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作物学报

作物学报 ›› 2024, Vol. 50 ›› Issue (1): 55-66.doi: 10.3724/SP.J.1006.2024.33011

所属专题： 玉米：遗传育种·种质资源·分子遗传学

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

控制玉米株高基因PHR1的基因克隆

杨晨曦1(), 周文期1,2,*(), 周香艳1,*(), 刘忠祥2, 周玉乾2, 刘芥杉1, 杨彦忠2, 何海军2, 王晓娟2, 连晓荣2, 李永生2   

  1. 1甘肃农业大学生命科学技术学院, 甘肃兰州, 730070
    2甘肃省农业科学院作物研究所, 甘肃兰州730070
  • 收稿日期:2023-02-24 接受日期:2023-06-29 出版日期:2024-01-12 网络出版日期:2023-07-25
  • 通讯作者: *周文期, E-mail: zhouwenqi850202@163.com; 周香艳, E-mail: zhouxy@gsau.edu.cn
  • 作者简介:E-mail: yangchenxi@qq.com
  • 基金资助:
    国家自然科学基金项目(32160490);国家自然科学基金项目(31960443);甘肃省重大专项(21ZD11NA005);甘肃省重大专项(21ZD10NF003);甘肃省农业科学院生物育种专项(2022GAAS04);甘肃农业大学青年导师基金项目(GSAU-QDFC-2021-14);中国工程院战略研究与咨询项目: 特色粮油作物优异基因精准鉴定及新种质创制(2021-DFZD-21-3)

Mapping and cloning of plant height gene PHR1 in maize

YANG Chen-Xi1(), ZHOU Wen-Qi1,2,*(), ZHOU Xiang-Yan1,*(), LIU Zhong-Xiang2, ZHOU Yu-Qian2, LIU Jie-Shan1, YANG Yan-Zhong2, HE Hai-Jun2, WANG Xiao-Juan2, LIAN Xiao-Rong2, LI Yong-Sheng2   

  1. 1College of Life Science and Technology, Gansu Agricultural University, Gansu 730070, Lanzhou, China
    2Crop Research Institute, Gansu Academy of Agriculture Sciences, Gansu 730070, Lanzhou, China
  • Received:2023-02-24 Accepted:2023-06-29 Published:2024-01-12 Published online:2023-07-25
  • Contact: *E-mail: zhouwenqi850202@163.com; E-mail: zhouxy@gsau.edu.cn
  • Supported by:
    National Natural Science Foundation of China(32160490);National Natural Science Foundation of China(31960443);Major Project of Gansu Province(21ZD11NA005);Major Project of Gansu Province(21ZD10NF003);Biological Breeding Project of Gansu Academy of Agricultural Sciences(2022GAAS04);Youth Mentor Fund Project of Gansu Agricultural University(GSAU-QDFC-2021-14);Strategic Research and Consulting Project of Chinese Academy of Engineering(2021-DFZD-21-3)

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摘要：

株高属于玉米理想株型育种的一个重要指标, 不但影响玉米机械化收获, 更与玉米的倒伏性和生物产量密切相关。本研究以低剂量快中子(4.19 Gy)辐照诱变玉米自交系KWS39获得的矮秆低穗位突变体为研究对象, 该突变体命名为plant height reducing mutant-1 (phr-1), 开展了表型性状的田间调查分析, 并利用phr-1×B73获得的F2分离群体, 借助极端性状混池测序分析法(BSA-seq)及目标区段重组交换鉴定的方法, 基于B73参考基因组对目标区段内的基因进行挖掘和功能注释, 定位候选基因。研究结果表明, 在1号染色体Bin1.06 区间可能存在变异位点, 进而利用大的分离群体结合目标区段多态性标记开发, 将目标区段精细定位分子标记到Umc1122和Umc1583a两个标记之间约600 kb区间, 该区段内存在一个控制株高的已知基因Brachytic2 (BR2), BR2编码一个调控玉米茎秆中生长素极性运输的糖蛋白。候选基因测序结果表明, phr-1是BR2基因在第4个外显子处插入了165 bp的序列, 导致第547位氨基酸变为终止子, 蛋白翻译提前终止。phr-1的基因突变位点和变异方式与已报道的br2-1单个碱基发生变异位点完全不同, 通过等位杂交实验证明了phr-1突变体就是br2-1的一个新等位突变体, 候选基因就是BR2基因。本研究为玉米BR2基因在玉米株高遗传改良中提供了新的种质资源。

关键词: 玉米, 株高, 穗位高, BSA混池测序, 基因定位, 功能分析

Abstract:

Plant height is an important index for the ideal maize breeding, which not only affects the mechanical harvest of maize, but also closely relates to the lodging resistance and the biological yield of maize. In this study, a plant height reducing mutant-1 (phr-1) was obtained by using low-dose fast neutron (4.19 Gy) irradiation to mutate the maize inbred line KWS39 at the low ear position. Phenotypic traits were investigated and analyzed in the field, and candidate genes were identified by mining and functional annotation of genes in the target region based on the B73 reference genome using the extreme trait pool sequencing analysis (BSA-seq) and the method of target region recombination exchange identification with F2 segregation population of phr-1×B73. These results indicated that there might be a variation site in Bin1.06 interval on chromosome 1, and the target region was precisely located between two markers, Umc1122 and Umc1583a, with a 600 kb interval by using the large segregation population and polymorphic markers. Brachytic2 (BR2), encoding a sugar protein regulating the polar transport of auxin in maize stems, is a known gene controlling plant height in this region. The sequencing of candidate genes revealed that phr-1 was a new allelic mutation of br2-1, with a 165 bp insertion in the fourth exon of the BR2 gene resulting in the amino acid at position 547 changing to a stop codon and premature protein translation termination. The mutation site and variation mode of phr-1 were completely different from those of br2-1 single base mutation site. The allelic hybridization experiment confirmed that the PHR1 candidate gene is a new allelic mutation of the BR2 gene. This study provides new germplasm resources for maize BR2 gene in plant height genetic improvement.

