黑麦NHX基因家族的鉴定及盐胁迫下的表达分析

doi:10.7668/hbnxb.20195534

摘要/Abstract

摘要：

植物钠氢反转运蛋白(NHX,Na⁺/H⁺ antiporter)在植物钠钾离子平衡、细胞pH值调节中起重要的作用。为探究黑麦耐盐性与NHX的关系,通过生物信息学流程对黑麦的NHX基因进行鉴定与分析,结合RT-qPCR技术对ScNHXs在盐胁迫下的表达模式进行检测,为探究NHX基因在黑麦中的潜在功能以及黑麦耐盐基因挖掘等研究提供参考信息。共鉴定获得10个黑麦NHX基因家族成员(ScNHX1~ScNHX10),系统进化树分析表明,其可分为Vac和Endo 2个亚家族,分别包含4,6个基因。其编码蛋白理化性质分析表明,分子质量为27.92~59.72 ku,氨基酸数目为253~546 aa,等电点为5.17~8.81,大多数蛋白属于酸性蛋白,均未预测到信号肽存在但均具有跨膜结构,亚细胞定位预测ScNHXs定位于质膜和液泡中。空间结构预测显示,其二级结构主要由α-螺旋和无规则卷曲构成。基因结构和motif分析发现,ScNHXs的外显子数为13~24,且均具有保守的Na⁺/H⁺交换结构域。此外,顺式作用元件分析表明,ScNHXs启动子区域中均发现众多与激素响应以及非生物胁迫响应等生物过程相关的元件。黑麦转录组数据分析发现,在黑麦的不同组织中ScNHXs表达模式存在明显差异。RT-qPCR分析表明,ScNHXs对不同浓度NaCl胁迫有不同程度的响应,并且在较长时间内能够持续响应盐胁迫。综上所述,ScNHXs可能参与黑麦对盐胁迫的生物调节相关过程。

关键词: 黑麦, NHX基因家族, 生物信息学, 盐胁迫, 基因表达

Abstract:

Plant sodium-hydrogen antiporter(NHX,Na⁺/H⁺ antiporter)plays a crucial role in plant sodium and potassium ion balance and cellular pH regulation.In order to investigate the relationship between salt tolerance and ScNHXs,it was conducted to identify and analyze the ScNHXs by bioinformatics process,and to examine the expression pattern of ScNHXs under salt stress by RT-qPCR,which can provide the reference information for the investigation of the potential functions of ScNHXs as well as the mining of salt tolerance genes in rye.A total of 10 rye NHX gene family members(ScNHX1—ScNHX10)were identified,and the phylogenetic tree analysis showed that they could be divided into two subfamilies,Vac and Endo,containing four and six genes,respectively.Physicochemical property analysis of the encoded proteins showed that most of the molecular weight ranged from 27.92 to 59.72 ku,the number of amino acids from 253 to 546 aa,and the isoelectric point between 5.17 and 8.81,with most of proteins being classified as acidic proteins.Signal peptide prediction indicated the absence of signal peptides in the members,and transmembrane structure analysis revealed that all members possessed transmembrane structures.The subcellular localization prediction indicated that ScNHXs were located in the plasma membrane and vesicles.Spatial structure prediction showed that their secondary structures mainly consisted of α-helices and irregular convolutions.Gene structure and motif analyses revealed that the number of exons of the ScNHXs varied from 13 to 24,and all of them possessed a conserved Na⁺/H⁺ exchange structural domain.In addition,cis-acting element analysis revealed that numerous elements related to hormone response and abiotic stresses were found in the promoter region of ScNHXs.Analysis of rye transcriptome data revealed significant differences in the expression patterns of ScNHXs in different tissues of rye.RT-qPCR analysis showed that ScNHXs responded differently to different concentrations of NaCl stress,and were able to persistently respond to salt stress over a long period of time.In summary,ScNHXs may be involved in the biological regulation during salt stress in rye.

