桃SAP基因家族鉴定及在非生物胁迫下砧木中的缓解响应

doi:10.16420/j.issn.0513-353x.2025-0634

摘要/Abstract

摘要： 采用生物信息方法对桃SAP基因进行鉴定，明确其结构特征和在逆境缓解后砧木中的表达模式。结果表明，在桃基因组中共鉴定出了11个PpSAP家族成员，分子量为16 268.42 Da（PpSAP3） ~ 31 983.27 Da（PpSAP8），只有3个成员（PpSAP4、PpSAP8和PpSAP9）含有内含子，亚细胞定位预测显示大部分成员定位于细胞核。系统发育树分析发现，PpSAP蛋白可分为ⅠA、ⅠC、ⅡA和ⅡB 4类。PpSAP家族多数成员结构域为A20-AN1，同结构域成员具有相似的Motif分布。顺式作用元件分析表明，PpSAP主要与光、激素和非生物胁迫反应有关。此外，串联重复事件有助于PpSAP基因的扩张。通过GO富集分析鉴定了PpSAP的13个GO ID，蛋白互作网络显示7个成员与4个蛋白存在互作。miRNA互作分析鉴定出175个靶向PpSAP蛋白的miRNA，其中miR172家族与多个成员存在特异性互作。对非生物胁迫缓解后PpSAP在桃砧木（白花山桃、中桃抗砧1号、SH、GF677）和樱桃李砧木（李1、李3、李5、RA）中的表达进行qRT-PCR分析显示，干旱胁迫下，PpSAP4在李3、GF677 + Spd和GF677 + PEG缓解处理后均高表达，PpSAP6在李1 + Spd处理后高表达，PpSAP7、PpSAP8在李3 + PEG及PpSAP9在SH + Spd处理后显著表达。低温胁迫下，PpSAP3在李5 + ALA、RA + ALA、RA + Spd和SH、RA + PEG缓解处理后显著表达，PpSAP4在GF677 + PEG处理后最高表达（1 404.05），PpSAP8在李5、RA的ALA和Spd处理后高表达。盐碱胁迫下，PpSAP1在中桃抗砧1号 + ALA缓解处理后高表达，PpSAP6在李1 + ALA处理下显著表达，而PpSAP9在SH + ALA和GF677 + PEG处理后均显著表达。说明外源缓解物质ALA、Spd和PEG能特异性诱导关键PpSAP的高表达，其调控效果因砧木类型与胁迫环境的不同而存在显著差异。综上，推测PpSAP可能是缓解非生物胁迫的关键基因家族。

关键词: 桃, 砧木, SAP基因家族, 生物信息学, 非生物胁迫

Abstract:

The peach SAP genes were identified using bioinformatics methods to determine their structural characteristics and expression patterns following stress alleviation. Results showed that 11 PpSAP family members were identified in the peach genome，with molecular weights ranging from 16 268.42 Da（PpSAP3）to 31 983.27 Da（PpSAP8）. Only three genes PpSAP4，PpSAP8，and PpSAP9 contained introns. Subcellular localization predictions indicated that most members are localized in the nucleus. Phylogenetic analysis revealed that PpSAP proteins can be classified into four groups：ⅠA，ⅠC，ⅡA，and ⅡB. Most PpSAP family members contain A20-AN1 domains，and proteins within the same group exhibit similar motif distributions. cis-Acting element analysis suggested that PpSAPs are primarily associated with light，hormone，and abiotic stress responses. Moreover，tandem duplication events contributed to the expansion of the PpSAP gene family. GO enrichment analysis identified 13 GO terms for PpSAPs，while a protein-protein interaction network indicated interactions between seven PpSAP members and four other proteins. miRNA interaction analysis identified 175 miRNAs targeting PpSAP proteins，among which the miR172 family showed specific interactions with multiple PpSAP members. qRT-PCR analysis of PpSAP expression in peach rootstocks（Baihua Shantao，Zhongtao Kangzhen 1，SH，GF677）and Prunus cerasifera rootstocks（Li 1，Li 3，Li 5，RA）after mitigation of abiotic stress revealed that under drought stress，PpSAP4 showed high expression in Li 3 and in GF677 treated with Spd or PEG，PpSAP6 was highly expressed in Li 1 treated with Spd，while PpSAP7 and PpSAP8 were significantly upregulated in Li 3 treated with PEG，and PpSAP9 was notably expressed in SH treated with Spd. Under low-temperature stress，PpSAP3 exhibited significant expression in Li 5 and RA treated with ALA，RA treated with Spd，and SH and RA treated with PEG；PpSAP4 reached its highest expression level（1 404.05）in GF677 treated with PEG；PpSAP8 showed high expression in Li 5 and RA after treatment with ALA or Spd. Under saline-alkali stress，PpSAP1 was highly expressed in Zhongtao Kangzhen 1 treated with ALA，PpSAP6 was significantly upregulated in Li 1 treated with ALA，while PpSAP9 showed notable expression in both SH treated with ALA and GF677 treated with PEG. These results indicate that the exogenous mitigators ALA，Spd，and PEG can specifically induce high expression of key PpSAP genes，and their regulatory effects vary significantly depending on the rootstock type and stress environment. In conclusion，PpSAP is hypothesized to be a key gene family involved in mitigating abiotic stress.

