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作物学报 ›› 2022, Vol. 48 ›› Issue (11): 2749-2764.doi: 10.3724/SP.J.1006.2022.14225

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

铝胁迫下甘蓝型油菜苗期光合相关性状的QTL定位及候选基因筛选

吴家怡(), 袁芳, 孟丽姣, 李晨阳, 史红松, 白岩松, 武晓如, 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:2021-12-01 接受日期:2022-02-25 出版日期:2022-11-12 网络出版日期:2022-03-31
  • 通讯作者: 周清元,崔翠
  • 作者简介:第一作者联系方式: E-mail: 2940116934@qq.com
  • 基金资助:
    本研究由财政部和农业农村部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 and candidate genes screening of photosynthesis-related traits in Brassica napus L. during seedling stage under aluminum stress

WU Jia-Yi(), YUAN Fang, MENG Li-Jiao, LI Chen-Yang, SHI Hong-Song, BAI Yan-Song, WU Xiao-Ru, LI Jia-Na, ZHOU Qing-Yuan*(), CUI Cui*()   

  1. College of Agronomy and Biotechnology, Southwestern University, Chongqing 400715, China
  • Received:2021-12-01 Accepted:2022-02-25 Published:2022-11-12 Published online:2022-03-31
  • Contact: ZHOU Qing-Yuan,CUI Cui
  • Supported by:
    The China Agriculture Research System of MOF and MARA(CARS-12);The Technological Innovation and the Application Development in Chongqing(cstc2019jscx-msxmX0383)

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摘要：

随着土壤酸化的加剧, 土壤中的Al3+已成为限制光合作用、影响作物生长和产量的主要因素之一。本文以2400 µmol L-1铝溶液处理由甘蓝型油菜品种中双11号及10D130构建的高世代重组自交系(RIL)群体为研究对象, 测定油菜铝毒胁迫下油菜苗期光合能力和叶绿素荧光相关性状, 计算其相对值(处理/对照)并定位其QTL, 根据QTL置信区间筛选合适的候选基因。研究结果表明, 光合能力性状间和叶绿素荧光参数间分别都存在着显著相关, 但是光合能力参数和叶绿素荧光参数之间相关性不显著; 除相对Tr外, 其余10个性状(相对Pn、相对Gs、相对Ci、相对ΦPSII、相对Fv/Fo、相对Fv'/Fm°、相对qP、相对ETR、相对NPQ和相对干重)共检测到16个与铝胁迫和光合相关性状QTL位点, 分别位于A03、A05、A06、A08、A09、A10、C02和C09染色体上, LOD值介于2.05~3.18, 能够解释的表型变异为8.1%~12.3%, 其中在A03染色体106.038~109.129 cM位置上, 相对Gs与相对Fv°/Fm°的位点完全重合, A06染色体上145.355~167.417 cM处检测到相对ΦPSII和相对ETR的QTL置信区间完全重合。筛选到47个可能与抗铝胁迫及光合相关的候选基因, 涉及到有机酸分泌、金属离子转运、激素调节、启动抗氧化保护以及叶绿体自身调节、磷元素摄入调节等方面。本研究为油菜铝抗性及相关光合基因功能挖掘和油菜耐铝高光合品种选育奠定基础。

关键词: 甘蓝型油菜, 铝胁迫, 光合性状, QTL, 候选基因筛选

Abstract:

Due to excessive soil acidification, aluminum in soil has become one of the main factors limiting crop growth and yield. In this study, a high generation of recombinant inbred line (RIL) population constructed from Brassica napus L. varieties Zhongshuang 11 (ZS11) and 10D130 treated with 2400 µmol L-1 aluminum solution was used as experimental materials. To explore the photosynthetic capacity and chlorophyll fluorescence related traits of rapeseed seedling stage under aluminum toxicity stress, we calculated their relative values (treatment/control) and mapped their QTLs, and screened suitable candidate genes according to the confidence interval of QTLs. The results showed that there was significantly correlation between photosynthetic capacity and chlorophyll fluorescence parameters, but there was no significant correlation between photosynthetic capacity and the chlorophyll fluorescence parameters. Relative Pn, relative Gs, relative Ci, relative ΦPSII, relative Fv/Fo, relative Fv′/Fm′, relative qP, relative ETR, relative NPQ, and relative dry weight were investigated and QTLs mapping were analyzed. A total of 16 QTLs related to aluminum stress and photosynthesis were detected. The locus was located on chromosomes A03, A05, A06, A08, A09, A10, C02, and C09, respectively. LOD value was 2.05-3.18, and the phenotypic variation explained was 8.1%-12.3%. The confidence intervals of QTLs for relative Gs and relative Fv'/Fm', relative ΦPSII and relative ETR were found completely overlap at 106.038-109.129 cM of chromosome A03, 145.355-167.417 cM of chromosome A06, respectively. 47 candidate genes that may be related to aluminum stress and photosynthesis were screened, involving organic acid secretion, metal ion transport, hormone regulation, initiation of antioxidant protection, chloroplast self-regulation, and phosphorus uptake regulation. This study lays the foundation for the function research of aluminum resistance and related photosynthetic genes and the breeding of aluminum tolerant and high photosynthetic efficiency rapeseed.