Key words: Rye, NHX gene family, Bioinformatics, Salt stress, Gene expression

中图分类号:

王昭懿, 崔原瑗, 韩梦荞, 刘正文, 邓习, 豆飞飞, 任毓昭, 刘彩霞, 刘凤楼, 王掌军, 孙洋洋, 任民, 李清峰. 黑麦NHX基因家族的鉴定及盐胁迫下的表达分析[J]. 华北农学报, 2025, 40(3): 9-18. doi: 10.7668/hbnxb.20195534.

WANG Zhaoyi, CUI Yuanyuan, HAN Mengqiao, LIU Zhengwen, DENG Xi, DOU Feifei, REN Yuzhao, LIU Caixia, LIU Fenglou, WANG Zhangjun, SUN Yangyang, REN Min, LI Qingfeng. Identification and Expression Analysis of the NHX Gene Family under Salt Stress in Rye[J]. Acta Agriculturae Boreali-Sinica, 2025, 40(3): 9-18. doi: 10.7668/hbnxb.20195534.

http://www.hbnxb.net/CN/Y2025/V40/I3/9

图/表 12

表1 RT-qPCR反应中使用的引物

Tab.1 The sequences of primers used in RT-qPCR reactions

引物名称 Primer name	引物序列(5'—3') Primer sequence(5'—3')	退火温度/℃ Annealing temperature
NHX1-F	GAGGAGAACCGCTGGCTC	59.8
NHX1-R	TCGCTGAAGACGAGCACG	60.1
NHX2-F	TTGTGAGCCATCCAGCCC	59.6
NHX2-R	TGACCGTGTTGGACTCGC	60.0
NHX3-F	CGTGGTGTCCATCTGCGT	60.0
NHX3-R	TGATGAGCGCGGTGATGG	60.2
NHX4-F	TGGTCATACTGCCGTGCG	60.1
NHX4-R	TTGAGATGTCGGCTGCGG	60.1
NHX5-F	TCGCTGGAGCTGAGCATG	59.8
NHX5-R	CTGCAGCAGGATCCCCAC	60.1
NHX6-F	TGGTGCTCTCCTTCGTGC	60.0
NHX6-R	CCGATGAGGAGGGATGCG	60.0
NHX7-F	GTTCGTGCTCAGCCACCT	60.0
NHX7-R	GCCCTCCCACGATCAACC	60.1
NHX8-F	TCAGAGGACTCCCAGCGT	59.6
NHX8-R	AGCCGATGTGTGACCAGC	60.0
NHX9-F	CCGCGCTGGACAGAAACT	60.4
NHX9-R	TCCACGTCTTGCAGCTCG	60.0
NHX10-F	ATCGCGGCCAGAACATCG	60.6
NHX10-R	AAGCGCAAAAGCCATGGC	60.0

表2 ScNHXs蛋白性质及亚细胞定位预测

Tab.2 Prediction of properties and subcellular localizations of ScNHXs proteins in rye

基因名称 Gene name	ID号 Gene identifier	氨基酸数/ aa Amino acids	分子质量/ku Molecular weight	等电点 pI	不稳定系数 Instability index	亲水性平均系数 Grand average of hydropathicity	亚细胞定位 Subcellular localization
ScNHX1	ScWN2R01G102600.1	538	59.03	8.41	34.13	0.612	质膜、液泡
ScNHX2	ScWN4R01G176300.1	541	59.49	8.81	34.85	0.573	质膜、液泡
ScNHX3	ScWN4R01G271700.1	527	58.03	8.41	33.55	0.603	质膜、液泡
ScNHX4	ScWN4R01G477800.1	546	59.72	8.13	31.26	0.679	质膜、液泡、高尔基体
ScNHX5	ScWN5R01G182900.1	534	58.35	5.13	47.85	0.477	质膜
ScNHX6	ScWN5R01G285100.1	533	59.02	5.20	43.63	0.389	质膜
ScNHX7	ScWN6R01G382600.1	527	58.15	5.59	45.89	0.446	质膜、液泡
ScNHX8	ScWN6R01G493000.1	344	37.93	5.17	46.36	0.099	质膜、液泡、内质网
ScNHX9	ScWNUn01G004700.1	533	58.73	5.29	47.70	0.377	质膜、液泡
ScNHX10	ScWNUn01G024600.1	253	27.92	5.36	40.77	0.109	质膜、液泡