Key words: peach, rootstock, SAP gene family, bioinformatics, abiotic stress

任家玄, 张帆, 王晨冰. 桃SAP基因家族鉴定及在非生物胁迫下砧木中的缓解响应[J]. 园艺学报, 2026, 53(3): 635-652.

REN Jiaxuan, ZHANG Fan, WANG Chenbing. Identification of the Peach SAP Gene Family and Alleviative Response in Rootstocks Under Abiotic Stress[J]. Acta Horticulturae Sinica, 2026, 53(3): 635-652.

图/表 11

表1 PpSAP家族成员引物信息

Table 1 Primer information of PpSAP family members

基因 Gene	上游引物（5′-3′） Forward primers	下游引物（5′-3′） Reverse primers
PpSAP1	CCCCGAAACTCTAACGCACT	AGATGAGGTTGCTGTGGTGG
PpSAP2	CCAAACAGGTGCGGAACTTG	CCTGAGCAGCACTTCGGTAA
PpSAP3	GGCTGTGGCTTTTATGGCTG	TCCTCTAGCGATGGAACCCA
PpSAP4	ACGCCTTCCTCATCGTTGTT	TCGCTCTTCCCGCATTCTTT
PpSAP5	CAAAGCTCGCTGCATCATCC	CCAGCAGCAACAACAGGTTC
PpSAP6	GTGCCAGGCTCCACAACTAT	TGAGCTGCAAGTCACGGTAG
PpSAP7	GAGAACCCTGAGAAGCGACC	AATCGAACACGCAGTTGTGC
PpSAP8	GATCTGGGTCGGCATTGCTA	GGCACAAAGGGGACAGATGA
PpSAP9	TGGGAGTAGAAACGGGCAAC	CTGCTGGATCACGAAACCCT
PpSAP10	AAGAGGGACCAAGCAGATGC	GCATCTTGACCAGCAGTCCT
PpSAP11	ACCGGGACTTTTGCCTCAAA	AGGAGCAGAGACAGAGGAGG
Actin	GATTCCGGTGCCCAGAAGT	CCAGCAGCTTCCATTCCAA

表2 桃SAP家族成员基本信息

Table 2 Basic information of peach SAP family members

基因 Gene	登录号 Gene ID	氨基酸数量 No. of Amino acid	相对分子质量/Da Molecular weight	等电点 pI	锌指结构 Zinc-finger	内含子数量 No. of introns	蛋白类型 Protein type	亚细胞定位 Subcellular localization
PpSAP1	Prupe.2G010300.1	165	18 190.51	9.40	A20-AN1	0	ⅠA	细胞核 Nuclear
PpSAP2	Prupe.2G010400.1	172	18 479.00	7.95	A20-AN1	0	ⅠA	胞外 Extracellular
PpSAP3	Prupe.2G145200.1	149	16 268.42	8.55	A20-AN1	0	ⅠA	细胞核 Nuclear
PpSAP4	Prupe.3G278700.1	192	21 620.69	9.01	AN1-AN1	1	ⅡB	细胞核 Nuclear
PpSAP5	Prupe.6G023300.1	173	18 483.90	7.55	A20-AN1	0	ⅠA	细胞核 Nuclear 胞外 Extracellular
PpSAP6	Prupe.6G127400.1	162	18 064.56	8.81	A20-AN1	0	ⅠA	细胞核 Nuclear
PpSAP7	Prupe.6G168300.1	168	18 831.31	8.98	A20-AN1	0	ⅠA	细胞核 Nuclear
PpSAP8	Prupe.7G063400.1	292	31 983.27	8.72	AN1-AN1-C2H2-C2H2	1	ⅡA	细胞核 Nuclear 胞外 Extracellular
PpSAP9	Prupe.7G063600.1	282	31 111.58	8.58	AN1-AN1-C2H2-C2H2	4	ⅡA	胞外 Extracellular
PpSAP10	Prupe.7G068900.1	172	18 502.89	6.79	A20-AN1	0	ⅠA	细胞核 Nuclear 胞外 Extracellular
PpSAP11	Prupe.7G113400.1	183	19 561.27	7.95	A20-AN1	0	ⅠA	细胞核 Nuclear

