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作物学报

作物学报 ›› 2020, Vol. 46 ›› Issue (3): 341-353.doi: 10.3724/SP.J.1006.2020.94076

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

净作和套作下大豆贮藏蛋白11S、7S组分相对含量的QTL分析

刘代铃,谢俊锋,何乾瑞,陈四维,胡跃,周佳,佘跃辉,刘卫国,杨文钰(),武晓玲()   

  1. 四川农业大学农学院 / 四川省作物带状复合种植工程技术研究中心, 四川成都 611130
  • 收稿日期:2019-05-20 接受日期:2019-09-26 出版日期:2020-03-12 网络出版日期:2019-10-09
  • 通讯作者: 杨文钰,武晓玲
  • 作者简介:E-mail: icyfish00@163.com
  • 基金资助:
    本研究由国家重点研发计划项目(2017YFD0101500);四川省科技厅育种攻关项目资助(2016NYZ0031)

QTL analysis for relative contents of glycinin and β-conglycinin fractions from storage protein in soybean seeds under monoculture and relay intercropping

Dai-Ling LIU,Jun-Feng XIE,Qian-Rui HE,Si-Wei CHEN,Yue HU,Jia ZHOU,Yue-Hui SHE,Wei-Guo LIU,Wen-Yu YANG(),Xiao-Ling WU()   

  1. College of Agronomy, Sichuan Agricultural University / Sichuan Engineering Research Center for Crop Strip Intercropping System, Chengdu 611130, Sichuan, China
  • Received:2019-05-20 Accepted:2019-09-26 Published:2020-03-12 Published online:2019-10-09
  • Contact: Wen-Yu YANG,Xiao-Ling WU
  • Supported by:
    the Crops Breeding Project in Sichuan Province(2017YFD0101500);the Crops Breeding Project in Sichuan Province(2016NYZ0031)

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摘要：

大豆贮藏蛋白的7S组分和11S组分占蛋白质总量的70%左右, 其含量直接与大豆种子的加工品质、营养品质及功能特性密切相关。目前, 在玉米-大豆带状套作模式下11S和7S组分的相对含量会如何变化尚不明确。本研究以南豆12和九月黄为亲本所构建的重组自交系(RILs)群体为材料, 结合已构建的SNP高密度遗传图谱, 在不同环境(E1: 2017年仁寿; E2: 2017年雅安; E3: 2016年仁寿)的净作及套作下, 对控制贮藏蛋白组分含量的相关性状进行QTL分析。结果表明, 在不同的环境和种植方式下, 亲本和RIL群体的大部分贮藏蛋白11S、7S组分相对含量差异显著或极显著; 检测到10个相关QTL, 分布于9条染色体, 表型贡献率为5.63%~9.68%。通过注释信息, 在定位到的QTL区段中初步筛选出65个与贮藏蛋白质含量相关的候选基因。上述结果为大豆品质育种提供了理论参考。

关键词: 大豆, 贮藏蛋白, 荫蔽, QTL定位

Abstract:

The relative contents of glycinin and β-conglycinin from storage protein are closely related to the quality and function of soybean seeds. However, it is not clear whether or how glycinin and β-conglycinin contents change in maize-soybean strip intercropping system. The glycinin and β-conglycinin relative contents of the recombinant inbred lines (RILs) derived from the cross of ‘Nandou 12’ and ‘Jiuyuehuang’ were measured under different environments (E1: 2017, Renshou; E2: 2017, Ya’an; E3: 2016, Renshou) and planting patterns (M: monoculture; RI: relay intercropping), among which the differences were significant or extremely significant in parents and RILs. Based on a genetic linkage map with 6366 SNP markers, we detected ten QTLs for glycinin and β-conglycinin relative contents which were distributed in nine linkage groups with the phenotypic variation of 5.63%-9.68%. According to the soybean genomic information, 65 candidate genes were screened in the region of above-mentioned QTLs. These results lay a theoretical foundation for soybean quality breeding.