表3 ScNHXs的蛋白质二级结构分析

Tab.3 The secondary structure of ScNHXs protein%

蛋白 Protein	α-螺旋 Alpha helix	β-转角 Beta turn	无规则卷曲 Random coil	延伸链 Extended strand
ScNHX1	45.54	2.97	33.27	18.22
ScNHX2	40.30	5.18	33.46	21.07
ScNHX3	44.78	4.17	30.36	20.68
ScNHX4	44.51	4.03	33.52	17.95
ScNHX5	46.44	3.93	32.96	16.67
ScNHX6	47.84	2.63	34.52	15.01
ScNHX7	49.34	2.09	33.21	15.37
ScNHX8	45.64	2.03	39.83	12.50
ScNHX9	53.73	3.19	30.39	13.70
ScNHX10	41.50	3.56	39.53	15.42

图1 NHX基因的系统进化树

Fig.1 The phylogenetic tree of NHX gene

图2 ScNHXs的染色体定位图不同颜色代表基因在染色体上的密度,由红到蓝表示密度由高到低。

Fig.2 Chromosome localization map of ScNHXs The density of genes on chromosomes is indicated by various colors,from red to blue indicating high density to low density.

图3 ScNHX基因家族基因结构分析

Fig.3 Gene structure analysis of the ScNHX gene family

图4 ScNHX基因家族motif分析

Fig.4 ScNHX gene family motif analysis

图5 黑麦NHXs蛋白中保守基序的Logo分析

Fig.5 Logo analysis of conserved motifs in rye NHXs proteins

图6 ScNHXs基因顺式作用元件预测顺式作用元件的数量用红色表示,红色越深代表数量越多。

Fig.6 Cis-acting elements prediction of ScNHXs genes Numbers of cis-acting elements are indicated in red,with darker red shading representing higher numbers.

图7 ScNHXs在不同组织和发育时期的表达不同颜色的方格表示不同组织中基因的相对表达水平:红色代表高阶,蓝色代表低阶。DAA.开花后天数。

Fig.7 Expression of ScNHXs in different tissues and developmental periods Different colored patches represent the relative expression levels of genes in different tissues:red for high level,blue for low level.DAA.Days after anthesis.

图8 ScNHXs在不同浓度NaCl处理下的相对表达量图中不同字母表示在P<0.05水平上存在显著差异(Tukey test)。图9同。

Fig.8 The relative expression levels of ScNHXs under varying NaCl concentrations Different letters denote significant difference at P<0.05 level(Tukey test).The same as Fig.9.

图9 ScNHXs在300 mmol/L NaCl处理下不同时间的相对表达量

Fig.9 The relative expression of ScNHXs at different time points following 300 mmol/L NaCl treatment