图1 PpSAP家族基因在染色体上的位置分布

Fig. 1 Chromosomal location distribution of PpSAP family genes

图2 番茄、拟南芥、水稻和桃SAP蛋白系统发育树

Fig. 2 Phylogenetic tree of SAP proteins in Solanum lycopersicum，Arabidopsis thaliana，Oryza sativa and Prunus persica

图3 PpSAP家族成员多序列比对红框依次代表A20、AN1和C2H2保守结构域

Fig. 3 Multiple sequence alignment of PpSAP family members The red boxes sequentially represent the conserved domains of A20，AN1，and C2H2

图4 PpSAP家族成员Motif（A）、结构域（B）和基因结构（C）

Fig. 4 Motifs（A），domains（B）and gene structures（C）of PpSAP family members

图5 PpSAP家族成员顺式作用元件类型（A）和分布（B）

Fig. 5 Types（A）and distribution（B）of cis-acting elements of PpSAP family members

图6 PpSAP家族成员之间（A）以及桃与番茄（Sl）、拟南芥（At）、水稻（Os）间（B）SAP基因的共线性

Fig. 6 Synteny analysis of SAP genes among PpSAP family members（A）and between Prunus persica，Solanum lycopersicum （Sl），Arabidopsis thaliana（At），and Oryza sativa（Os）（B）

图7 PpSAP家族成员GO富集（A）和蛋白互作网络（B）

Fig. 7 GO enrichment（A）and protein interaction network（B）of PpSAP family members

图8 PpSAP家族成员与互作miRNA

Fig. 8 Members of the PpSAP family and interacting miRNA

图9 桃砧木在自然干旱（A）、4 ℃低温（B）和盐碱胁迫（C）后外源物质缓解下的PpSAP表达水平外源缓解物质分别为清水（对照）、ALA 50 mg · L-1（T1）、Spd 1.0 mmol · L-1（T2）和PEG-6000 100 g · L-1（T3）

Fig. 9 Expression levels of PpSAP in peach rootstocks alleviated by exogenous substances after natural drought（A），4 ℃ low temperature （B），and saline-alkali stress（C） The exogenous alleviative substances were clean water（Control），ALA 50 mg · L-1（T1），Spd 1.0 mmol · L-1（T2），and PEG-6000 100 g · L-1（T3）