Key words: Soybean (Glycine max L.), storage protein, shade, QTL mapping

表1

不同环境及种植方式下亲本的贮藏蛋白11S、7S组分相对含量"

性状
Trait		 种植方式
Planting pattern		 E1 E2 E3
南豆12
ND12		 九月黄
JYH		 Sig. 南豆12
ND12		 九月黄
JYH		 Sig. 南豆12
ND12		 九月黄
JYH		 Sig.
7S (%) M 20.18±0.22 b 24.47±0.30 a ** 21.77±1.28 a 21.55±0.63 a NS 22.84±0.20 a 22.96±0.47 a NS
RI 23.42±1.40 a 25.25±2.05 a NS 23.36±1.64 a 21.25±1.16 a NS 22.43±2.21 a 22.21±1.45 a NS
11S (%) M 33.74±0.29 a 42.47±0.31 A ** 36.80±2.06 a 38.05±0.97 a NS 37.99±0.69 a 36.08±0.76 a *
RI 35.57±1.64 a 36.63±0.30 B NS 38.09±1.37 a 35.35±1.02 b * 33.30±3.60 a 38.12±2.03 a NS
11S+7S (%) M 53.92±0.51 b 66.94±0.27 a ** 58.58±3.33 a 59.60±1.59 a NS 60.84±0.88 a 59.04±1.18 a NS
RI 58.99±3.04 a 61.88±2.16 b NS 61.45±1.01 a 56.60±2.15 a * 55.73±5.80 a 60.33±3.48 a NS
11S/7S M 1.67±0.00 a 1.74±0.03 A * 1.69±0.01 a 1.77±0.01 a ** 1.66±0.02 A 1.57±0.02 B **
RI 1.52±0.02 b 1.46±0.11 B NS 1.64±0.16 a 1.67±0.05 b NS 1.48±0.02 B 1.72±0.02 A **
α' (%) M 6.67±0.03 B 8.35±0.06 a ** 7.06±0.22 a 7.12±0.17 a NS 7.08±0.54 a 8.01±0.16 A *
RI 7.66±0.26 A 8.84±0.96 a NS 7.29±0.83 a 6.74±0.83 a NS 7.68±0.94 a 6.90±0.36 B NS
Α (%) M 6.84±0.18 a 9.49±0.28 a ** 7.92±0.75 a 7.63±0.69 a NS 8.38±0.50 a 8.14±0.81 a NS
RI 8.29±0.93 a 9.89±0.89 a NS 8.50±0.92 a 7.25±0.15 a NS 8.45±0.92 a 7.72±0.72 a NS
Β (%) M 6.67±0.20 B 6.63±0.42 a NS 6.80±0.38 b 6.81±0.12 b NS 7.39±0.91 a 6.81±0.30 b NS
RI 7.47±0.23 A 6.52±0.66 a NS 7.57±0.27 a 7.26±0.19 a NS 6.30±0.60 a 7.59±0.37 a *
A3 (%) M 5.27±0.15 a 7.43±0.76 a ** 5.04±0.71 a 4.79±0.11 a NS 5.61±0.23 a 5.28±1.24 a NS
RI 4.71±0.61 a 6.52±1.07 a NS 5.47±0.21 a 4.65±0.31 a * 5.37±0.15 a 6.00±0.46 a NS
Acid (%) M 13.01±0.88 a 17.17±0.78 a ** 15.79±2.58 a 14.68±1.07 a NS 17.46±1.65 a 15.77±1.65 a NS
RI 14.14±0.65 a 16.35±2.29 a NS 16.98±1.68 a 14.56±1.20 a NS 13.72±1.33 b 17.39±0.44 a **
Basic (%) M 8.65±0.44 a 12.48±1.28 a ** 8.93±0.08 B 9.82±0.57 a NS 8.27±0.47 a 8.66±0.11 a NS
RI 9.36±1.42 a 9.45±2.24 a NS 9.90±0.15 A 9.24±0.96 a NS 8.95±0.71 a 9.13±0.45 a NS
A5 (%) M 6.81±0.34 a 5.39±0.66 a * 7.04±0.91 a 8.75±1.31 a NS 6.71±2.71 a 6.37±1.59 a NS
RI 7.36±0.92 a 4.42±1.10 a * 5.74±0.58 a 6.9±0.94 a NS 5.27±2.86 a 5.60±1.65 a NS