Key words: aluminum stress, photosynthetic traits, QTLs, candidate genes screening, Brassica napus L.

图1

铝胁迫对ZS11和10D130的净光合速率的影响 CK为对照; 800、1600、2400、3200和4000分别表示铝处理浓度为800、1600、2400、3200和4000 μmol L-1。柱上不同小写字母表示同一品种处理间差异显著(P < 0.05)。ZS11: 中双11号。"

表1

亲本及重组自交系群体11个性状的性状分析"

性状
Trait		 亲本 Parents RIL群体 RIL population 遗传率
Heritability (%)
ZS11 10D130 均值
Mean		 范围
Range		 极差
Range value		 标准差
SD		 变异系数
CV (%)
RPn 1.048 0.751 0.617** 0.285-1.589 1.304 0.249 40.357 95.4
RGs 1.040 0.985 0.472** 0.184-1.521 1.337 0.213 45.127 93.2
RCi 0.957 0.945 0.957** 0.827-1.106 0.279 0.054 5.643 81.8
RTr 1.860 0.989 0.537** 0.215-2.324 2.109 0.275 51.210 84.0
RΦPS 0.997 0.920 0.910** 0.531-1.322 0.791 0.114 12.527 90.7
Rfv 0.857 1.062 0.937** 0.681-1.114 0.433 0.069 7.364 82.4
Rfv° 1.006 0.971 0.943** 0.793-1.114 0.321 0.051 5.408 80.0
RqP 0.994 0.952 0.969** 0.706-1.293 0.587 0.081 8.359 87.0
RETR 0.997 0.920 0.909** 0.682-1.168 0.486 0.078 8.581 81.8
RNPQ 1.012 0.952 1.733** 0.952-3.512 2.560 0.498 28.736 89.2
RDW 0.869 1.439 0.551** 0.828-1.720 1.525 0.270 49.002 97.3

图2

铝处理下重组自交系各性状相对值频数分布图 缩写同表1。"

表2

重组自交系群体11个性状相关性分析"

测定指标Trait RPn RGs RCi RTr RΦPS Rfv Rfv' RqP RETR RNPQ
RGs 0.274**
RCi 0.036 0.118
RTr 0.272** 0.584** 0.272**
RΦPS -0.078 0.021 -0.034 0.031
Rfv -0.047 0.130 0.081 0.204* 0.188*
Rfv° -0.034 -0.061 -0.188* 0.001 0.427** 0.060
RqP -0.032 0.056 0.071 0.003 0.404** 0.201* 0.085
RETR -0.007 0.000 0.001 0.061 0.698** 0.268** 0.540** 0.484**
RNPQ -0.003 0.100 0.161 -0.030 -0.304** 0.012 -0.516** -0.095 -0.388**
RDW 0.197* 0.038 -0.077 0.087 0.075 -0.030 -0.093 0.125 0.052 0.011

表3

重组自交系群体10个性状的QTL定位"