参考文献 48

[1]	Khan Z, Jan R, Asif S, Farooq M, Jang Y H, Kim E G, Kim N, Kim K M. Exogenous melatonin induces salt and drought stress tolerance in rice by promoting plant growth and defense system[J]. Scientific Reports, 2024, 14:1214.doi:10.1038/s41598-024-51369-0.
[2]	Yang L, Han R, Duan Y K, Li J Y, Gou T Y, Zhou J, Zhu H J, Xu Z M, Guo J, Gong H J. Exogenous application of silicon and selenium improves the tolerance of tomato plants to calcium nitrate stress[J]. Plant Physiology and Biochemistry, 2024, 207:108416.doi:10.1016/j.plaphy.2024.108416.
[3]	Zhang K Y, Chang L, Li G H, Li Y F. Advances and future research in ecological stoichiometry under saline-alkali stress[J]. Environmental Science and Pollution Research, 2023, 30(3):5475-5486.doi:10.1007/s11356-022-24293-x.
[4]	Deinlein U, Stephan A B, Horie T, Luo W, Xu G H, Schroeder J I. Plant salt-tolerance mechanisms[J]. Trends in Plant Science, 2014, 19(6):371-379.doi:10.1016/j.tplants.2014.02.001. pmid: 24630845
[5]	Hu Y C, Schmidhalter U. Opportunity and challenges of phenotyping plant salt tolerance[J]. Trends in Plant Science, 2023, 28(5):552-566.doi:10.1016/j.tplants.2022.12.010. pmid: 36628656
[6]	Zhu X L, Wang B Q, Wang X, Wei X H. Identification of the CIPK-CBL family gene and functional characterization of CqCIPK14 gene under drought stress in quinoa[J]. BMC Genomics, 2022, 23(1):447.doi:10.1186/s12864-022-08683-6.
[7]	Liang W J, Ma X L, Wan P, Liu L Y. Plant salt-tolerance mechanism:a review[J]. Biochemical and Biophysical Research Communications, 2018, 495(1):286-291.doi:10.1016/j.bbrc.2017.11.043.
[8]	Isayenkov S V, Dabravolski S A, Pan T, Shabala S. Phylogenetic diversity and physiological roles of plant monovalent cation/H⁺ antiporters[J]. Frontiers in Plant Science, 2020, 11:573564.doi:10.3389/fpls.2020.573564.
[9]	Pehlivan N, Sun L, Jarrett P, Yang X J, Mishra N, Chen L, Kadioglu A, Shen G X, Zhang H. Co-overexpressing a plasma membrane and a vacuolar membrane sodium/proton antiporter significantly improves salt tolerance in transgenic Arabidopsis plants[J]. Plant and Cell Physiology, 2016, 57(5):1069-1084.doi:10.1093/pcp/pcw055. pmid: 26985021
[10]	Cao B N, Xia Z Q, Liu C Y, Fan W, Zhang S, Liu Q, Xiang Z H, Zhao A C. New insights into the structure-function relationship of the endosomal-type Na⁺,K⁺/H⁺ antiporter NHX6 from mulberry(Morus notabilis)[J]. International Journal of Molecular Sciences, 2020, 21(2):428.doi:10.3390/ijms21020428.
[11]	Sze H, Chanroj S. Plant endomembrane dynamics:studies of K⁺/H⁺ antiporters provide insights on the effects of pH and ion homeostasis[J]. Plant Physiology, 2018, 177(3):875-895.doi:10.1104/pp.18.00142.
[12]	Bassil E, Coku A, Blumwald E. Cellular ion homeostasis:emerging roles of intracellular NHX Na⁺/H⁺ antiporters in plant growth and development[J]. Journal of Experimental Botany, 2012, 63(16):5727-5740.doi:10.1093/jxb/ers250. pmid: 22991159
[13]	Huertas R, Rubio L, Cagnac O, García-Sánchez M J, De Dios Alché J, Venema K, Fernández J A, Rodríguez-Rosales M P. The K⁺/H⁺ antiporter LeNHX2 increases salt tolerance by improving K⁺ homeostasis in transgenic tomato[J]. Plant,Cell & Environment, 2013, 36(12):2135-2149.doi:10.1111/pce.12109.
