小麦倒春寒响应基因<i>TaJAZ6</i>的克隆、表达模式及亚细胞定位分析

doi:10.7668/hbnxb.20194434

摘要/Abstract

摘要：

JAZ蛋白在植物生长发育和胁迫信号通路中具有关键作用。为探究JAZ蛋白在小麦倒春寒中的调控机制,从小麦幼穗中克隆了 TaJAZ6基因,并对其分子特征、表达特性与亚细胞定位进行分析。结果表明,该基因CDS 序列全长为549 bp,编码178个氨基酸,预测编码蛋白质的分子质量为 18.376 ku,理论等电点为 9.37,不稳定系数为62.44,属于不稳定蛋白。该基因编码蛋白质具有1个 TIFY 结构域和1个CCT_2结构域。系统进化树分析发现,该蛋白质与野生二粒小麦和乌拉尔图小麦TIFY 11b蛋白亲缘关系最近。TaJAZ6基因启动子区除含有CAAT-box、TATA-box等基础作用元件外,还含有激素类响应元件、光响应元件、低温响应元件、防御和应激反应元件等。实时荧光定量PCR分析发现,TaJAZ6 基因在根、茎、叶和幼穗中均有表达,根系中表达量最高。TaJAZ6 基因还受低温和茉莉酸甲酯( MeJA) 的诱导表达,低温胁迫下,TaJAZ6在矮抗58(耐倒春寒)和郑麦366(倒春寒敏感)根、茎和叶中表达趋势相同,均显著升高;喷施300,350 μmol/L的MeJA之后再低温处理TaJAZ6在2个小麦品种中表达量均显著下降。TaJAZ6在低温胁迫后的幼穗中表达量出现相反的趋势,在矮抗58幼穗中表达量显著下降,在郑麦366幼穗中则显著上升,推测该基因可能负调控了小麦倒春寒胁迫防御反应。通过喷施MeJA显著降低了低温胁迫下2个小麦品种幼穗中TaJAZ6基因的相对表达量,同时提高了小麦的结实粒数。亚细胞定位试验显示,TaJAZ6 蛋白定位于细胞核中。以上研究结果表明,TaJAZ6 可能在小麦响应倒春寒胁迫过程中发挥重要作用。

关键词: 小麦, TaJAZ6基因, 表达分析, 低温胁迫, 亚细胞定位

Abstract:

JAZ protein plays a key role in plant growth and development and stress signaling pathways.In order to explore the regulatory mechanism of JAZ protein in wheat late spring cold,TaJAZ6 gene was cloned from young spike of wheat,and its molecular characteristics,expression characteristics and subcellular localization were analyzed.The results showed that the full-length CDS sequence of the gene was 549 bp,encoding 178 amino acids.The predicted molecular weight of the encoded protein was 18.376 ku,the theoretical isoelectric point was 9.37,and the instability coefficient was 62.44,so it was an unstable protein.The protein encoded by this gene had a TIFY domain and a CCT_2 domain.Phylogenetic tree analysis showed that the protein had the closest relationship with TIFY 11b proteins of Triticum dicoccoides and Triticum urartu.In addition to the basic response elements such as CAAT-box and TATA-box,the promoter region of TaJAZ6 gene also contained hormone response elements,light response elements,low temperature response elements,defense and stress response elements.Real-time Quantitative PCR analysis showed that TaJAZ6 gene was expressed in roots,stems,leaves and young panicles,with the highest expression in roots.The expression of TaJAZ6 gene was also induced by low temperature and methyl jasmonate (MeJA).Under low temperature stress,the expression trends of TaJAZ6 in roots,stems and leaves of Aikang 58 (tolerant to late spring cold ) and Zhengmai 366 ( sensitive to late spring cold ) were the same trend,which significantly increased.After spraying 300,350 μmol/L MeJA,the expression of TaJAZ6 in plants treated with low temperature decreased significantly in both wheat varieties.The expression level of TaJAZ6 in the young panicles after low temperature stress showed an opposite trend.The expression level of TaJAZ6 in the young panicles of Aikang 58 decreased significantly,and increased significantly in the young panicles of Zhengmai 366.It was speculated that the gene might negatively regulate the defense response of wheat to late spring cold stress.By spraying MeJA,the relative expression of TaJAZ6 gene in young spikes of two wheat varieties under low temperature stress was significantly reduced,and the grain number of wheat was increased.Subcellular localization assay showed that TaJAZ6 protein was localized in the nucleus.The results above indicate that TaJAZ6 may play an important role in the response of wheat to late spring cold stress.

Key words: Wheat, TaJAZ6, Expression analysis, Cold stress, Subcellular localization

张自阳, 周谦, 王依, 王志伟, 朱启迪, 茹振钢, 刘明久. 小麦倒春寒响应基因TaJAZ6的克隆、表达模式及亚细胞定位分析[J]. 华北农学报, 2024, 39(1): 27-36. doi: 10.7668/hbnxb.20194434.

