小麦TaSPX1基因的克隆、表达及耐低氮逆境的功能研究

doi:10.3724/SP.J.1006.2024.31025

作物学报 ›› 2024, Vol. 50 ›› Issue (3): 576-589.doi: 10.3724/SP.J.1006.2024.31025

• 作物遗传育种·种质资源·分子遗传学 • 上一篇下一篇

小麦TaSPX1基因的克隆、表达及耐低氮逆境的功能研究

张宝华¹^,²(), 刘佳静¹^,², 田晓¹^,², 田旭钊¹^,², 董阔¹, 武郁洁¹, 肖凯³^,^*(), 李小娟¹^,²^,^*()

¹河北农业大学生命科学学院, 河北保定 071001
²河北省植物生理和分子病理学重点实验室, 河北保定 071001
³河北农业大学农学院, 河北保定 071001

收稿日期:2023-04-06 接受日期:2023-09-13 出版日期:2024-03-12 网络出版日期:2023-10-09
通讯作者: *肖凯, E-mail: xiaokai@hebau.edu.cn; 李小娟, E-mail: lxjlixiaojuan@126.com
作者简介:E-mail: zbh1383306@163.com
基金资助:
国家自然科学基金项目(32071935)

Cloning, expression, and functional analysis of wheat (Triticum aestivum L.) TaSPX1 gene in low nitrogen stress tolerance

ZHANG Bao-Hua¹^,²(), LIU Jia-Jing¹^,², TIAN Xiao¹^,², TIAN Xu-Zhao¹^,², DONG Kuo¹, WU Yu-Jie¹, XIAO Kai³^,^*(), LI Xiao-Juan¹^,²^,^*()

¹College of Life Sciences, Hebei Agricultural University, Baoding 071001, Hebei, China
²Key Laboratory of Plant Physiology and Molecular Pathology, Baoding 071001, Hebei, China
³College of Agronomy, Hebei Agricultural University, Baoding 071001, Hebei, China

Received:2023-04-06 Accepted:2023-09-13 Published:2024-03-12 Published online:2023-10-09
Contact: *E-mail: xiaokai@hebau.edu.cn; E-mail: lxjlixiaojuan@126.com
Supported by:
National Natural Science Foundation of China(32071935)

摘要/Abstract

摘要：

包含SPX、SPX-EXS、SPX-MFS和SPX-RING四个亚族的植物SPX基因家族在磷信号应答中发挥重要功能, 但迄今对小麦的该家族基因成员功能了解尚少。本研究前期从小麦(Triticum aestivum)中鉴定得到一个SPX亚族成员基因TaSPX1 (GenBank No. Ak332300), 亚细胞定位分析发现其定位于细胞核。对TaSPX1和来自小麦、拟南芥和水稻SPX家族的同源蛋白进行系统进化分析, 结果表明, 其与水稻SPX亚族的OsSPX1亲缘关系较近。应用RT-qPCR技术研究发现, TaSPX1的表达量在低氮胁迫下显著增加。构建烟草(Nicotiana tabacum)过表达转基因系(overexpression lines, OE), 应用MS营养液培养对野生型(WT)和OE株系OE3和OE4植株表型进行鉴定。发现在低氮胁迫下, OE3和OE4较WT表现明显的生长优势, 植株鲜重、根重和叶面积显著增加; 包括光合速率、胞间CO₂浓度、气孔导度和蒸腾速率在内的光合参数, 以及氮含量、可溶性糖、可溶性蛋白和叶绿素含量也都较WT显著增加。对氮吸收和同化相关基因的表达和酶活性测定结果表明, 上述基因的部分成员在OE植株中的表达量和氮同化酶活性升高。此外, 对包括SOD、POD和CAT在内的植株活性氧清除相关酶活力和MDA含量测定表明, 与WT相比, OE植株中保护酶的活性均明显提高, MDA含量降低。同时发现OE植株中部分保护酶基因成员的表达水平也较WT明显升高。这些结果初步证实了TaSPX1通过改善光合参数、增强氮吸收和转运以及加强保护酶系统等在介导植株抵御低氮胁迫中发挥重要作用。本项研究对小麦SPX家族成员抵御非生物逆境功能增加了新认识, 为作物抗低氮营养逆境的遗传改良提供了理论依据。

关键词: 小麦, TaSPX1, 亚细胞定位, 低氮胁迫, 功能分析

Abstract:

The SPX gene family includes four subgroups: SPX, SPX-EXS, SPX-MFS, and SPX-RING, which play an important role in phosphate signal response, but so far, little is known about the functions of this family in wheat. Previously, we identified a gene TaSPX1 (GenBank No. Ak332300), belonged to SPX subfamily from wheat (Triticum aestivum). Subcellular localization analysis showed that it targeted onto nucleus. Phylogenetic tree of TaSPX1 and its homologous proteins from the wheat, Arabidopsis, and rice SPX families showed that it was closely related to OsSPX1, a member of rice SPX subfamily. The relative expression level of TaSPX1 significantly increased under low nitrogen (low-N) stress when investigated by RT-qPCR. Transgenic tobacco (Nicotiana tabacum) overexpression lines were generated. Using the culture methods of Murashige & Skoog (MS) hydroponic solution, the phenotype of WT and OE under low-N stress treatment was investigated. We found that the plants of OE3 and OE4, two OE lines overexpressing TaSPX1, displayed increased growth vigor and leaf area, together with the enhanced plant fresh weight and root weight, and elevated photosynthetic parameters including photosynthetic rate (P_n), intercellular carbon dioxide concentration (C_i), stomatal conductance (G_s), and transpiration rate (T_r), along with the increased contents of nitrogen, soluble sugar, soluble protein, and chlorophyll content upon low-N stress with respect to WT. Studies on transport and assimilation related parameters showed that under low-N stress, the relative expression level of some related genes and the activities of nitrogen assimilation-related enzymes were increased. Assays on the SOD, POD, and CAT, the enzymes functional as cellular protector, revealed the higher activities of them in OE plants than those in WT. On the contrary, MDA content was decreased. Further RT-qPCR analysis indicated the expression levels of several protection enzymes mentioned above were higher in OE plants than those of WT under low-N stress. Therefore, TaSPX1 played an important role in mediating plant resistance to low-N stress by improving photosynthetic parameters, enhancing nitrogen absorption and transport, and strengthening the protective enzyme system. The results enrich new understanding on the function of wheat SPX family members involved in abiotic stress, and provide a theoretical basis for genetic improvement of crops against low-N stress.

Key words: wheat, TaSPX1, subcellular localization, low nitrogen stress, functional analysis

张宝华, 刘佳静, 田晓, 田旭钊, 董阔, 武郁洁, 肖凯, 李小娟. 小麦TaSPX1基因的克隆、表达及耐低氮逆境的功能研究[J]. 作物学报, 2024, 50(3): 576-589.

ZHANG Bao-Hua, LIU Jia-Jing, TIAN Xiao, TIAN Xu-Zhao, DONG Kuo, WU Yu-Jie, XIAO Kai, LI Xiao-Juan. Cloning, expression, and functional analysis of wheat (Triticum aestivum L.) TaSPX1 gene in low nitrogen stress tolerance[J]. Acta Agronomica Sinica, 2024, 50(3): 576-589.

图/表 10

表1

用于RT-qPCR的引物"