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)
RPn qRPn-A10 AX-177828527-AX182175987 3.18 12.3 -0.0904037 9.474-39.386
RGs qRgs-A03-1 AX-95635223-AX-177830413 2.20 8.7 -0.0655227 96.668-100.169
qRgs-A03-2 AX-177913067-AX-95507652 2.11 8.3 -0.0643332 106.038-109.129
RCi qRCi-C02 AX-86224097-AX-179307085 2.05 8.1 -0.0157819 0.468-3.458
RΦPS qRΦPS-A06 AX-95503227-AX-177632560 2.84 11.0 0.0472636 145.355-167.417
Rfv qRfv-C09 AX-95506407-AX-177910786 2.37 9.3 -0.0296313 38.45-86.514
Rfv° qRfv'-A03 AX-95638185-AX-182141998 2.41 9.4 -0.0158728 103.15-118.351
qRfv'-A09-1 AX-177835940-AX177912292 2.36 9.2 -0.0166685 87.81-96.271
qRfv'-A09-2 AX-95506944-AX-182127697 2.19 8.6 0.0154628 15.138-15.609
RqP qRqP-A03 AX-95509347-AX-177828121 2.62 10.2 -0.0252692 198.519-228.654
RETR qRETR-A06 AX-95503227-AX-177632560 2.96 11.4 0.0314658 145.355-167.417
qRETR-A08 AX-95505568-AX-179307653 2.19 8.6 0.0234311 31.074-45.768
RNPQ qRNPQ-A09-1 AX-182127697-AX-177632723 2.65 10.3 -0.1639910 15.138-36.592
qRNPQ-A09-2 AX-177912943-AX-182155511 2.55 9.9 -0.1615910 0-9.664
RDW qRDW-A03 AX-95502575-AX-182145123 2.37 9.3 -0.0737873 54.689-75.861
qRDW-A05 AX-95665384-AX-95666447 2.78 10.8 -0.0638520 59.152-61.883

表4

铝胁迫及光合相关的候选基因"

