。Abbreviations correspond with these given in Table 1."

Table 2

Complex variance for six ear related traits in F2:3 populations （LTpop/CTpop）under eight watering envrionments"

Table 2

Phenoty and genetic correlation analysis among six ear relaits in F2:3 populations （LTpop/CTpop）"

图3

采用CIM和MCIM法不同水分环境下F2:3群体（LTpop/CTpop）6个穗部性状QTL在染色体上的分布 "

表4

多环境下采用MCIM法对F2:3群体(LTPOP/CTPOP) 6个穗部性状联合QTL及QTL×E分析"

性状Trait QTL Chr. QTL位置 QTL position A AE1/
AE5		 AE2/
AE6		 AE3/
AE7		 AE4/
AE8		 h2A
(%)		 h2AE
(%)
cM Mb 标记区间
Marker interval
LTPOP群体 LTPOP population
EW qEW-Ch.1-2 1 60.7 0.05 umc2025-umc1395 -2.07 12.56
qEW-J2-1 2 89.2 4.40 bnlg1520-umc1736 -1.55 -0.81 -0.95 9.43 7.08
qEW-Ch.4-1 4 183.6 46.16 umc2041-umc2287 -0.98 7.18
qEW-Ch.9-1 9 54.4 16.69 umc1120-umc2134 -1.70 6.33
qEW-J10-1 10 2.7 0.26 umc1319-bnlg1451 1.41 0.66 0.89 12.95 6.59
CW qCW-Ch.1-1 1 35.8 22.79 umc2224-bnlg1484 1.09 0.86 10.20 8.12
qCW-Ch.2-1 2 23.0 2.71 umc1555-umc1024 0.30 4.98
qCW-J2-1 2 87.5 4.40 bnlg1520-umc1736 -1.37 -0.79 -0.65 9.03 6.84
qCW-Ch.4-1 4 179.9 46.16 umc2041-umc2287 0.24 6.16
qCW-Ch.9-1 9 67.3 23.41 umc1120-umc2346 -1.58 8.54
GW qGW-J1-1 1 114.2 5.03 phi308707-umc1847 -1.20 -0.64 -0.37 0.45 8.14 5.01
qGW-J2-1 2 10.5 0.39 umc2363-umc2403 -0.53 4.85
qGW-J2-2 2 101.8 0.24 bnlg1520-umc1736 -1.55 -0.86 -0.40 -0.71 9.31 6.86
qGW-Ch.4-1 4 181.9 1.32 umc2041-umc2287 -1.31 8.32
qGW-Ch.8-1 8 44.2 0.01 bnlg1863-umc2075 -0.35 -0.26 -0.17 2.79 1.24
KW qKW-Ch.1-2 1 114.2 5.03 phi308707-umc1847 -1.06 -0.63 -0.20 -0.48 8.02 2.87
qKW-Ch.4-1 4 181.0 4.16 umc2041-umc2287 -1.28 8.17
qKW-J6-1 6 94.9 20.76 bnlg2191-mmc0523 0.72 5.48
qKW-Ch.6-1 6 119.7 11.12 umc2040-bnlg1174a 0.41 3.56
性状Trait QTL Chr. QTL位置 QTL position A AE1/
AE5		 AE2/
AE6		 AE3/
AE7		 AE4/
AE8		 h2A
(%)		 h2AE
(%)
cM Mb 标记区间
Marker interval
KR qKR-Ch.1-1 1 40.7 1.56 bnlg1484-umc1917 1.11 11.69
qKR-J1-1 1 95.4 2.30 bnlg1025-mmc0041 0.60 0.37 5.30 2.73
qKR-Ch.6-1 6 119.8 0.03 umc2040-bnlg1174a 0.47 3.51
qKR-Ch.7-1 7 110.5 1.01 umc1708-umc1768 0.65 5.37
qKR-J8-1 8 88.3 0.54 umc2356-umc1607 0.84 0.26 0.43 -0.31 8.14 2.20
EL qEL-Ch.9-1 9 66.5 23.41 umc1120-umc2346 0.64 5.36
qEL-Ch.10-1 10 50.1 2.85 umc1345-umc2016 0.82 6.49
CTPOP群体 CTPOP population
EW qEW-Ch.1-1 1 138.4 17.57 bnlg1025-mmc0041 -0.90 5.06
qEW-J2-1 2 124.3 4.40 bnlg1520-umc1736 -1.76 -0.88 -0.74 10.93 6.00
qEW-Ch.5-1 5 236.0 2.96 umc2216-umc1072 1.03 4.71
qEW-Ch.6-1 6 81.3 22.97 mmc0523-umc2141 -0.77 4.84
qEW-J8-1 8 8.8 4.96 umc1327-bnlg1194 1.19 0.95 4.90 4.63
qEW-J10-1 10 4.9 0.26 umc1319-bnlg1451 1.01 0.81 0.58 0.66 10.25 7.11
CW qCW-Ch.1-1 1 27.1 22.79 umc2224-bnlg1484 1.10 0.73 0.58 6.49 4.57
qCW-J1-1 1 159.0 19.78 phi308707-umc2289 1.13 -0.79 -0.43 0.68 5.40 4.34
qCW-J5-1 5 113.8 15.76 umc1226-umc1815 -0.88 -0.54 4.92 3.59
qCW-Ch.8-1 8 107.0 32.89 umc2218-umc2356 -1.20 7.93
qCW-Ch.9-1 9 51.8 16.69 umc1120-umc2134 -1.82 9.29
qCW-J10-1 10 3.3 0.26 umc1319-bnlg1451 1.07 0.55 6.05 4.18
GW qGW-J1-1 1 154.9 17.51 mmc0041-phi308707 -2.03 -0.84 -0.66 -0.41 -0.68 11.07 5.50
qGW-J2-1 2 38.6 2.01 umc2363-umc1024 -0.88 7.31
qGW-J2-2 2 127.1 0.24 bnlg1520-umc1736 -1.63 -0.89 -0.41 -0.64 -0.50 9.95 4.19
qGW-Ch.4-1 4 120.3 0.66 umc2041-umc2188 -1.77 10.14
qGW-J6-1 6 97.4 0.03 umc2040-bnlg1174a -1.05 8.96
qGW-J8-1 8 40.7 0.01 bnlg1863-umc2075 -0.84 -0.36 -0.52 7.20 4.47
KW qCW-J1-1 1 155.0 17.51 mmc0041-phi308707 -0.78 -0.58 7.10 6.03
qKW-Ch.4-1 4 `120.5 0.66 umc2041-umc2188 -1.02 8.95
qKW-Ch.6-1 6 81.0 22.97 mmc0523-umc2141 0.44 4.06
KR qKR-J1-1 1 139.3 2.30 bnlg1025-mmc0041 0.51 0.26 0.39 4.15 2.93
qKR-J6-1 6 88.1 0.17 umc2141-umc2040 0.58 4.20
qKR-J7-1 7 118.0 1.01 umc1708-umc1768 0.73 6.12
qKR-Ch.8-1 8 114.9 1.96 umc2356-phi233376 0.62 0.37 0.18 5.04 2.11
EL qEL-J1-1 1 154.6 17.51 mmc0041-phi308707 -0.79 -0.66 7.04 6.18
qEL-Ch.4-1 4 120.3 0.66 umc2041-umc2188 -1.32 9.16
qEL-Ch.10-1 10 142.6 0.75 bnlg1839-umc1249 0.36 3.41

Table 2

Epistatic interactions among QTLS for six ear related traits in F2:3 population （LTpop/CTpop）under multiple environments"

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