图 1

大豆贮藏蛋白亚基的SDS-PAGE图谱 1: 南豆12; 2: 九月黄; 3: 158号家系材料; 4: 104号家系材料。缩写同表1。"

图2

RIL群体贮藏蛋白亚基含量频数分布图 缩写同表1。"

表2

RIL群体贮藏蛋白11S、7S组分相对含量基本统计量分析"

性状
Trait		 环境
E		 种植方式
Planting
pattern		 平均数±标准差
Mean±SD		 最大值
Max.		 最小值
Min.		 极差
Range		 变异系数
CV (%)		 偏度
Skew.		 峰度
Kurt.		 P (S-W)
7S (%) E1 M 22.34±1.81 30.03 19.73 10.30 8.11 1.73 4.07 0.00
RI 24.30±2.67 29.35 19.17 10.18 10.99 -0.10 -1.30 0.00
E2 M 21.74±1.63 26.88 17.49 9.39 7.52 0.49 0.33 0.01
RI 22.44±1.45 26.24 18.16 8.08 6.45 0.03 0.06 0.74
E3 M 24.67±1.92 29.92 19.60 10.32 7.78 0.23 -0.37 0.21
RI 25.22±1.92 30.18 19.89 10.29 7.60 0.14 -0.04 0.35
11S (%) E1 M 36.16±2.68 42.62 30.03 12.59 7.42 0.14 -0.79 0.01
RI 35.65±2.27 41.83 29.24 12.59 6.37 0.15 -0.17 0.75
E2 M 37.73±2.05 42.69 32.39 10.30 5.43 0.26 -0.35 0.05
RI 36.57±2.30 44.09 29.91 14.18 6.28 0.25 0.52 0.20
E3 M 37.47±2.62 46.31 30.46 15.85 6.99 0.14 0.01 0.34
RI 36.22±2.74 44.36 30.87 13.49 7.56 0.81 0.48 0.00
11S+7S (%) E1 M 58.50±2.81 65.08 52.05 13.03 4.80 -0.03 -0.79 0.00
RI 59.95±2.85 67.16 53.25 13.91 4.76 -0.01 -0.30 0.59
E2 M 59.46±2.23 65.27 54.60 10.67 3.75 0.05 -0.74 0.04
RI 59.01±2.59 65.31 51.99 13.32 4.39 -0.04 0.01 0.30
E3 M 62.15±2.65 72.35 55.68 16.67 4.27 0.21 0.28 0.10
RI 61.45±2.76 69.69 55.35 14.34 4.49 0.66 0.08 0.00
11S/7S E1 M 1.63±0.19 2.10 1.10 1.00 11.67 -0.17 0.01 0.11
RI 1.49±0.22 2.03 1.06 0.97 14.67 0.29 -0.99 0.00
E2 M 1.75±0.18 2.37 1.32 1.05 10.36 0.26 0.35 0.06
RI 1.64±0.16 2.08 1.26 0.82 9.46 0.31 -0.13 0.22
E3 M 1.53±0.18 2.17 1.10 1.07 12.07 0.22 0.19 0.47
RI 1.45±0.18 1.93 1.09 0.84 12.32 0.30 -0.42 0.03
α' (%) E1 M 7.45±0.79 9.99 6.10 3.89 10.59 1.19 1.71 0.00
RI 8.35±1.22 11.12 6.02 5.10 14.61 -0.07 -1.06 0.00
E2 M 7.10±0.65 10.01 5.61 4.40 9.09 0.57 1.79 0.00
RI 7.43±0.61 9.87 5.81 4.06 8.18 0.24 1.72 0.00
E3 M 8.39±0.77 10.46 6.66 3.80 9.21 0.42 -0.32 0.00
RI 8.53±0.85 11.05 5.98 5.07 9.95 0.01 0.41 0.14
α (%) E1 M 8.12±0.73 11.71 6.81 4.90 9.00 1.91 5.37 0.00
RI 9.03±1.17 11.33 6.73 4.60 12.94 -0.17 -1.10 0.00
E2 M 7.92±0.72 10.24 6.20 4.04 9.09 0.60 0.36 0.00
RI 8.04±0.63 9.81 6.32 3.49 7.87 -0.03 -0.02 0.61
E3 M 9.28±0.89 11.93 6.81 5.12 9.59 0.22 -0.27 0.13
RI 9.43±0.94 12.60 6.97 5.63 9.98 0.29 0.47 0.14
β (%) E1 M 6.77±0.58 9.48 5.31 4.17 8.54 1.72 5.30 0.00
RI 6.92±0.61 8.33 4.97 3.36 8.78 0.07 0.08 0.02
E2 M 6.72±0.61 8.73 5.46 3.27 9.13 0.30 0.02 0.11
RI 6.97±0.60 8.95 5.49 3.46 8.63 0.46 0.80 0.01
E3 M 7.01±0.76 9.37 5.23 4.14 10.85 0.11 -0.05 0.75
RI 7.26±0.59 8.71 5.54 3.17 8.15 0.03 0.09 0.64
A3 (%) E1 M 5.50±0.38 6.48 4.25 2.23 6.92 -0.28 0.39 0.21
RI 5.06±0.55 6.25 3.71 2.54 10.89 -0.41 -0.24 0.00
E2 M 5.64±0.61 8.20 4.30 3.90 10.86 0.77 1.84 0.00
RI 5.35±0.54 7.02 3.16 3.86 10.11 -0.37 1.65 0.00
E3 M 5.42±0.62 7.15 3.45 3.70 11.35 -0.36 0.34 0.04
RI 5.60±0.72 7.56 3.94 3.62 12.85 0.33 -0.14 0.10