Key words: maize, plant height, ear height, the bulked segregant analysis, map-based localization, function analysis

表1

本研究所用到的引物"

引物名称
Primer name		 正向序列
Forward sequence (5′-3′)		 反向序列
Reverse sequence (5′-3′)
GAPDH CCATCATGCCACACAGAAAAC AGGAACACGGAAGGACATACCAG
BR2-Q1F/Q1R TGAGCAACTCCAGCTCTTAACCA GCGAACTGGACATCCTCAGATTA
BR2-Z1F/Z1R ATTCACGCAGAGCAGAAGAGC GCATAAGGTTGGACAGGGAAAG
BR2-H1F/H1R AGGTTGGGGAGCGCGGCCTGCAG ACCCATCCATCCATCCATTCCGTCCT
RT-Br2-F1/R1 ACGAGCATCAGGGAGAACC GCTCCCCAACCTGCGTGTCG

图1

野生型KWS39和突变体phr-1植株表型 A, B: KWS39和phr-1株型比较, phr-1比KWS39株型矮小; 标尺为10 cm。C: PH和EH统计, t检测; D: KWS39和phr-1节间比较; E: KWS39和phr-1节间数相同, 共11节, 节间从根到尖用1, 2, 3, …, 9, 10, 11表示, 但phr-1每一节之间的长度比对照减小, 标尺为10 cm。"

图2

野生型KWS39和突变体phr-1植株表型 A: KWS39和phr-1萌芽5 d, 株高及根长无明显差异; B: 从左往右分别是倒三叶、倒二叶和剑叶; C: KWS39和phr-1根系发育正常, phr-1根系更为发达; D: phr-1和KWS39套袋自交的果穗性状; E: KWS39和phr-1籽粒、百粒重都无显著变化。标尺均为10 cm。"

表2

突变体phr-1和野生型KWS39农艺性状分析"

类型
Type		 性状
Trait		 野生型KWS39
Wild type KWS39		 突变体phr-1
Mutant phr-1		 P-value N
株型Plant architecture 株高Plant height (cm) 182.8±11.0 93.3±2.3** 3.88E-09 15
穗位高Ear height (cm) 66.4±17.3 23.2±1.7** 0.000106 15
雄穗Tassel 雄穗分支数 Tassel branch number 5.1±1.4 4.7±1.6 0.582953 15
雄穗长Tassel length (cm) 31.65±2.3 27.68±1.9* 0.031581 15
雌穗Ear 穗长Ear length (cm) 13.81±1.3 13.08±0.5 0.447437 15
穗粗Ear diameter 3.97±0.1 3.7±0.2 0.094702 15
穗行数 Kernel row number 14.4±0.8 13.4±0.9 0.052177 15
行粒数 Grains row number 24.1±4.1 22.6±4.6 0.057321 15
秃尖长Bald tip length (cm) 1.5±0.6 1.31±0.4 0.078416 15
叶型Blade profile 倒3叶长Inverted 3 leaves long (cm) 47.25±3.8 44.3±3.6* 0.045242 15
倒3叶宽Inverted 3 leaves wide (cm) 6.61±0.7 6.36±0.6 0.486148 15

表3

F2分离群体的卡方测验"

群体
Population		 观察值Observations 期望值Expectations 卡方检验
Chi-square
野生型Wild type 突变体Mutant 野生型Wild type 突变体Mutant
phr-1×B73 F2 762 238 760 240 0.88

图3

KWS39、phr-1、杂交F1和F2矮秆植株表型 标尺为10 cm。"

图4

phr-1候选基因的连锁定位图 A: 黑色点和灰色点为ΔSNP-index; B: 蓝色线为ΔSNP-index划窗后的拟合线; C: 橙黄色线为95%置信线; D: 红色线为99%置信线。"

表4

初步定位标记与表型的交换频率"

标记名称
Molecular marker		 交换频率
Crossing-over value
umc2235 0.38
umc1035 0.29
umc2560 0.17
umc2569 0.15
umc1122 0.08
umc1583a 0.11
umc1278 0.21
umc2387 0.36

图5

BR2转录本在KWS39和phr-1突变体中的序列分析"

图6

等位鉴定突变体F1代表型一致 A: B73与突变体br2-1、phr-1和(bra2-1× phr-1) F1的植株表型; B: B73与野生型KWS39、突变体phr-1和(bra2-1×phr-1) F1植株表型。标尺为10 cm。"

图7

BR2基因在4个玉米自交系中的相对表达 数据经过t检测, **表示差异极显著(P < 0.01)。"

表5

BR2在不同物种中的基因编号及功能注释"

物种
Species		 同源基因
Orthologous gene		 功能预测
Putative function
玉米Zea mays GRMZM2G315375 PGP1
拟南芥Arabidopsis thaliana AT1G10680 P-glycoprotein 10
水稻Oryza sativa LOC_Os02g46680 Multidrug resistance protein, putative, expressed
白杨树Populus trichocarpa POPTR_0001s02200 PGP2; ATPase, coupled to transmembrane movement of substances
高粱Sorghum bicolor Sb04g023730 P-glycoprotein 1
葡萄Vitis vinifera GSVIVG0009946001 Multidrug resistance protein 1, 2

图8

ZmBR2同源蛋白氨基酸序列比对及系统进化分析 A: ZmBR2同源蛋白氨基酸序列比对; B: ZmBR2同源树; C: ZmBR2系统发育树。"

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