参考文献 43

[1]	An J, Li H, Liu Z, Wang D R, You C X, Han Y. 2023. The E3 ubiquitin ligase SINA1 and the protein kinase BIN2 cooperatively regulate PHR1 in apple anthocyanin biosynthesis. Journal of Integrative Plant Biology, 65 (9):2175-2193. doi: 10.1111/jipb.13538
[2]	Baidyussen A, Aldammas M, Kurishbayev A, Myrzabaeva M, Zhubatkanov A, Sereda G, Porkhun R, Sereda S, Jatayev S, Langridge P, Schramm C, Jenkins C L D, Soole K L, Shavrukov Y. 2020. Identification,gene expression and genetic polymorphism of zinc finger A20/AN1 stress-associated genes,HvSAP,in salt stressed barley from Kazakhstan. BMC Plant Biology, 20 (Suppl 1):156. doi: 10.1186/s12870-020-02332-4 pmid: 33050881
[3]	Barberis E, Marengo E, Manfredi M. 2021. Protein subcellular localization prediction. Methods in Molecular Biology,2361:197-212.
[4]	Billah S A, Khan N Z, Ali W, Aasım M, Usman M, Alezzawi A, Ullah H. 2022. Genome-wide in silico identification and characterization of the stress associated protein(SAP)gene family encoding A20/AN1 zinc-finger proteins in potato(Solanum tuberosum L.). PLoS ONE, 17 (8):e0273416. doi: 10.1371/journal.pone.0273416 URL
[5]	Capella-Gutierrez S, Silla-Martinez J M, Gabaldon T. 2009. trimAl:a tool for automated alignment trimming in large-scale phylogenetic analyses. Bioinformatics,25:1972-1973.
[6]	Chen C, Wu Y, Li J, Wang X, Zeng Z, Xu J, Liu Y, Feng J, Chen H, He Y, Xia R. 2023. TBtools-II:a“one for all,all for one”bioinformatics platform for biological big-data mining. Molecular Plant, 16 (11):1733-1742. doi: 10.1016/j.molp.2023.09.010 URL
[18]	Li Yuxin, Li Miao, Du Xiaofen, Han Kangni, Lian Shichao. 2025. Identification and expression analysis of SiSAP gene family in foxtail millet(Setaria italica). Biotechnology Bulletin, 41 (1):143-156. (in Chinese) doi: 10.13560/j.cnki.biotech.bull.1985.2024-0576
	李禹欣, 李苗, 杜晓芬, 韩康妮, 连世超. 2025. 谷子SiSAP基因家族的鉴定与表达分析. 生物技术通报, 41 (1):143-156. doi: 10.13560/j.cnki.biotech.bull.1985.2024-0576
[19]	Liu S, Wang J, Jiang S, Wang H, Gao Y, Zhang H, Li D, Song F. 2019. Tomato SlSAP3,a member of the stress-associated protein family,is a positive regulator of immunity against Pseudomonas syringae pv. tomato DC300. Molecular Plant Pathology, 20 (6):815-830. doi: 10.1111/mpp.2019.20.issue-6 URL
[20]	Livak K J, Schmittgen T D. 2001. Analysis of relative gene expression data using real-time quantitative PCR and the 2^-ΔΔCT method. Methods, 25 (4):402-408. doi: 10.1006/meth.2001.1262 pmid: 11846609
[21]	Lu S, Zhuge Y, Hao T, Liu Z, Zhang M, Fang J. 2022. Systematic analysis reveals O-methyltransferase gene family members involved in flavonoid biosynthesis in grape. Plant Physiology and Biochemistry,173:33-45.
[22]	Lu Y Y, Fricke W. 2023. Salt stress-regulation of root water uptake in a whole-plant and diurnal context. International Journal of Molecular Sciences, 24 (9):8070. doi: 10.3390/ijms24098070 URL
[23]	Mittler R, Zandalinas S I, Fichman Y, Breusegem F V. 2022. Reactive oxygen species signalling in plant stress responses. Nature Reviews Molecular Cell Biology, 23 (10):663-679. doi: 10.1038/s41580-022-00499-2
[24]	Mukhopadhyay A S, Vij S, Tyagi A K. 2004. Overexpression of a zinc-finger protein gene from rice confers tolerance to cold,dehydration,and salt stress in transgenic tobacco. Proceedings of the National Academy of Sciences of the United States of America, 101 (16):6309-6314.
[25]	Nguyen L T, Schmidt H A, von Haeseler A, Minh B Q. 2015. IQ-TREE:a fast and efective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evolution, 32 (1):268-274. doi: 10.1093/molbev/msu300 URL