表2

RIL群体贮藏蛋白11S、7S组分相对含量基本统计量分析"

性状
Trait		 环境
E		 种植方式
Planting
pattern		 平均数±标准差
Mean±SD		 最大值
Max		 最小值
Min		 极差
Range		 变异系数
CV (%)		 偏度
Skew.		 峰度
Kurt.		 P (S-W)
Acid (%) E1 M 15.64±1.54 20.78 12.51 8.27 9.83 0.69 0.11 0.00
RI 14.38±1.52 17.99 8.70 9.29 10.57 -0.60 0.90 0.00
E2 M 15.86±1.92 21.60 11.33 10.27 12.10 0.50 0.26 0.01
RI 15.16±1.68 19.19 9.57 9.62 11.08 -0.07 0.07 0.43
E3 M 15.38±1.25 18.93 10.92 8.01 8.15 -0.29 0.98 0.04
RI 14.90±1.38 19.04 11.33 7.71 9.26 -0.19 -0.19 0.22
Basic (%) E1 M 10.36±1.95 16.15 6.46 9.69 18.79 0.56 -0.26 0.00
RI 10.18±2.83 20.32 5.35 14.97 27.78 0.84 1.50 0.00
E2 M 10.77±2.21 16.66 6.33 10.33 20.52 0.16 -0.62 0.03
RI 9.89±2.25 20.84 5.37 15.47 22.75 1.78 4.17 0.00
E3 M 8.82±2.31 23.59 4.81 18.78 26.24 1.90 7.63 0.00
RI 8.09±1.79 15.48 4.50 10.98 22.09 0.96 2.47 0.00
A5 (%) E1 M 4.66±1.53 11.43 2.88 8.55 32.86 1.95 4.15 0.00
RI 6.02±2.32 15.45 0.88 14.57 38.44 1.21 1.57 0.00
E2 M 5.45±1.63 11.61 3.01 8.60 29.84 1.22 1.56 0.00
RI 6.17±1.76 11.42 2.83 8.59 28.56 0.30 -0.21 0.00
E3 M 7.85±3.36 17.74 1.09 16.65 42.85 0.65 -0.13 0.00
RI 7.64±3.94 18.83 1.75 17.08 51.52 1.03 0.31 0.00