ZHANG Ziyang, ZHOU Qian, WANG Yi, WANG Zhiwei, ZHU Qidi, RU Zhengang, LIU Mingjiu. Cloning,Expression Pattern and Subcellular Localization Analysis of Late Spring Cold Response Gene TaJAZ6 in Wheat[J]. Acta Agriculturae Boreali-Sinica, 2024, 39(1): 27-36. doi: 10.7668/hbnxb.20194434.

http://www.hbnxb.net/CN/Y2024/V39/I1/27

图/表 15

表1 引物序列

Tab.1 Primer sequences

引物名称 Primer name	引物序列(5'—3') Primer sequence(5'—3')	用途 Function
TaJAZ6-F	CCTCGAGAGAGAACAAGTGTTGCCATG	克隆基因
TaJAZ6-R	GACTAGTCGCCTCAAGACATCGTTT
TaJAZ6-qPCR-F	AGCAGAAGATGTCCTCCT	荧光定量PCR
TaJAZ6-qPCR-R	CTCACCATCTTCTACGGC
Tublin-F	GTGGAACTGGCTCTGGC	内参基因
Tublin-R	CGCTCAATGTCAAGGGA
GFP-TaJAZ6-F	CCTCGAGAGAGAACAAGTGTTGCCATG	亚细胞定位
GFP-TaJAZ6-R	GACTAGTCGCCTCAAGACATCGTTT

图1 不同处理对2个小麦品种结实粒数的影响不同小写字母表示0.05水平差异显著。图12—13 同。

Fig.1 Effects of different treatments on grain number of two wheat varieties Different lowercase letters indicate significant differences at 0.05 level.The same as Fig.12—13.

图2 小麦幼穗总RNA

Fig.2 Total RNA extracted from young panicle of wheat

图3 小麦TaJAZ6基因PCR产物

Fig.3 PCR amplification products of TaJAZ6 in wheat 1.TaJAZ6;M.DL2000 Marker。

图4 TaJAZ6蛋白亲疏水性分析

Fig.4 Hydrophilicity analysis of TaJAZ6 protein

图5 TaJAZ6蛋白的保守结构域分析

Fig.5 Conserved domain analysis of TaJAZ6 protein

图6 TaJAZ6蛋白质二级结构预测

Fig.6 Secondary structure analysis of TaJAZ6 protein

图7 TaJAZ6蛋白的三级结构预测

Fig.7 Tertiary structure analysis of TaJAZ6 protein

图8 TaJAZ6蛋白信号肽分析

Fig.8 Signal peptide analysis of TaJAZ6 protein

表2 TaJAZ6基因启动子的顺式作用元件分析

Tab.2 Analysis of cis-acting elements of TaJAZ6 promoter

元件名称 Element	序列 Sequence	功能描述 Function description	数量/个 Number
CAAT-box	CAAAT	启动子和增强子区域的共同顺式作用元件	8
	CAAT		10
	CCAAT		2
TATA-box	TATA	-30 bp处左右基因核心启动子元件	7
	ATATAT		1
	CCTATAAAAA		1
	TATATAAATC		1
	TATATA		1
ABRE	ACGTG	ABA应答元件	3
	CACGTG		1
G-Box	CACGTT	参与光反应的顺式调控元件	1
	CACGTG		2
	TAAACGTG		2
	CACGTC		2
Box4	ATTAAT	参与光响应的保守 DNA 模块的一部分	1
CGTCA-motif	CGTCA	参与 MeJA 响应的顺式作用调节元件	1
LTR	CCGAAA	低温响应元件	1
TGACG-motif	TGACG	MeJA 响应元件	1
TCT-motif	TCTTAC	光响应元件的一部分	2
TCCC-motif	TCTCCCT	光响应元件的一部分	1
CAT-box	GCCACT	与分生组织表达相关的顺式调节元件	1
MSA-like	AACGG	参与细胞周期调控	1
TC-rich repeats	GTTTTCTTAC	参与防御和应激反应的顺式作用元件	1

图9 TaJAZ6蛋白同源序列比对 Td.野生二粒小麦;Tu.乌拉尔图小麦;Lr.硬直黑麦草;Hv.大麦;At.节节麦;Ob.短药野生稻;Zm.玉米;Bd.二穗短柄草。图10同。

Fig.9 Homologous sequence alignment of TaJAZ6 protein Td.Triticum dicoccoides;Tu.Triticum urartu;Lr.Lolium rigidum;Hv.Hordeum vulgare;At.Aegilops tauschii subsp. strangulata; Ob.Oryza brachyantha;Zm.Zea mays;Bd.Brachypodium distachyon.The same as Fig.10.

图10 不同物种JAZ同源蛋白进化分析

Fig.10 Phylogenetic analysis of JAZ homologous proteins in different species

图11 TaJAZ6-GFP蛋白在洋葱表皮细胞中的定位 A.空白载体荧光;B.空白载体明场;C.空白载体叠加;D.表达载体荧光;E.表达载体明场;F.表达载体叠加。

Fig.11 TaJAZ6-GFP protein localization in onion epidermal cells A.Non transfection vector fluorescence;B.Non-transfection vector bright field;C.Non-transfection vector superposition;D.Expression vector fluorescence;E.Expression vector bright field;F.Expression vector superposition.

图12 TaJAZ6基因的组织差异性表达分析

Fig.12 Tissue-specific expression analysis of TaJAZ6 gene

图13 TaJAZ6基因的外源胁迫表达分析

Fig.13 Expression analysis of TaJAZ6 gene under exogenous stress

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[41]	Hu Y R, Jiang Y J, Han X, Wang H P, Pan J J, Yu D Q. Jasmonate regulates leaf senescence and tolerance to cold stress:crosstalk with other phytohormones[J]. Journal of Experimental Botany, 2017, 68(6):1361-1369.doi:10.1093/jxb/erx004. URL

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小麦倒春寒响应基因TaJAZ6的克隆、表达模式及亚细胞定位分析

Cloning,Expression Pattern and Subcellular Localization Analysis of Late Spring Cold Response Gene TaJAZ6 in Wheat

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小麦倒春寒响应基因TaJAZ6的克隆、表达模式及亚细胞定位分析

Cloning,Expression Pattern and Subcellular Localization Analysis of Late Spring Cold Response Gene TaJAZ6 in Wheat

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