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作物学报 ›› 2020, Vol. 46 ›› Issue (6): 832-843.doi: 10.3724/SP.J.1006.2020.94154

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

铝毒胁迫下甘蓝型油菜种子萌发期相关性状的QTL定位

王瑞莉,王刘艳,叶桑,郜欢欢,雷维,吴家怡,袁芳,孟丽姣,唐章林,李加纳,周清元(),崔翠()   

  1. 西南大学农学与生物科技学院, 重庆 400715
  • 收稿日期:2019-10-16 接受日期:2020-01-15 出版日期:2020-06-12 网络出版日期:2020-02-19
  • 通讯作者: 周清元,崔翠
  • 作者简介:E-mail: 1525731297@qq.com
  • 基金资助:
    国家重点研发计划项目(2018YFD0100500);国家现代农业产业技术体系建设专项(CARS-12);重庆市技术创新与应用发展项目(cstc2019jscx-msxmX0383)

QTL mapping of seed germination-related traits in Brassica napus L. under aluminum toxicity stress

WANG Rui-Li,WANG Liu-Yan,YE Sang,Gao Huan-Huan,LEI Wei,WU Jia-Yi,YUAN Fang,MENG Li-Jiao,TANG Zhang-Lin,LI Jia-Na,ZHOU Qing-Yuan(),CUI Cui()   

  1. College of Agronomy and Biotechnology, Southwestern University, Chongqing 400715, China
  • Received:2019-10-16 Accepted:2020-01-15 Published:2020-06-12 Published online:2020-02-19
  • Contact: Qing-Yuan ZHOU,Cui CUI
  • Supported by:
    National Key Research and Development Program of China(2018YFD0100500);China Agriculture Research System(CARS-12);Technological Innovation and Application Development in Chongqing(cstc2019jscx-msxmX0383)

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摘要：

随着土壤酸化的日益加重, 铝毒已成为影响作物种子萌发质量以及作物产量的重要胁迫因子之一。作物耐铝相关性状的QTL定位和候选基因筛选已有许多报道, 但铝胁迫下甘蓝型油菜萌发期相关性状的QTL定位报道较少。本文以80 μg mL -1的铝胁迫浓度处理重组自交系(10D130×中双11号)群体进行种子萌发试验, 处理3 d时调查发芽势, 7 d时调查发芽率, 测定其根长、芽长和干重, 并计算各性状相对值。基于6K SNP芯片, 结合高密度遗传连锁图谱对油菜萌发期的5个性状进行QTL定位, 共检测到23个QTL。其中与相对发芽势、相对发芽率、相对根长、相对芽长和相对干重相关的QTL分别有9个、1个、4个、5个和4个, 覆盖了A、C基因组, LOD值介于3.00~5.26, 可解释的表型变异为7.70%~13.10%。根据各QTL置信区间序列筛选, 与铝胁迫相关的候选基因共30个。ALMT1基因和MATE基因与有机酸的合成和分泌有关, 主要通过苹果酸、柠檬酸和草酸等有机酸的分泌来增强植物的耐铝性; STOP1基因、NAC基因和RAP2.4基因均属于转录激活因子, 通过诱导耐铝基因的表达增强植株的抗性; ABC转运蛋白、膜蛋白转运体、GDSL脂肪酶通过减少有毒物质在质膜上的积累或将有毒物质排出体外等途径增强植物的耐铝性; 过氧化物酶和细胞色素P450均属于氧化胁迫相关基因, 具有防止植物细胞氧化损伤、抵御逆境胁迫的功能; 另外, 还有部分编码逆境蛋白的基因, 均在各种胁迫反应中起重要作用。本研究的结果将为培育耐铝油菜品种及后续基因的功能研究提供理论依据。

关键词: 甘蓝型油菜, 萌发期, 单核苷酸多态性, 铝胁迫, QTL, 候选基因筛选

Abstract:

With the aggravation of soil acidification, aluminum toxicity has become one of the important stress factors that affect crop seed germination quality and crop yield. There have been many reports on QTL mapping and candidate gene screening for aluminum tolerance related traits in crops, but few reports on QTL mapping for aluminum tolerance during germination in Brassica napus L. The seed germination test was conducted on the recombinant inbred line population treated with 80 μg mL -1 aluminum solution and with distilled water as control. Germination potential was investigated at 3 d, germination rate was investigated at 7 d, root length, bud length and dry weight were measured, and the relative values of various characters were calculated. Based on 6K SNP chip and combined with high density genetic linkage map, we detected 23 QTLs for five traits in rape germination period. QTLs for relative germination potential, relative germination rate, relative root length, relative bud length and relative dry weight were 9, 1, 4, 5, and 4 respectively, covering A and C genomes, with LOD values from 3.00 to 5.26, and interpretable phenotypic variation from 7.70% to 13.10%. According to the confidence interval sequence of each QTL, 30 candidate genes related to aluminum stress were screened. ALMT1 gene and MATE gene are related to the synthesis and secretion of organic acids, which mainly enhance the aluminum tolerance of plants through the secretion of organic acids such as malic acid, citric acid and oxalic acid. STOP1 gene, NAC gene and RAP2.4 gene all belong to transcription activation factors, which enhance the resistance of plants by inducing the expression of aluminum tolerance genes. ABC transporter, membrane protein transporter, GDSL lipase enhance aluminum tolerance of plants by reducing accumulation of toxic substances on plasma membrane or discharging toxic substances out of body. Peroxidase and cytochrome P450 are genes related to oxidative stress, which have the functions of preventing oxidative damage of plant cells and resisting stress. At the same time, many genes encoding stress proteins were found, which play an important role in various stress reactions. The results of this study provide a theoretical basis for the cultivation of aluminum-tolerant rape varieties and the functional research of subsequent genes.

Key words: Brassica napus L., germination, single nucleotide polymorphism, aluminum stress, QTL, candidate gene

图1

不同浓度铝对油菜相对根长的影响"

表1

重组自交系群体及亲本油菜在铝胁迫环境下各性状的分布特征"

相关性状
Relative trait		 亲本 Parents RIL群体 RIL population
ZS11 10D130 均值Mean 极差Range 标准差SD 变异系CV(%)
相对发芽势 RGV 0.982 0.967 1.032** 1.667 0.230 22.3
相对发芽率 RGR 0.983 0.982 1.043** 1.030 0.140 13.4
相对根长 RRL 0.621 0.472 0.465** 0.733 0.146 31.3
相对芽长 RBL 0.955 0.987 0.994** 1.158 0.178 17.9
相对干重 RDW 0.987 0.990 0.920** 1.125 0.175 19.0

图2

最适铝胁迫处理下油菜品系各性状相对值的频率分布图 RGV: 相对发芽势; RGR: 相对发芽率; RRL: 相对根长; RBL: 相对芽长; RDW: 相对干重。"

表2

油菜种子铝胁迫处理后各性状间的相关性分析"

测定指标Index 相对发芽势RGV 相对发芽率RGR 相对根长RRL 相对芽长RBL 相对干重RDW
相对发芽势RGV 1
相对发芽率RGR 0.404** 1
相对根长RRL 0.101 0.053 1
相对芽长RBL -0.108 -0.109 0.062 1
相对干重RDW -0.148* -0.252** -0.075 -0.020 1

图3

甘蓝型油菜耐铝胁迫QTL在SNP连锁群上的分布情况 RGV: 相对发芽势; RGR: 相对发芽率; RRL: 相对根长; RBL: 相对芽长; RDW: 相对干重。"

表3

利用复合区间作图法检测到油菜萌发期在铝胁迫下的QTL"

性状
Trait		 QTL 标记区间
Marker interval		 LOD 表观贡献率
PVE (%)		 加性效应
Additive effect		 置信区间
Confidence interval (cM)
相对发芽势RGV qRGV-A01-1 AX-177912497-AX-179306831 4.33 10.90 -0.0282 37.410-43.975
qRGV-A01-2 AX-182171878-AX-182150271 3.90 9.90 -0.0217 141.657-155.683
qRGV-A03-1 AX-95507427-AX-177832788 4.20 10.60 0.0064 179.611-222.454
qRGV-A03-2 AX-95504605-AX-177831059 3.74 9.50 0.0033 93.552-97.113
qRGV-A03-3 AX-95681175-AX-95665748 3.23 8.20 -0.0236 121.496-124.157
qRGV-A08 AX-95506362-AX-182175648 5.26 13.10 -0.0402 8.146-12.834
qRGV-C01-1 AX-182087666-AX-95638042 3.43 8.70 -0.0123 160.226-163.199
qRGV-C01-2 AX-177910610-AX86223402 3.00 7.70 -0.0072 1.826-6.674
qRGV-C03 AX-95665064-AX-105309102 4.17 10.50 -0.0022 6.340-10.132
相对发芽率RGR qRGR-C04 AX-95636072-AX-182161180 3.12 8.00 0.0329 34.378-38.45
相对根长RRL qRRL-A03-1 AX-182087696-X-182147367 3.14 8.00 -0.0349 144.851-146.986
qRRL-A03-2 AX-95509347-AX-95509401 3.11 8.00 -0.0366 207.579-211.5
qRRL-A09 AX-95635531-AX-182177329 3.36 8.60 -0.0348 59.898-79.773
qRRL-C03 AX-95662770-AX-177910217 3.23 8.20 -0.0274 218.487-221.508
相对芽长RBL qRBL-A03 AX-95636441-AX-177911244 3.12 8.00 -0.0038 32.513-47.814
qRBL-A08-1 AX-95663434-AX-95506362 3.34 8.50 -0.0158 2.275-8.146
qRBL-A09-1 AX-177912380-AX-177829538 4.07 10.30 0.0216 178.878-188.216
qRBL-A09-2 AX-177832428-AX-182127697 3.53 9.00 0.0224 6.106-15.138
qRBL-C08 AX-95665611-AX-86227922 4.75 11.90 0.0115 94.661-132.074
相对干重RDW qRDW-A09-1 AX-182158057-AX-182154686 3.60 7.80 0.0196 127.938-128.931
qRDW-A09-2 AX-95682972-AX-182133711 3.05 9.10 0.0067 275.002-280.965
qRDW-A10-1 AX-182087443-AX-65636575 4.09 10.30 0.0050 129.25-131.783
qRDW-A10-2 AX-177829985-AX-182146546 3.90 9.90 0.0036 160.157-160.711