表3

大豆贮藏蛋白11S、7S组分相对含量的方差分析"

性状
Trait		 变异来源 Variability source
E P G E×P E×G P×G E×P×G
7S (%) 2719.17** 1130.16** 16.35** 198.05** 12.31** 15.24** 9.05**
11S (%) 232.79** 390.23** 9.34** 22.11** 9.01** 8.57** 6.90**
11S+7S (%) 577.47** 1.81 5.13** 90.72** 4.71** 4.39** 4.42**
11S/7S 3509.46** 3081.43** 34.63** 68.89** 29.83** 35.37** 20.09**
α' (%) 1534.95** 670.80** 10.08** 169.27** 6.65** 8.86** 6.62**
α (%) 1726.56** 423.291** 8.68** 181.27** 6.97** 7.90** 5.07**
β (%) 131.30** 162.88** 6.03** 4.20* 4.76** 4.04** 3.80**
A3 (%) 60.23** 91.30** 3.89** 93.41** 3.23** 2.47** 2.70**
Acid (%) 43.73** 324.46** 5.05** 26.16** 3.87** 3.09** 4.02**
Basic (%) 490.46** 116.33** 7.57** 14.65** 5.40** 5.82** 4.28**
A5 (%) 541.06** 97.84** 5.93** 52.02** 6.24** 5.73** 4.71**

表5

贮藏蛋白11S、7S组分相对含量相关QTL"

环境
Environ.		 种植方式
Planting pattern		 QTL 标记区间
Marker interval		 LOD 贡献率
PVE(%)		 加性效应
Add. effect		 置信区间
Confidence
interval
E1 M qAcid-6-131 Marker359561-Marker361355 5.63 8.06 0.53 130.5-131.5
qAcid-10-80 Marker584033-Marker584037 4.03 5.63 -0.46 79.5-80.5
qAcid-14-155 Marker713356-Marker710168 4.06 5.81 -0.47 153.5-158.0
RI qGpC-18-85 Marker1158309-Marker1157431 5.61 6.62 1.06 84.5-85.5
qBeta-19-5 Marker1175558-Marker1175562 4.03 9.01 -0.18 4.5-5.5
E2 M qBeta-1-135 Marker69562-Marker69768 3.21 7.49 -0.18 134.5-135.5
RI qAlpha-8-27 Marker460291-Marker460320 4.40 9.68 0.20 26.5-27.5
qBeta-1-13 Marker2130-Marker2656 3.22 7.14 -0.17 11.5-15.5
E3 M qAcid-4-67 Marker270606-Marker271196 3.68 8.20 -0.40 66.5-70.5
RI qGpC-20-110 Marker1242459-Marker1242433 3.65 8.60 0.82 109.5-110.5

表6

贮藏蛋白含量相关的候选基因分类"