性状
Trait		 物理区间
Physical
interval		 甘蓝型油菜
基因编号
Gene ID in
B. napus		 拟南芥
基因登录号
Gene
accession		 基因注释
Gene annotation
RPn 1006640-1008278 BnaA10g01990D AT1G03550 分泌载体膜家族蛋白
Secretory carrier membrane protein (SCAMP) family protein
1017170-1017425 BnaA10g02020D AT1G03600 PSB27
1159799-1160267 BnaA10g02210D AT1G03850 谷氧还家族蛋白
Glutaredoxin family protein
1416448-1417208 BnaA10g02700D AT1G04560 AWPM-19家族蛋白
AWPM-19-like family protein
1420412-1425818 BnaA10g02720D AT1G04580 醛氧化酶 4
Aldehyde oxidase 4 (AO4)
270316-271150 BnaA10g00540D AT1G01360 脱落酸受体调节成分1
Regulatory component of ABA receptor 1 (RCAR1)
2297229-2298311 BnaA10g04360D AT1G06680 光系统II亚基 P-1
Photosystem II subunit P-1
2428863-2433922 BnaA10g04550D AT1G07110 果糖-2,6-二磷酸酯酶
Fructose-2,6-bisphosphatase (F2KP)
4967798-4968762 BnaA10g06520D AT5G53160 脱落酸受体调节成分3
Regulatory components of ABA receptor 3 (RCAR3)
5777349-5778320 BnaA10g07350D AT5G54270 捕光色素叶绿素a/b结合蛋白 3
Light-harvesting chlorophyll a/b-binding protein 3 (LHCB3)
RGs 11450896-11455054 BnaA03g23880D AT4G09020 异淀粉酶3
Isoamylase 3 (ISA3)
11592964-11594227 BnaA03g24110D AT4G09670 氧化还原酶家族蛋白
Oxidoreductase family protein
11703183-11704457 BnaA03g24350D AT4G10340 捕光色素叶绿素a/b结合蛋白5
Light harvesting chloroplyll a/b-binding protein 5 (LHCB5)
性状
Trait		 物理区间
Physical
interval		 甘蓝型油菜
基因编号
Gene ID in
B. napus		 拟南芥
基因登录号
Gene
accession		 基因注释
Gene annotation
12051498-12054303 BnaA03g24930D AT4G12020 WRKY19
RCi 1050779-1052616 BnaC02g02350D AT5G05340 过氧化物酶超家族蛋白
Peroxidase superfamily protein
1312700-1315286 BnaC02g02870D AT5G03860 苹果酸合酶
Malate synthase (MLS)
1338228-1339653 BnaC02g02940D AT2G18960 H(+)-ATP酶
H(+)-ATPase 1 (HA1)
RΦPS 23332213-23333764 BnaA06g35540D AT5G47180 植物 VAMP (囊泡相关膜蛋白)家族蛋白
Plant VAMP (vesicle-associated membrane protein) family protein
23354413-23359357 BnaA06g35590D AT5G47110 叶绿素a/b结合家族蛋白
Chlorophyll a/b binding family protein
24314147-24316076 BnaA06g37230D AT4G38970 果糖二磷酸醛缩酶2
Fructose-bisphosphate aldolase 2 (FBA2)
24206087-24209026 BnaA06g37120D AT5G42480 叶绿体的积累和复制6
Accumulation and replication of chloroplasts 6 (ARC6)
Rfv 44394910-44396026 BnaC09g43010D AT3G58810 金属耐受蛋白A2
Metal tolerance protein A2 (MTPA2)
44536233-44537627 BnaC09g43140D AT5G14040 磷酸盐转运蛋白3
Phosphate transporter 3
46193139-46193697 BnaC09g46220D AT3G21490 重金属转运/解毒超家族蛋白
Heavy metal transport/detoxification superfamily protein
48190280-48191717 BnaC09g50250D AT5G05000 叶绿体外膜蛋白转运机器的构件蛋白34
Translocon at the outer envelope membrane of chloroplasts 34 (TOC34)
Rfv° 13146862-13148728 BnaA03g26740D AT4G00910 铝活化的苹果酸盐转运家族蛋白
Aluminium activated malate transporter family protein
4120082-4121722 BnaA09g08390D AT3G32980 过氧化物酶超家族蛋白
Peroxidase superfamily protein
4519220-4519695 BnaA09g09080D AT2G34430 捕光叶绿素蛋白复合物II亚基
Light-harvesting chlorophyll-protein complex II subunit B1 (LHB1B1)
RqP 23778823-23781902 BnaA03g46370D AT4G23980 生长素应答因子9
Auxin response factor 9 (ARF9)
27082166-27084124 BnaA03g52000D AT4G32320 抗坏血酸过氧化物酶6
Ascorbate peroxidase 6 (APX6)
27439846-27442123 BnaA03g52600D AT5G51470 生长素应答GH3家族蛋白
Auxin-responsive GH3 family protein
27829209-27831558 BnaA03g53180D AT4G35090 过氧化氢酶2
Catalase 2 (CAT2)
RETR 23332213-23333764 BnaA06g35540D AT5G47180 植物 VAMP (囊泡相关膜蛋白)家族蛋白
Plant VAMP (vesicle-associated membrane protein) family protein
23354413-23359357 BnaA06g35590D AT5G47110 叶绿素a/b结合家族蛋白
Chlorophyll a/b binding family protein
24206087-24209026 BnaA06g37120D AT5G42480 叶绿体的积累和复制 6
Accumulation and replication of chloroplasts 6 (ARC6)
24314147-24316076 BnaA06g37230D AT4G38970 果糖二磷酸醛缩酶2
Fructose-bisphosphate aldolase 2 (FBA2)
性状
Trait		 物理区间
Physical
interval		 甘蓝型油菜
基因编号
Gene ID in
B. napus		 拟南芥
基因登录号
Gene
accession		 基因注释
Gene annotation
11764739-11766413 BnaA08g13620D AT4G28610 磷饥饿响应1
Phosphate starvation response 1 (PHR1)
RNPQ 525671-526016 BnaA09g00850D AT2G02930 谷胱甘肽S-转移酶 F3
Glutathione S-transferase F3 (GSTF3)
530729-531954 BnaA09g00880D AT4G02530 叶绿体类囊体腔蛋白
Chloroplast thylakoid lumen protein
607437-608064 BnaA09g01080D AT4G02770 光系统I亚基D-1
Photosystem I subunit D-1 (PSAD-1)
392076-395981 BnaA09g00710D AT4G02280 蔗糖合酶3
Sucrose synthase 3 (SUS3)
775432-776934 BnaA09g01470D AT4G03280 光合电子转移C
Photosynthetic electron transfer C (PETC)
RDW 7333238-7334039 BnaA03g15830D AT2G34430 捕光叶绿素蛋白复合物II亚基
Light-harvesting chlorophyll-protein complex II subunit B1 (LHB1B1)
5050885-5053547 BnaA03g11260D AT5G55700 β-淀粉酶4
Beta-amylase 4 (BAM4)
5090429-5091422 BnaA03g11320D AT5G55560 蛋白激酶超家族蛋白
Protein kinase superfamily protein
6332750-6334339 BnaA03g13820D AT2G30250 WRKY DNA结合蛋白25
WRKY DNA-binding protein 25 (WRKY25)
7108373-7109471 BnaA03g15320D AT2G03610 F-box家族蛋白
F-box family protein

图3

甘蓝型油菜10个相对性状QTL在连锁群上分布图 缩写同表1。"

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