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作物学报 ›› 2024, Vol. 50 ›› Issue (2): 403-413.doi: 10.3724/SP.J.1006.2024.31016

所属专题： 小麦：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

小麦TabHLH112-2B基因克隆及每穗小穗数相关功能标记开发

范子培1,2(), 李龙2, 史雨刚1, 孙黛珍1,*(), 李超男2,*(), 景蕊莲2   

  1. 1山西农业大学农学院, 山西太谷 030801
    2中国农业科学院作物科学研究所, 北京 100081
  • 收稿日期:2023-02-27 接受日期:2023-05-24 出版日期:2024-02-12 网络出版日期:2023-06-20
  • 通讯作者: *孙黛珍, E-mail: sdz64@126.com;李超男, E-mail: lichaonan@caas.cn
  • 作者简介:E-mail: fan20202090@163.com
  • 基金资助:
    国家重点研发计划项目(2022YFD1200201);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(小麦, CARS-03-5)

Cloning of TabHLH112-2B gene and development of its functional marker associated with the number of spikelet per spike in wheat

FAN Zi-Pei1,2(), LI Long2, SHI Yu-Gang1, SUN Dai-Zhen1,*(), LI Chao-Nan2,*(), JING Rui-Lian2   

  1. 1College of Agriculture, Shanxi Agricultural University, Taigu 030801, Shanxi, China
    2Institute of Crop Sciences, Chinese Academy of Agricultural Sciences, Beijing 100081, China
  • Received:2023-02-27 Accepted:2023-05-24 Published:2024-02-12 Published online:2023-06-20
  • Contact: *E-mail: sdz64@126.com;lichaonan@caas.cn
  • Supported by:
    National Key Research and Development Program of China(2022YFD1200201);China Agriculture Research System of MOF and MARA(Wheat, CARS-03-5)

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摘要：

bHLH (basic Helix-Loop-Helix)转录因子在植物生长发育过程中发挥着重要作用。本研究克隆了小麦TabHLH112-2B, 该基因由7个外显子和6个内含子构成, 编码444个氨基酸, 在315~364位氨基酸处具有一个典型的HLH保守结构域。组织表达模式分析表明TabHLH112-2B在小麦幼苗期、拔节期、抽穗期和开花期的各个组织中均有表达, 其中在叶和根中表达量较高。顺式作用元件分析发现TabHLH112-2B启动子区含有植物激素应答、胁迫响应、与分生组织发育相关的多种顺式作用元件, qRT-PCR结果显示其表达响应ABA、IAA、MeJA等植物激素和干旱、高盐、低温、高温等胁迫处理。基因组序列多态性分析检测到启动子区域的2个SNP, 分为2种单倍型。根据SNP-682开发分子标记并进行关联分析, 发现该标记与干旱、高温等多种环境下的每穗小穗数显著相关, Hap-2B-2为每穗小穗数多的优异单倍型。研究结果为培育高产广适小麦新品种的分子标记辅助育种提供了优异基因资源和技术支撑。

关键词: 小麦, TabHLH112-2B, 分子标记, 关联分析, 每穗小穗数, 优异单倍型

Abstract:

The bHLH (basic Helix-Loop-Helix) transcription factor plays an important role in plant growth and development. In this study, wheat gene TabHLH112-2B was cloned, which consists of seven exons and six introns, encoding 444 amino acids, and has a typical HLH conserved domain at 315-364 amino acids. The tissue expression pattern analysis showed that TabHLH112-2B was expressed in all tissues at seedling stage, jointing stage, heading stage, and flowering stages. Among them, the relative expression levels in leaves and roots were higher. The cis-acting element analysis showed that the promoter region of TabHLH112-2B contained a variety of cis-acting elements related to plant hormone responses, stress responses, and meristem development. The qRT-PCR exhibited that the relative expression level of TabHLH112-2B was responsive to plant hormones (such as ABA, IAA, MeJA) and abiotic stresses (such as drought, salt, low and high temperatures). Two SNPs were detected in its promoter region by genomic sequence polymorphism, which were classified into two haplotypes. A molecular marker was developed based on SNP-682, and association analysis showed that the marker was significantly correlated with the number of spikelet per spike in various environments such as drought and high temperature. Hap-2B-2 was a favorable haplotype with more spikelets per spike. These results of this study provide the valuable genetic resources and technical support for molecular marker-assisted breeding of wheat varieties with high yield and wide adaptability.

Key words: wheat, TabHLH112-2B, molecular marker, association analysis, the number of spikelet per spike, favorable haplotype

表1

生物信息学分析所用软件的网址列表"

软件
Software		 用途
Function		 网址
Website
SMART 结构域分析
Domain analysis		 http://smart.embl.de/
ProtParam 蛋白理化性质分析
Physical and chemical properties		 https://web.expasy.org/protparam/
ExPASy 蛋白亲疏水性分析
Hydrophilicity analysis		 https://web.expasy.org/protscale/SWISS-MODEL
SignalP-5.0 蛋白信号肽预测
Signal peptide prediction		 https://services.healthtech.dtu.dk/service.php?SignalP-5.0
NetPhos-3.1 磷酸化位点分析
Phosphorylation site analysis		 https://services.healthtech.dtu.dk/service.php?NetPhos-3.1
SOPMA 蛋白二级结构预测
Protein secondary structure		 https://npsa-prabi.ibcp.fr/cgi-bin/npsa_automat.pl?page=npsa_sopma.html
phyre2 蛋白三级结构预测
Protein tertiary structure		 http://www.sbg.bio.ic.ac.uk/phyre2/html/page.cgi?id=index
PlantCARE 顺式作用元件分析
Cis-element analysis		 https://bioinformatics.psb.ugent.be/webtools/plantcare/html/