分组
Group		 生物过程
Biological process		 基因ID (净作/套作)
Gene ID (Monoculture/Relay intercropping)		 注释信息
Annotation description
I 激素调节
Regulation of
hormone		 Glyma.01G179900; Glyma.14G193800, 060300, 195200, 195300, 196000, 196600, 197100, 197200, 197500, 197600, 198000, 198100, 198300, 198600, 199200, 200200, 202000, 202600, 202700, 203000 / Glyma.01G025300, 025400, 025500, 025600, 027400; Glyma.18G251800, 252200, 252300, 252400 脱落酸激活的信号通路、激素介导的信号通路、乙烯激活的信号通路、生长素反应、脱落酸反应
Abscisic acid-activated signaling pathway, hormone-mediated signaling pathway, ethylene-activated signaling pathway, response to auxin, response to abscisic acid
II 氨基酸合成
Amino acid
synthesis		 Glyma.01G180100; Glyma.04G211400; Glyma.06G243800; Glyma.14G195300, 196600, 197100, 198700, 202000, 202700, 203000, 203700 / Glyma.01G025600, 027100, 027400, 028100, 028700; Glyma.08G210500; Glyma.18G250600, 251800 蛋白丝氨酸/苏氨酸激酶活性、丝氨酸家族氨基酸代谢过程
Protein serine/threonine kinase activity, serine family amino acid metabolic process
III 蛋白质磷酸化
Protein
phosphorylation		 Glyma.06G243800; Glyma.14G195200, 196600, 202700, 203000 / Glyma.01G027100, 028000, 028700; Glyma.18G249400, 249700, 249900, 250500, 250600, 251800 蛋白质磷酸化、蛋白质丝氨酸/苏氨酸磷酸酶活性、蛋白质脱磷酸
Protein phosphorylation, protein serine/threonine phosphatase activity, protein dephosphorylation
IV 细胞器生物过程
Organelle
biological process		 Glyma.04G212200, 212400; Glyma.14G194300, 194500, 196600, 197600, 198000, 198100, 198600, 199000, 202000, 202400, 203000 / Glyma.01G025200, 026000, 026200, 026700, 028100; Glyma.18G248400, 251300, 251800, 253200 内质网、液泡、内质网膜
E ndoplasmic reticulum, vacuole, endoplasmic reticulum membrane
V 种子发育
Seed development		 Glyma.01G179800; Glyma.14G197100, 197500, 199000, 202100 / Glyma.01G025200, 025300, 025400, 026200, 027400; Glyma.18G249400, 249700, 249800, 250000, 250100, 250600, 250700, 251200, 251800 子叶发育、以种子休眠结束的胚胎发育
Cotyledon development, embryo development ending in seed dormancy

表7

3个环境6月至10月气象资料"

月
Month		 旬
Ten-day periods		 日平均温度
Daily mean temperature		 日最高温度
Daily maximum temperature		 日最低温度
Daily minimum temperature
E1 E2 E3 E1 E2 E3 E1 E2 E3
6月
June		 上旬 The first ten days 25.40 23.13 25.80 30.47 28.02 31.86 21.59 19.87 20.99
中旬 The middle ten days 23.94 22.68 27.40 28.07 26.92 32.54 21.12 19.82 22.43
下旬 The last ten days 25.77 24.24 25.90 30.39 29.45 30.41 22.08 20.61 22.53
7月
July		 上旬 The first ten days 27.83 26.09 27.20 33.93 31.38 31.26 23.67 22.73 23.57
中旬 The middle ten days 29.23 27.78 27.80 35.00 34.31 32.80 24.45 23.50 23.20
下旬 The last ten days 30.41 28.08 27.90 36.13 35.23 32.84 25.88 24.15 24.23
8月
August		 上旬 The first ten days 29.41 28.11 27.00 35.17 34.20 31.98 25.55 24.43 23.79
中旬 The middle ten days 27.71 26.10 30.40 32.25 31.20 35.66 24.23 23.24 26.33
下旬 The last ten days 26.16 24.36 28.20 29.99 28.63 33.05 23.40 21.88 24.72
9月
September		 上旬 The first ten days 23.95 22.54 23.10 27.66 27.38 27.12 21.31 19.81 20.43
中旬 The middle ten days 23.68 22.29 22.00 27.40 26.03 25.40 21.22 20.30 19.82
下旬 The last ten days 23.04 21.55 22.30 26.43 25.89 25.95 20.94 19.24 19.88
10月
October		 上旬 The first ten days 19.97 18.77 22.00 23.73 23.03 25.59 17.70 16.85 19.47
中旬 The middle ten days 17.07 15.75 18.30 19.41 18.36 20.94 15.65 14.28 16.55
下旬 The last ten days 16.02 14.97 17.70 18.60 17.70 20.75 14.33 13.58 15.76
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