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作物学报 ›› 2023, Vol. 49 ›› Issue (4): 966-977.doi: 10.3724/SP.J.1006.2023.21023

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

小麦幼苗活性LTR反转录转座子筛选及其对非生物胁迫的响应

周宾寒1(), 杨竹1, 王书平1, 方正武1, 胡赞民2, 徐兆师3, 张迎新1,*()   

  1. 1长江大学农学院/主要粮食作物产业化湖北省协同创新中心, 湖北荆州 434025
    2中国科学院遗传与发育生物学研究所, 北京 100101
    3中国农业科学院作物科学研究所/农作物基因资源与基因改良国家重大科学工程/农业部麦类生物学与遗传育种重点实验室, 北京 100081
  • 收稿日期:2022-03-29 接受日期:2022-07-21 出版日期:2023-04-12 网络出版日期:2022-08-22
  • 通讯作者: *张迎新, E-mail: zhangyingxin1985@126.com
  • 作者简介:E-mail: z18171732900@126.com
  • 基金资助:
    国家重点研发计划项目(2017YFD0100800);国家转基因生物新品种培育重大专项(2018ZX0800909B);湖北省重点研发计划项目(2022BBA0035);湖北省科学技术重大创新专项(2018ABA085)

Screening of active LTR retrotransposons in wheat (Triticum aestivum L.) seedlings and analysis of their responses to abiotic stresses

ZHOU Bin-Han1(), YANG Zhu1, WANG Shu-Ping1, FANG Zheng-Wu1, HU Zan-Min2, XU Zhao-Shi3, ZHANG Ying-Xin1,*()   

  1. 1College of Agriculture, Yangtze University, Hubei Collaborative Innovation Center for Grain Industry, Jingzhou 434025, Hubei, China
    2Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing 100101, China
    3Institute of Crop Sciences, Chinese Academy of Agricultural Sciences/National Key Facility for Crop Gene Resources and Genetic Improvement/Key Laboratory of Biology and Genetic Improvement of Triticeae Crops, Ministry of Agriculture, Beijing 100081, China
  • Received:2022-03-29 Accepted:2022-07-21 Published:2023-04-12 Published online:2022-08-22
  • Contact: *E-mail: zhangyingxin1985@126.com
  • Supported by:
    National Key Research and Development Programs of China(2017YFD0100800);National Major Project for Developing New GM Crops(2018ZX0800909B);Key Research and Development Program of Hubei Province(2022BBA0035);Major Project for Special Technology Innovation of Hubei Province(2018ABA085)

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摘要：

LTR (长末端重复, long terminal repeat)反转录转座子占小麦基因组的60%以上, 筛选小麦基因组中具有转座活性的LTR反转录转座子, 并分析其在非生物胁迫下的响应, 对研究反转录转座子在小麦抗逆境胁迫中的作用具有重要意义。本研究通过生物信息学分析, 从转座子数据库(TREP database)中筛选出4个具有完整结构的LTR反转录转座子Fatima、Wis、Angela和Babara; 同时利用实时荧光定量PCR (qRT-PCR)、甲基化特异PCR (methylmion specific PCR, MSP)和转座子展示(transposon display, TD)技术分别分析了它们在盐、ABA、H2O2和干旱等处理的小麦幼苗期(二叶一心)叶和根中的表达水平、甲基化水平和转座活性变化。结果表明, 这4个反转录转座子在正常条件下均存在基础水平的转录, 并且能够响应上述4种胁迫而发生转录水平的变化, 且在相同胁迫条件下表达水平变化趋势一致。Fatima、Angela和Babara在非生物胁迫处理下表达水平的提高与其甲基化水平的降低有关, Wis则相反。反转录转座子LTR序列含有胁迫响应顺式作用元件, 但在非生物胁迫条件下顺式作用元件对这4个反转录转座子的调控作用不显著。与叶相比, 这4个反转录转座子在根中对胁迫的响应程度更高, 且在盐和ABA处理下转座活性更强。本研究将有助于进一步揭示LTR反转录转座子对非生物胁迫的响应规律, 为进一步研究利用反转录转座子进行小麦抗逆育种的遗传改良积累资料。

关键词: 小麦, 非生物胁迫, LTR反转录转座子, 相对表达水平, 转座

Abstract:

LTR (long terminal repeat) retrotransposable elements account for more than 60% of rye genome. It is of great significance to screen active LTR retrotransposons from wheat genome and analyze their transposition, the relative expression levels, and methylation levels under abiotic stress in exploring the role of retrotransposable elements in improving wheat anti-stress ability. Four active LTR retrotransposons (Fatima, Wis, Angela, and Babara) with complete structure were selected from the TREP database by bioinformatics analysis. The relative expression levels, methylation dynamics, and transposition activity of the four LTR retrotransposons were then analyzed at seedling stage (two leaves with one heart) leaves and roots of the stress-treated (NaCl, ABA, H2O2, and PEG) wheat seedlings by qRT-PCR, methylation-specific polymerase chain reaction (MSP), and transposon display method. The results suggested that the four retrotransposons had basic transcription activity under normal conditions, which could be changed under stress conditions, and their variation tendency was similar in the same stress conditions. The up-regulation of Fatima, Angela, and Babara was due to their down-regulated methylation levels, while Wis was opposite. The 3′LTR of LTR retrotransposons contained many stress-responsive cis-elements, but the regulatory effect of which were not obvious under stress treatments. Compared with that in leaves, the four LTR retrotransposons in roots were more sensitive to stresses and had higher transposition activity. This study will help to further reveal the response regularity of LTR retrotransposons under stress conditions and accumulate data for improving of stress resistance breeding wheat by using transposable elements.

Key words: wheat (Triticum aestivum L.), abiotic stress, LTR retrotransposons, the relative expression level, transposition

表1

荧光定量引物序列"

基因
Gene name		 正向引物
Forward sequence (5′-3′)		 反向引物
Reverse sequence (5′-3′)
TaActin CTCCCTCACAACAACCGC TACCAGGAACTTCCATACCAAC
Fatima AAAAAGACCAGGCTGCAACG GCGCTGATATGTTGTGCTGG
Wis CGACTTGTGGCGAAGAGTTTG GCGATGCTTAAGTCCGTCAG
Angela GTTGGCGATTGCCGCATTTT TCAACAAAACCTTCCGGTTGC
Babara GGATTACGTTGCTGCTGAGG AGCAGCTGTGCCAAATTCCT

表2

甲基化分析引物序列"

引物名称
Primer name		 正向引物
Forward sequence (5'-3')		 反向引物
Reverse sequence (5'-3')
M-Fatima GGTTCGAATTTGGGTAAAAATATC AACATCCTTCCTAATAACGACGAC
U-Fatima GGTTCGAATTTGGGTAAAAATATC AAACATCCTTCCTAATAACAACAAC
M-Wis TAAGATTGAATAAGGTATTGGGATATC GACTCTCTAATCAACCAACAACGA
U-Wis AATAAGATTGAATAAGGTATTGGGATATT CAACTCTCTAATCAACCAACAACAA
M-Angela GATATTTTGTTGGATCGGAGTTC AAAAATAATCGTACGTCTTCACGAT
U-Angela GATATTTTGTTGGATTGGAGTTTG AAAAATAATCATACATCTTCACAAT
M-Babara TATTTAAGTGAGGTAAGTTTTTCGT AATAATACTATCATACATCCACGTT
U-Babara TATTTAAGTGAGGTAAGTTTTTTGT AATAATACTATCATACATCCACATT

表3

转座子展示分析引物序列"

引物名称
Primer name		 序列
Sequence (5'-3')		 引物名称
Primer name		 序列
Sequence (5'-3')
Fatima-1 CTCCCTCAGTCAGTGTCAAAA M-CA GACGATGAGTCCTGAGTAACA
Fatima-2 GGTCCCGAACTGTGCGTCT M-CC GACGATGAGTCCTGAGTAACC
Wis-1 ATGATTGTTGTTCGTCCTAT M-CG GACGATGAGTCCTGAGTAACG
Wis-2 CTCCCTCAGTCAGTGTCAAAA M-CT GACGATGAGTCCTGAGTAACT
Angela-1 ACAATTCTAGCATGAATAA M-GA GACGATGAGTCCTGAGTAAGA
Angela-2 AACTTTATTATTGCCTCTA M-GC GACGATGAGTCCTGAGTAAGC
Babara-1 TTAGCCAATCCTTTGTCCT M-GG GACGATGAGTCCTGAGTAAGG
Babara-2 TTGAAGGAGTTCCCACAAC M-GT GACGATGAGTCCTGAGTAAGT
M-0 GACGATGAGTCCTGAGTAA M-TA GACGATGAGTCCTGAGTAATA
M-AA GACGATGAGTCCTGAGTAAAA M-TC GACGATGAGTCCTGAGTAATC
M-AC GACGATGAGTCCTGAGTAAAC M-TG GACGATGAGTCCTGAGTAATG
M-AG GACGATGAGTCCTGAGTAAAG M-TT GACGATGAGTCCTGAGTAATT
M-AT GACGATGAGTCCTGAGTAAAT