引物名称 Primer name	登录号 Accession number	上游引物 Forward sequence (5′-3′)	下游引物 Reverse sequence (5′-3′)
Ntaction	U91563	ATTACGAGGATGAAGAGGAAGC	TAAAAGCCTGAAGTTTGAAGA
Tatubulin	U76558	CATGCTATCCCTCGTCTCGACCT	CGCACTTCATGATGGAGTTGTAT
TaSPX1	Ak332300	ACTGGGGCTCTTATCCGTCT	CAGCCTGCTCGATCACAGAT
NtSOD1	KJ874395	GGACAAGGAGTTCCGGATGATT	CCATATTGACTTGAGCCAAGCG
NtSOD2	EU123521	TTGGGGAAGATGGTACTGCATC	TCATCAGGATCAGCGTGAACAA
NtFeSOD	KF724056	CAGGCCTGGAATCATCAGTT	TTTCACCAAGGCAAGCTTTT
NtMnSOD1	X14482	ACCACCAGAATCACCATCAGAC	CTATGCAATTTGGCGACGGTAG
NtMnSOD2	AB093097	AAGCCCTTGAACAGCTACATGA	TCGAGTGGTTAATGTGACCTCC
NtCAT	NTU07627	CAAGGATCTCTACGACTCGATT	CTTGAGGGCAAATAATCCACCT
NtCAT1	EF532799	TCGGAGGATAAGCTTCTCCAGA	ATGAGCACACTTGGGAGCATTA
NtCAT1;1	NTU93244	CAATGCTTGCCGATTTCTCT	AGCAGGATCGGTATGGTCAG
NtCAT1;2	HF564632	GAAGTTCCCTGACATGGTCCAT	AAGTGAACATGTGCAGGCTTTC
NtCAT1;3	HF564631	TGGCAAACGAGAGAAGTGTG	AAGCAAGCTTTTGACCCAGA
NtCAT3	HF564633	GACCCCAGAGGATTTGCTGTAA	ACCATGTCAGGGAACTTCATCC
NtPOD1;1	L02124	ATTAGGCTAAAGGTCCGTCGTG	GCAAGTGTTTCAAAGGGGCTAG
NtPOD1;2	AB044154	GAAGGTTCAGACGCTGAGAGAA	CAGGAAACAACTCCAGGACAGA
NtPOD1;3	AB044153	ATGGTCGTGGAGTTCTGGAATC	GAATGTCAACCCAAGCAATCCC
NtPOD1;4	D11396	TCCTAATGTAGGTGCAGGAGGA	CTCCAATTTCAGATGCAAGGGC
NtPOD1;5	AB178953	AGACCTCACCACCCAACAAC	GCATCTCTTCTCCCAAGTGC
NtPOD1;6	AB027753	CTTACTGGTGCGCACACTATTG	GCTGAGGAAGAAAGGTTGCATC
NtPOD1;7	AB027752	AATGGGTGCTTCTCTTCTTCGT	CCCTTTTCTCTCCTGTGAAGCT
NtPOD4	AY032675	TTCCATCAGGGGATTTGAAG	TGGTGCAGGAATGTTTGTGT
NtPOD9	AY032674	GAAACCCTAGCCAACCTTCC	TCGTGTTCGTGGAGTTTGAG
NtNRT1.1-s	AB102805	TACCGGTTTGTCGACGTGTC	TCTCTTCTCCTTGTACACATAC
NtNRT1.1-t	AB102806	CCGGCTTCATTGACACTCTT	CCTCTTCTCCTTGTACACATAC
NtNRT1.2-s	AB102807	GGGTTATCGTTCCCATTTGTCG	TCAGCAAGTCTCTTCTCCTTGT
NtNRT1.2-t	AB102808	GCCCTAACAGAGGTTAAGAGG	TCCCCATTTCAGCAAGTCTC
NtNRT2.1	AJ557583	TAGCCGTCACATTCATGATCCT	GATCGGCAGTTCTCGGCGAA
NtNRT2.2	AJ557584	CGTCGATCGTTAGGTATAATC	ATTAACTACTCACACTTGGGTAA
NtNR1	X06134	ATCAGGTGGATGGATGGCGA	ACAACCAACTCGAAGTACCC
NtNR2	X14059	ATGACTGGACCGTGGAAGTC	AAAACACCGCACCGTTTTAG
NtNR3	JN384020	GGACGAAGGTACCGCTGATA	GACTTCCACGGTCCAGTCAT
NtNR4	JN384019	TGTTTGAGCATCCGACTCAAC	GGAGGTGGTCCACAAGCCA
NtNR5	XM_016606384	ATGGATATTACCGGCCGGCA	TTACGTACCATCAGCATTAACA
NtNR6	XM_016583028	TTGAGGCACTGCTCAAAGAGC	TCCCAAATGTGAATCACTCCC
NtNIR1	NM_001324935	ATGGCATCTTTTTCTGTTAAAT	TTAATCTTCTGCTTCTTCTCTT
NtNIR2	X66145	GGGTTCCAGCTGATGATGTT	TGGCATTCTCTTCTCGACCT
NtNIR3	X66147	TGGCAATTCACGAGGCAATC	TCGTGGAACTGCACCAAAGT
NtNIR4	XR_001649867	AGTTGGGTTGACCAGTTTGC	ACACGGATTCCACTTCCTTG
NtNIR5	XM_016583028	AGCATCTCCAGAAGACTTGG	CCCAAATGTGAATCACTCCC
NtGS1	NM_001325250	ATTATGTCTCCGCTTTCAGATC	ATTGAGCAAACCAGAAACAAGC
NtGS2	X95932	TCTTGGTGGTTTTCCTGGAC	TGCTGAAATGCCAACTGAAG
NtGS3	XM_016584731	AAGATGGCTCAGATCTTGGCTC	CTTTAAACATTCAATGCGAGCT
NtGS4	XM_016579636	GATGTGCAAACCCACACCTT	CTGCAAATTCCTCGTCTGGT
NtGS5	XM_016640903	CGGGCATGATCAGAATTATTC	AGATTGGTCGGGATATGAACC
NtGS6	XM_016640901	AACTAGGCTGGGATTGACG	AGTCCCTCTCAGGCTCACAA
NtGS7	XM_016631331	CATTATGTCTCTGCTTTCAGAT	TCATGGCAGTAACGACGTATGG

表1

图1

图2

图3

表2

图4

图5

图6

图7

图8

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顺式元件 Cis-element	目标序列 Target sequence (5′-3′)	数目 Number	功能 Function
TATA-BOX	TATATA	23	胁迫响应元件和核心启动元件 Stress response elements and core promoter element
CAAT-BOX	CAAT	39	MYBHv1结合位点 MYBHv1 binding site
MYC	CATGTG/CATTTG	4	干旱和盐响应元件 Drought and salt response element
MYB	CAACAG/CAACTG/CAACCA	2	干旱和盐响应元件 Drought and salt response element
ABRE	TACGTGTC/ACGTG	7	ABA响应元件 ABA-responsive element
Root motif	ATATT	2	根系生长 Root growth
G-box	CACGTT/TACGTG/GCCACGTGGA	4	参与光响应顺式作用调节元件 cis-acting regulatory element involved in light responsiveness
LTRE	AGTCGG	2	低氮胁迫响应元件 Response to low nitrogen stress
/	NANGAG	4	低氮胁迫响应元件 Response to low nitrogen stress

小麦TaSPX1基因的克隆、表达及耐低氮逆境的功能研究

Cloning, expression, and functional analysis of wheat (Triticum aestivum L.) TaSPX1 gene in low nitrogen stress tolerance

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