[14]	Ohnishi M, Fukada-Tanaka S, Hoshino A, Takada J, Inagaki Y, Iida S. Characterization of a novel Na⁺/H⁺ antiporter gene InNHX2 and comparison of InNHX2 with InNHX1,which is responsible for blue flower coloration by increasing the vacuolar pH in the Japanese morning glory[J]. Plant and Cell Physiology, 2005, 46(2):259-267.doi:10.1093/pcp/pci028. pmid: 15695437
[15]	Fu X K, Lu Z Y, Wei H L, Zhang J J, Yang X, Wu A M, Ma L, Kang M, Lu J H, Wang H T, Yu S X. Genome-wide identification and expression analysis of the NHX(sodium/hydrogen antiporter)gene family in cotton[J]. Frontiers in Genetics, 2020, 11:964.doi:10.3389/fgene.2020.00964.
[16]	Ayadi M, Martins V, Ben Ayed R, Jbir R, Feki M, Mzid R, Géros H, Aifa S, Hanana M. Genome wide identification,molecular characterization,and gene expression analyses of grapevine NHX antiporters suggest their involvement in growth,ripening,seed dormancy,and stress response[J]. Biochemical Genetics, 2020, 58(1):102-128.doi:10.1007/s10528-019-09930-4.
[17]	Sharma P, Mishra S, Pandey B, Singh G. Genome-wide identification and expression analysis of the NHX gene family under salt stress in wheat(Triticum aestivum L.)[J]. Frontiers in Plant Science, 2023, 14:1266699.doi:10.3389/fpls.2023.1266699.
[18]	艾迪, 魏永成, 孟景祥, 张捷, 仲崇禄, 张勇. 木麻黄NHX基因家族的鉴定及盐胁迫下的表达分析[J]. 西北植物学报, 2023, 43(5):750-760.doi:10.7606/j.issn.1000-4025.2023.05.0750.
	Ai D, Wei Y C, Meng J X, Zhang J, Zhong C L, Zhang Y. Identification of NHX gene family and their responses to salt stress in Casuarina equisetifolia[J]. Acta Botanica Boreali-Occidentalia Sinica, 2023, 43(5):750-760.
[19]	张业猛, 朱丽丽, 陈志国. 藜麦NHX基因家族鉴定及盐胁迫下表达分析[J]. 生物技术通报, 2022, 38(12):184-193.doi:10.13560/j.cnki.biotech.bull.1985.2022-0404.
	Zhang Y M, Zhu L L, Chen Z G. Identification and expression analysis of NHX gene family in quinoa under salt stress[J]. Biotechnology Bulletin, 2022, 38(12):184-193.
[20]	Pires I S, Negrão S, Pentony M M, Abreu I A, Oliveira M M, Purugganan M D. Different evolutionary histories of two cation/proton exchanger gene families in plants[J]. BMC Plant Biology, 2013, 13(1):97.doi:10.1186/1471-2229-13-97.
[21]	Kumar K, Jha S K, Kumar V, Sagar P, Tripathi S, Rathore M, Singh A K, Soren K R, Dixit G P. Identification and characterization of NHX gene family for their role under salt stress in Vigna mungo[J]. Physiologia Plantarum, 2024, 176(5):e14563.doi:10.1111/ppl.14563.
[22]	Parveen, Anwar-Ul-Haq M, Aziz T, Aziz O, Maqsood L. Potassium induces carbohydrates accumulation by enhancing morpho-physiological and biochemical attributes in soybean under salinity[J]. Archives of Agronomy and Soil Science, 2021, 67(7):946-959.doi:10.1080/03650340.2020.1769075.
[23]	Wu X X, Li J, Wu X D, Liu Q, Wang Z K, Liu S S, Li S N, Ma Y L, Sun J, Zhao L, Li H Y, Li D M, Li W B, Su A Y. Ectopic expression of Arabidopsis thaliana Na⁺(K⁺)/H⁺ antiporter gene,AtNHX5,enhances soybean salt tolerance[J]. Genetics and Molecular Research, 2016, 15(2):1-12.doi:10.4238/gmr.15027483.
[24]	张慧军, 张万科, 俞嘉宁, 孔祥强, 陈受宜, 王丹, 张国强, 董合忠. 过量表达TaNHX2基因提高转基因棉花的抗旱耐盐性[J]. 东北农业科学, 2021, 46(1):31-35,71.doi:10.16423/j.cnki.1003-8701.2021.01.009.