图1

小麦中 4个LTR反转录转座子的结构分析 GAG: 群特异性抗原; AP: 天冬氨酸酶; INT: 整合酶; RT: 反转录酶; RH: 核糖核酸酶H。"

图2

不同非生物胁迫下小麦幼苗叶和根中的转座子表达模式分析 A: Fatima; B: Wis; C: Angela; D: Babara; Mock: 对照。采用Tukey法进行检验, *表示在0.05概率水平差异显著; **表示在0.01概率水平差异显著; ***表示在0.001概率水平差异显著。"

表4

小麦反转录转座子中胁迫应答相关顺式作用元件分析"

转座子
Exist in Tes		 胁迫应答
Stress response		 顺式
作用因子
CREs		 序列(核苷酸)
Sequence
(nucleotides)		 参考文献
Reference
Fatima 光学响应元件 Light response element Box 4 ATTAAT
无氧反应响应元件 Anaerobic reaction response element ARE AAACCA
Wis 脱落酸响应元件 Cis-acting element involved in abscisic acid responsiveness ABRE ACGTG [26]
ABRE TACGTGTC [27]
干旱响应元件 Drought response element MYC CATGTG [30]
MYC CANNTG [31]
光学响应元件 Light response element ACE GACACGTATG
G-Box CACGTT
G-box CACGAC
无氧反应响应元件 Anaerobic reaction response element ARE AAACCA
Angela 脱落酸响应元件 Cis-acting element involved in abscisic acid responsiveness DRE1 ACCGAGA [28]
LTRE CCGAC [29]
干旱响应元件 Drought response element MYC CATGTG [30]
MYC CANNTG [31]
LTRE CCGAC [29]
光学响应元件 Light response element G-box CACGAC
低温响应元件 Low temperature response element LTR CCGAAA
Babara 脱落酸响应元件 Cis-acting element involved in abscisic acid responsiveness ABRE ACGTG [26]
DRE1 ACCGAGA [28]
干旱响应元件 Drought response element MYC CATGTG [30]
MYC CANNTG [31]
光学响应元件 Light response element G-Box CACGTT
无氧反应响应元件 Anaerobic reaction response element ARE AAACCA
低温响应元件 Low temperature response element LTR CCGAAA
赤霉素响应元件 Gibberellin response element TATC-box TATCCCA

图3

不同非生物胁迫下小麦幼苗叶和根中反转录转座子的甲基化程度比较分析 A: Fatima; B: Wis; C: Angela; D: Babara; m: marker; M: 甲基化条带; U: 去甲基化条带。"

图4

小麦幼苗叶和根中Fatima和Wis转座活性分析 M: marker; 1, 5: 盐胁迫; 2, 6: ABA胁迫; 3, 7: H2O2胁迫; 4, 8: 干旱胁迫。图中箭头指示新的转座插入位点, AA、AT、TT、CG、GG、AC、AG、CA、CT、GT、TC、TG、TA、CC、GC和GA分别表示扩增引物的最后2个碱基。"

图5

小麦幼苗叶和根中Angela和Babara转座活性分析 M: marker; 1, 5: 盐胁迫; 2, 6: ABA胁迫; 3, 7: H2O2胁迫; 4, 8: 干旱胁迫。图中箭头指示新的转座插入位点, AA、AT、TT、CG、GG、AC、AG、CA、CT、GT、TC、TG、TA、CC、GC和GA分别表示扩增引物的最后2个碱基。"

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