表4

甘蓝型油菜基因组中QTL置信区间候选基因与拟南芥逆境胁迫相关基因的比对"

性状
Trait		 物理区间
Physical interval		 甘蓝型油菜基因编号
Gene ID in B. napus		 基因登录号
Gene accession		 基因注释
Gene annotation		 参考文献
Reference
相对发芽势RGV 18835423-22795371 BnaA01g26740D AT3G18440 Aluminum-activated malate transporter 9 (ALMT9) [14]
20061150-26605700 BnaA03g57410D AT3G06130 Heavy metal-associated domain (HMA) [23]
BnaA03g39800D AT5G61240 Leucine-rich repeat (LRR) family protein [24]
14408144-15038093 BnaA03g03270D AT5G11250 Disease resistance protein (TIR-NBS-LRR class)
BnaA03g14650D AT2G31660 Super sensitive to ABA and drought2 (SAD2) [25]
11435749-11630658 BnaA03g13730D AT2G30140 UDP-Glycosyltransferase superfamily protein [26]
1012939-2022032 BnaA08g30510D AT4G19960 K+ uptake permease 9 (KUP9) [27]
BnaA08g19190D AT1G24620 EF hand calcium-binding protein family (CML25) [28]
BnaA08g26970D AT1G08430 Aluminum-activated malate transporter 1 (ALMT1)
676816-1129401 BnaC01g23380D AT3G51490 Metallothionein 3 (MT3) [29]
BnaC01g01650D AT4G37270 Response to toxic substance (HMA)
3925832-4284222 BnaC03g08920D AT5G18370 Disease resistance protein (TIR-NBS-LRR class) family [30]
相对发芽率RGR 3627529-9882343 BnaC04g12580D AT2G27780 Transcription factor IIS family protein [31]
相对根长RRL 17577932-2005963 BnaA03g35920D AT3G20750 Zinc ion binding (GATA) [32]
24406111-29734315 BnaA03g35900D AT5G26717 Putative membrane lipoprotein [33]
BnaA03g54580D AT1G34370 Sensitive to proton rhizotoxicity 1 (STOP1) [15]
866809-2227131 BnaA09g11920D AT1G64065 Late embryogenesis abundant (LEA) [34]
BnaA09g14610D AT3G43570 GDSL-like Lipase/Acylhydrolase superfamily protein [35]
51210223-55837858 BnaC03g04310D AT5G44050 MATE efflux family protein [16]
BnaC03g65810D AT4G34710 Response to osmotic stress (ADC2) [36]
相对芽长RBL 3249677-4604524 BnaA03g39360D AT5G02490 Heat shock protein 70 (Hsp 70) family protein [37]
598224-1012939 BnaA08g01480D AT1G52490 F-box associated interaction domain [38]
22081699-23394648 BnaA09g02290D AT3G28415 ABC transporter family protein [39]
BnaA09g30790D AT1G22220 F-box family protein
BnaA09g30810D AT1G22190 Response to toxic substance (RAP2.4) [40]
575326-705822 BnaA08g28220D AT1G05250 Peroxidase superfamily protein [41]
19580994-23049951 BnaC08g07240D AT1G33110 MATE efflux family protein
相对干重RDW 32408555-33029777 BnaA09g48330D AT1G49360 F-box family protein
16531371-16654826 BnaA10g24620D AT3G26180 Cytochrome P450 proteins [42]
BnaA10g25760D AT5G04410 Regulation of flavonoid biosynthetic process (NAC) [43]
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