表2

本实验所用引物"

引物名称
Primer name		 上游引物序列
Forward sequence (5'-3')		 下游引物序列
Reverse sequence (5'-3')
TabHLH112-2B ACACTACCCCTCATCTTAGCTCC CTATGACATAATTCAGCCCTCTCA
TabHLH112-pro-2B GAGGCGACTATAGTACACGCAAA CGAAGGGTAAACTGATCCAGC
TabHLH112-2B-qRT CGGCTGAGCAGACTAGCGT AGACGAGCTGTTGGAGTGCC
TaTUB CGTGCTGTCTTTGTAGATCTCG GACCAGTGCAGTTGTCTGAAAG
TabHLH112-2B-BamH I GACAATCGAGTTAAAGAAATTAGGA TCTGTATGAATCTCTGACCGCA

图1

TabHLH112-2B基因序列和蛋白结构预测分析 A: 基因结构; B: 蛋白结构域; C: 蛋白的二级结构; D: 蛋白的三级结构。"

图2

TabHLH112-2B蛋白性质和磷酸化位点预测 A: 蛋白的氨基酸组成。A: 丙氨酸; R: 精氨酸; N: 天冬酰胺; D: 天冬氨酸; C: 半胱氨酸; Q: 谷氨酰胺; E: 谷氨酸; G: 甘氨酸; H: 组氨酸; I: 异亮氨酸; L: 亮氨酸; K: 赖氨酸; M: 蛋氨酸; F: 苯丙氨酸; P: 脯氨酸; S: 丝氨酸; T: 苏氨酸; W: 色氨酸; Y: 酪氨酸; V: 缬氨酸。B: 蛋白的亲/疏水性分析。C: 蛋白的磷酸化位点预测。D: 磷酸化预测值高于0.99的氨基酸位点。"

图3

小麦TabHLH112-2B在不同发育时期的组织表达谱 TabHLH112-2B在A: 幼苗期; B: 拔节期; C: 抽穗期和D: 开花期不同组织中的表达模式。L: 叶; R: 根; RB: 根基; S: 穗; YL: 心叶; P: 穗下节; PN: 倒二节; AN: 倒三节; FL: 旗叶; 误差线代表±标准差。不同的小写字母表示在P < 0.05水平上差异显著。"

图4

小麦幼苗根中TabHLH112-2B在不同激素处理下的表达谱 A: TabHLH112-2B启动子区顺式作用元件预测; B: 50 μmol L-1 ABA处理; C: 0.1 mmol L-1 IAA处理; D: 0.1 μmol L-1 MeJA处理2周龄幼苗根中TabHLH112-2B表达水平。误差线代表±标准差。*: P < 0.05; **: P < 0.01; ***: P < 0.001。"

图5

不同胁迫处理下小麦幼苗中TabHLH112-2B的表达谱 A: 16.1% PEG; B: 250 mmol L-1 NaCl; C: 低温胁迫; D: 高温胁迫处理下TabHLH112-2B的表达水平。误差线代表±标准差。*: P < 0.05; **: P < 0.01; ***: P < 0.001。"

图6

TabHLH112-2B的序列多态性及其dCAPS分子标记 A: TabHLH112-2B启动子区SNP位点与分子标记示意图, 红框表示酶切位点, 红色字母表示变异位点, 红点表示错配碱基。B: 2种单倍型的琼脂糖凝胶电泳图。M: 100 bp DNA ladder; C: Hap-2B-1; T: Hap-2B-2。"

表3

TabHLH112-2B单倍型与16种环境下每穗小穗数的关联分析结果"

环境
Environment		 P值
P-value		 环境
Environment		 P值
P-value
E1 0.011228526 E9 0.000000386
E2 0.113354414 E10 0.000000934
E3 0.048992919 E11 0.008263578
E4 0.031388574 E12 0.098658098
E5 0.002171438 E13 0.016524218
E6 0.000019440 E14 0.000638932
E7 0.000003438 E15 0.000032654
E8 0.000227778 E16 0.000022470

图7

TabHLH112-2B单倍型与每穗小穗数的关联分析结果 E1’E16缩写同表3。误差线代表±标准差。*: P < 0.05; **: P < 0.01; ***: P < 0.001。"

图8

TabHLH112-2B优异单倍型在我国小麦育种历史中的应用 TabHLH112-2B两种单倍型在我国十大麦区地方品种(A)和现代育成品种(B)中的分布频率。I: 北方冬麦区; II: 黄淮冬麦区; III: 长江中下游麦区; IV: 西南冬麦区; V: 华南冬麦区; VI: 东北春麦区; VII: 北部春麦区; VIII: 西北春麦区; IX: 青藏春冬麦区; X: 新疆冬春麦区。C: 不同年代品种在2005-2006 (2005), 2010-2011 (2010)两个生长季的每穗小穗数。D: 不同年代品种中2种单倍型的频率。"

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