[7]	Chenna R, Sugawara H, Koike T, Lopez R, Gibson T J, Higgins D G, Thompson J D. 2003. Multiple sequence alignment with the Clustal series of programs. Nucleic Acids Research, 31 (13):3497-3500. doi: 10.1093/nar/gkg500 pmid: 12824352
[8]	Ding Lan, Gu Bao, Li Peiying, Shu Xin, Zhang jianxia. 2019. Genome-wide identification and expression analysis of SAP family in grape. Scientia Agricultura Sinica, 52 (14):2500-2514. (in Chinese)
	丁兰, 顾宝, 李培楹, 舒欣, 张剑侠. 2019. 葡萄SAP家族的鉴定与表达分析. 中国农业科学, 52 (14):2500-2514. doi: 10.3864/j.issn.0578-1752.2019.14.009
[9]	Dong Q, Duan D, Zhao S, Xu B, Luo J, Wang Q, Huang D, Liu C, Li C, Gong X, Mao K, Ma F. 2018. Genome-wide analysis and cloning of the apple stress-associated protein gene family reveals MdSAP15,which confers tolerance to drought and osmotic stresses in transgenic Arabidopsis. International Journal of Molecular Sciences, 19 (9):2478. doi: 10.3390/ijms19092478 URL
[10]	Fattel L, Psaroudakis D, Yanarella C F, Chiteri K O, Dostalik H A, Joshi P, Starr D C, Vu H T H, Wimalanathan K, Lawrence‐Dill C J. 2022. Standardized genome-wide function prediction enables comparative functional genomics:a new application area for gene ontologies in plants. GigaScience,11:1-18.
[11]	Gimeno-Gilles C, Gervais M L, Planchet E, Satour P, Limami A M, Lelievre E. 2011. A stress-associated protein containing A20/AN1 zing-finger domains expressed in Medicago truncatula seeds. Plant Physiology and Biochemistry, 49 (3):303-310. doi: 10.1016/j.plaphy.2011.01.004 pmid: 21296585
[12]	Giri J, Dansana P K, Kothari K S, Sharma G, Vij S, Tyagi A K. 2013. SAPs as novel regulators of abiotic stress response in plants. BioEssays, 35 (7):639-648. doi: 10.1002/bies.201200181 pmid: 23640876
[26]	Opipari A W, Boguski M S, Dixit V M. 1990. The A20 cDNA induced by tumor necrosis factor alpha encodes a novel type of zinc finger protein. Journal of Biological Chemistry, 265 (25):14705-14708. pmid: 2118515
[27]	Saad R B, Romdhane W B, Čmiková N, Baazaoui N, Bouteraa M T, Akacha B B, Chouaibi Y, Maisto M, Hsouna A B, Garzoli S, Wiszniewska A, Kačániová M. 2024. Research progress on plant stress-associated protein(SAP)family:master regulators to deal with environmental stresses. BioEssays, 46 (11):2400097. doi: 10.1002/bies.v46.11 URL
[28]	Saad R B, Zouari N, Ramdhan W B, Azaza J, Meynard D, Guiderdoni E, Hassair A. 2010. Improved drought and salt stresstolerance in transgenic tobacco overexpressing a novel A20/AN1zinc-finger“AlSAP”gene isolated from the halophyte grass Aeluropus littoralis. Plant Molecular Biology,72:171-190.
[29]	Shukla V, Choudhary P, Rana S, Muthamilarasan M. 2021. Structural evolution and function of stress associated proteins in regulating biotic and abiotic stress responses in plants. Journal of Plant Biochemistry and Biotechnology, 30 (4):779-792. doi: 10.1007/s13562-021-00704-x
[30]	Soma F, Takahashi F, Yamaguchi‐Shinozaki K, Shinozaki K. 2021. Cellular phosphorylation signaling and gene expression in drought stress responses:ABA-dependent and ABA-independent regulatory systems. Plants, 10 (4):756. doi: 10.3390/plants10040756 URL
[31]	Sreedharan S, Shekhawat U K S, Ganapathi T R. 2012. MusaSAP1,a A20/AN1 zinc finger gene from banana functions as a positive regulator in different stress responses. Plant Molecular Biology,80:503-517.
[32]	Su A, Qin Q, Liu C, Zhang J, Yu B, Cheng Y, Wang S, Tang J, Si W. 2022. Identification and analysis of stress-associated proteins(SAPs)protein family and drought tolerance of ZmSAP8 in transgenic Arabidopsis. International Journal of Molecular Sciences, 23 (22):14109. doi: 10.3390/ijms232214109 URL
[33]	Tang D, Chen M, Huang X, Zhang G, Zeng L, Zhang G, Wu S, Wang Y. 2023. SRplot:a free online platform for data visualization and graphing. PLoS ONE, 18 (11):e0294236. doi: 10.1371/journal.pone.0294236 URL