	Zhang H J, Zhang W K, Yu J N, Kong X Q, Chen S Y, Wang D, Zhang G Q, Dong H Z. The overexpression of TaNHX2 gene can improve salt and drought tolerance of transgenic cotton[J]. Journal of Northeast Agricultural Sciences, 2021, 46(1):31-35,71.
[25]	Crespo-Herrera L A, Garkava-Gustavsson L, Åhman I. A systematic review of rye(Secale cereale L.)as a source of resistance to pathogens and pests in wheat(Triticum aestivum L.)[J]. Hereditas, 2017, 154(1):14.doi:10.1186/s41065-017-0033-5.
[26]	Finn R D, Coggill P, Eberhardt R Y, Eddy S R, Mistry J, Mitchell A L, Potter S C, Punta M, Qureshi M, Sangrador-Vegas A, Salazar G A, Tate J, Bateman A. The pfam protein families database:towards a more sustainable future[J]. Nucleic Acids Research, 2016, 44(D1):D279-D285.doi:10.1093/nar/gkv1344.
[27]	Potter S C, Luciani A, Eddy S R, Park Y, Lopez R, Finn R D. HMMER web server:2018 update[J]. Nucleic Acids Research, 2018, 46(W1):W200-W204.doi:10.1093/nar/gky448.
[28]	Singh J, Varshney V, Tak N, Jha S. Genome-wide identification and expression analysis of glycogen synthase kinase encoding genes in foxtail millet(Setaria italica L.) under salinity,dehydration,and oxidative stress[J]. Plant Stress, 2023, 8:100165.doi:10.1016/j.stress.2023.100165.
[29]	丛郁, 杨顺瑛, 宋志忠, 郝东利, 苏彦华. 葡萄AMT基因家族生物信息学分析[J]. 中国农学通报, 2011, 27(25):193-199.
	Cong Y, Yang S Y, Song Z Z, Hao D L, Su Y H. Bioinformatics analysis of AMT protein family in grape[J]. Chinese Agricultural Science Bulletin, 2011, 27(25):193-199. doi: 10.11924/j.issn.1000-6850.2011-1057
[30]	Mann S, Li J Y, Chen Y P. A pHMM-ANN based discriminative approach to promoter identification in prokaryote genomic contexts[J]. Nucleic Acids Research, 2007, 35(2):e12.doi:10.1093/nar/gkl1024. pmid: 17170007
[31]	Letunic I, Bork P. Interactive tree of life(iTOL)v4:recent updates and new developments[J]. Nucleic Acids Research, 2019, 47(W1):W256-W259.doi:10.1093/nar/gkz239.
[32]	Chen C J, Chen H, Zhang Y, Thomas H R, Frank M H, He Y H, Xia R. TBtools:an integrative toolkit developed for interactive analyses of big biological data[J]. Molecular Plant, 2020, 13(8):1194-1202.doi:10.1016/j.molp.2020.06.009.
[33]	Hu B, Jin J P, Guo A Y, Zhang H, Luo J C, Gao G. GSDS 2.0:an upgraded gene feature visualization server[J]. Bioinformatics, 2015, 31(8):1296-1297.doi:10.1093/bioinformatics/btu817.
[34]	Bailey T L, Boden M, Buske F A, Frith M, Grant C E, Clementi L, Ren J Y, Li W W, Noble W S. MEME Suite:tools for motif discovery and searching[J]. Nucleic Acids Research, 2009, 37(S2):W202-W208.doi:10.1093/nar/gkp335.
[35]	Lescot M, Déhais P, Thijs G, Marchal K, Moreau Y, Van de Peer Y, Rouzé P, Rombauts S. PlantCARE,a database of plant cis-acting regulatory elements and a portal to tools for in silico analysis of promoter sequences[J]. Nucleic Acids Research, 2002, 30(1):325-327.doi:10.1093/nar/30.1.325.
[36]	Livak K J, Schmittgen T D. Analysis of relative gene expression data using real-time quantitative PCR and the 2^-ΔΔCt method[J]. Methods, 2001, 25(4):402-408.doi:10.1006/meth.2001.1262. pmid: 11846609