[34]	Tyagi H, Jha S, Sharma M, Giri J, Tyagi A K. 2014. Rice SAPs are responsive to multiple biotic stresses and overexpression of OsSAP1,an A20/AN1 zinc-finger protein,enhances the basal resistance against pathogen infection in tobacco. Plant Science,225:68-76.
[35]	Verde I, Abbott A G, Scalabrin S, Jung S, Shu S, Marroni F. 2013. The high-quality draft genome of peach(Prunus persica)identifies unique patterns of genetic diversity,domestication and genome evolution. Nature Genetics, 45 (5):487-494. doi: 10.1038/ng.2586
[36]	Verde I, Jenkins J, Dondini L, Micali S, Pagliarani G, Vendramin E. 2017. The peach v2.0 release:high-resolution linkage mapping and deep resequencing improve chromosome-scale assembly and contiguity. BMC Genomics,18:225.
[37]	Vij S, Tyagi A K. 2006. Genome-wide analysis of the stress associated protein(SAP)gene family containing A20/AN1 zinc-finger(s)in rice and their phylogenetic relationship with Arabidopsis. Molecular Genetics and Genomics, 276 (6):565-575. doi: 10.1007/s00438-006-0165-1 URL
[38]	Wan F, Xu Y, Wang S, Gao J, Lu D, Zhou C, Liao Y, Ma Y, Zheng Y. 2022. Identification and expression analysis of zinc finger A20/AN1 stress-associated genes SmSAP responding to abiotic stress in eggplant. Horticulturae, 8 (2):108. doi: 10.3390/horticulturae8020108 URL
[39]	Wang H, Zhang S, Wang Z, Li D, Yan L, Feng Y, Liu X, Chen R, Fan W, Sun L, Zhao Z. 2024. Resistance index and browning mechanism of apple peel under high temperature stress. Horticultural Plant Journal, 10 (2):305-317. doi: 10.1016/j.hpj.2022.10.013 URL
[13]	Huang J, Teng L, Li L X, Liu T, Li L Y, Chen D Y, Xu L G, Zhai Z H, Shu H B. 2004. ZNF 216 is an A20-like and IκB kinase γ-interacting inhibitor of NFκB activation. Journal of Biological Chemistry, 279 (16):16847-16853. doi: 10.1074/jbc.M309491200 pmid: 14754897
[14]	Huang Y, Du L, Wang M, Ren M, Yu S, Yang Q. 2022. Multifaceted roles of zinc finger proteins in regulating various agronomic traits in rice. Frontiers in Plant Science,13:974396.
[15]	Jeffares D C, Penkett C J, Bähler J. 2008. Rapidly regulated genes are intron poor. Trends in Genetics, 24 (8):375-378. doi: 10.1016/j.tig.2008.05.006 pmid: 18586348
[16]	Kang M, Lee S, Abdelmageed H, Reichert A, Lee H K, Fokar M, Mysore K S, Allen R D. 2017. Arabidopsis stress associated protein 9 mediates biotic and abiotic stress responsive ABA signaling via the proteasome pathway. Plant,Cell and Environment, 40 (5):702-716. doi: 10.1111/pce.v40.5 URL
[17]	Li Y, Pan H, Hao Y, Zhu Y, Chang Q, Ren J. 2024. Genome-wide identification and expression analysis of the NRL(NPH3/RPT2-Like)family in Vitis vinifera. Journal of Plant Growth Regulation,44:2106-2121.
[40]	Wang K, Ding Y, Cai C, Chen Z, Zhu C. 2019. The role of C2H 2 zinc finger proteins in plant responses to abiotic stresses. Physiologia Plantarum, 165 (4):690-700. doi: 10.1111/ppl.12728
[41]	Wang Yinghua, Wang Xiaojing. 2011. Relationship between stress associated proteins and stress tolerance in plants. Journal of South China Normal University(Natural Science Edition),(1):9-12,18. (in Chinese)
	王瑛华, 王小菁. 2011. 胁迫相关蛋白(SAP)与植物的抗逆性. 华南师范大学学报(自然科学版),(1):9-12,18.
[42]	Zhang L, Li X, Li D, Sun Y, Liu Y, Luo Q, Liu Z, Wang J, Li X, Zhang H, Lou Z, Yang Y. 2018. CARK 1 mediates ABA signaling by phosphorylation of ABA receptors. Cell Discovery,4:30.
[43]	Zhao X, Wang R, Zhang Y, Li Y, Yue Y, Zhou T, Wang C. 2021. Comprehensive analysis of the stress associated protein(SAP)gene family in Tamarix hispida and the function of ThSAP6 in salt tolerance. Plant Physiology and Biochemistry,165:1-9.