[37]	Fukuda A, Nakamura A, Hara N, Toki S, Tanaka Y. Molecular and functional analyses of rice NHX-type Na⁺/H⁺ antiporter genes[J]. Planta, 2011, 233(1):175-188.doi:10.1007/s00425-010-1289-4.
[38]	Ji Y H, Liu Z, Liu C, Shao Z Y, Zhang N, Suo M Q, Liu Y H, Wang L. Genome-wide identification and drought stress-induced expression analysis of the NHX gene family in potato[J]. Frontiers in Genetics, 2024, 15:1396375.doi:10.3389/fgene.2024.1396375.
[39]	罗建, 许春苗, 张国斌, 郁继华. 辣椒NHX基因家族的鉴定和表达分析[J]. 华北农学报, 2021, 36(3): 15-24. doi: 10.7668/hbnxb.20191999.
	Luo J, Xu C M, Zhang G B, Yu J H. Bioinformation analysis and expression analysis of NHX genes family in pepper[J]. Acta Agriculturae Boreali-Sinica, 2021, 36(3): 15-24. doi: 10.7668/hbnxb.20191999
[40]	Wang Y, Ying J L, Zhang Y, Xu L, Zhang W T, Ni M, Zhu Y L, Liu L W. Genome-wide identification and functional characterization of the cation proton antiporter(CPA)family related to salt stress response in radish(Raphanus sativus L.)[J]. International Journal of Molecular Sciences, 2020, 21(21):8262.doi:10.3390/ijms21218262.
[41]	杨仕梅, 谢恩俊, 罗春芳, 饶萍, 欧珍贵, 杨龙. 木薯NHX基因家族鉴定及特征分析[J]. 贵州农业科学, 2024, 52(8):19-26.doi:10.3969/j.issn.1001-3601.2024.08.003.
	Yang S M, Xie E J, Luo C F, Rao P, Ou Z G, Yang L. Identification and characteristics analysis of NHX gene family in Manihot esculenta[J]. Guizhou Agricultural Sciences, 2024, 52(8):19-26.
[42]	Parveen K, Abu Bakar Saddique M, Rehman S U, Ali Z, Aziz I, Shamsi I H, Muneer M A. Identification and characterization of salt stress-responsive NHX gene family in chickpea[J]. Plant Stress, 2023, 10:100266.doi:10.1016/j.stress.2023.100266.
[43]	邱全胜. 拟南芥NHX5和NHX6:离子平衡与蛋白质运输[J]. 中国科学(生命科学), 2017, 47(8):839-846.doi:10.1360/N052016-00351.
	Qiu Q S. Arabidopsis NHX5 and NHX6:ion homeostasis and protein transport[J]. Scientia Sinica(Vitae), 2017, 47(8):839-846.
[44]	Ghorbel M, Brini F, Sharma A, Landi M. Role of jasmonic acid in plants:the molecular point of view[J]. Plant Cell Reports, 2021, 40(8):1471-1494.doi:10.1007/s00299-021-02687-4. pmid: 33821356
[45]	Pan J J, Hu Y R, Wang H P, Guo Q, Chen Y N, Howe G A, Yu D Q. Molecular mechanism underlying the synergetic effect of jasmonate on abscisic acid signaling during seed germination in Arabidopsis[J]. The Plant Cell, 2020, 32(12):3846-3865.doi:10.1105/tpc.19.00838.
[46]	高玉龙, 宋中邦, 李梅云, 李文正, 王丙武, 李永平. 烟草NtNHX1-3基因的克隆及表达特性[J]. 西北植物学报, 2018, 38(12):2201-2206.doi:10.7606/j.issn.1000-4025.2018.12.2201.
	Gao Y L, Song Z B, Li M Y, Li W Z, Wang B W, Li Y P. Cloning and expression characteristics of tobacco NtNHX1-3 gene[J]. Acta Botanica Boreali-Occidentalia Sinica, 2018, 38(12):2201-2206.
[47]	刘佳欣, 朱建峰, 杨秀艳, 张会龙, 张华新. 木本盐生植物西伯利亚白刺NHX基因家族全基因组鉴定与表达分析[J]. 基因组学与应用生物学, 2023, 42(7):698-714.doi:10.13417/j.gab.042.000698.
	Liu J X, Zhu J F, Yang X Y, Zhang H L, Zhang H X. Genome identification and expression analysis of NHX gene family in woody halophyte Nitraria sibirica[J]. Genomics and Applied Biology, 2023, 42(7):698-714.
[48]	Wu G Q, Wang J L, Li S J. Genome-wide identification of Na⁺/H⁺ antiporter(NHX)genes in sugar beet(Beta vulgaris L.) and their regulated expression under salt stress[J]. Genes, 2019, 10(5):401.doi:10.3390/genes10050401.

选择文件类型/文献管理软件名称

选择包含的内容