Roles of Plasma Membrane Na+/H+ Antiporter SOS1 in Maintaining Ionic Homeostasis of Plants

doi:10.13560/j.cnki.biotech.bull.1985.2023-0793

Abstract

Abstract:

Plants regulate ion homeostasis to adapt to saline environment through a series of complex transport systems. SOS（salt over sensitive）signal pathway is the major signal pathway for plants to respond to abiotic stress, which is mainly composed of plasma membrane Na⁺/H⁺ antiporter SOS1, serine/threonine protein kinase SOS2, and calcium sensor SOS3. As one of the main members of SOS signaling pathway, SOS1 widely exists in higher plants. Due to the early evolutionary differences, the structural, physical, and chemical properties of SOS1 from different species had certain specificity. The SOS1 protein is a homodimer, and each monomer is composed of transmembrane and intracellular domains, which provides a stable docking platform for integrating signals from different pathways and regulating Na⁺ transport. Transcription level of the SOS1 gene was regulated by different stress conditions. SOS1 activity was inhibited or activated through Ca²⁺ signal regulation, phosphorylation, self-inhibition, and synergistic regulation with other ion transporters. SOS1 has shown to regulate circadian rhythm and pH, and maintain ion homeostasis in plants, which plays an important role in the response of plant to abiotic stress. In this review, the structure, function, regulation mechanism of SOS1 and its role in maintaining plant ion homeostasis are reviewed, and its future research direction is also prospected. The information in this review provide a theoretical support and excellent genetic resources for the genetic improvement of crops to produce new, stress-resistant varieties.

Key words: SOS1, Na⁺ extrusion, Na⁺ long-distance transport, ion homeostasis, SOS signaling pathway, salt tolerance, oxidative stress

ZHU Ye-sheng, WU Guo-qiang, WEI Ming. Roles of Plasma Membrane Na⁺/H⁺ Antiporter SOS1 in Maintaining Ionic Homeostasis of Plants[J]. Biotechnology Bulletin, 2023, 39(12): 16-32.

Figures/Tables 4

References 124

[1]	Ma L, Liu XH, Lv WJ, et al. Molecular mechanisms of plant responses to salt stress[J]. Front Plant Sci, 2022, 13: 934877. doi: 10.3389/fpls.2022.934877 URL
[2]	Foster KJ, Miklavcic SJ. A comprehensive biophysical model of ion and water transport in plant roots. II. clarifying the roles of SOS1 in the salt-stress response in Arabidopsis[J]. Front Plant Sci, 2019, 10: 1121. doi: 10.3389/fpls.2019.01121 pmid: 31620152
[3]	Zhu JK. Salt and drought stress signal transduction in plants[J]. Annu Rev Plant Biol, 2002, 53: 247-273. doi: 10.1146/arplant.2002.53.issue-1 URL
[4]	Zhu M, Shabala L, Cuin TA, et al. Nax loci affect SOS1-like Na⁺/H⁺ exchanger expression and activity in wheat[J]. J Exp Bot, 2016, 67(3): 835-844. doi: 10.1093/jxb/erv493 URL
[5]	Ishikawa T, Shabala S. Control of xylem Na⁺ loading and transport to the shoot in rice and barley as a determinant of differential salinity stress tolerance[J]. Physiol Plant, 2019, 165(3): 619-631. doi: 10.1111/ppl.12758 pmid: 29761494
[6]	Wu HH, Zhang XC, Giraldo JP, et al. It is not all about sodium: revealing tissue specificity and signalling roles of potassium in plant responses to salt stress[J]. Plant Soil, 2018, 431(1): 1-17. doi: 10.1007/s11104-018-3770-y
[7]	Isayenkov SV, Maathuis FJM. Plant salinity stress: many unanswered questions remain[J]. Front Plant Sci, 2019, 10: 80. doi: 10.3389/fpls.2019.00080 pmid: 30828339
[8]	Rubio F, Nieves-Cordones M, Horie T, et al. Doing ‘business as usual’ comes with a cost: evaluating energy cost of maintaining plant intracellular K⁺ homeostasis under saline conditions[J]. New Phytol, 2020, 225(3): 1097-1104. doi: 10.1111/nph.v225.3 URL
[9]	Wu N, Li Z, Wu F, et al. Sex-specific photosynthetic capacity and Na⁺ homeostasis in Populus euphratica exposed to NaCl stress and AMF inoculation[J]. Front Plant Sci, 2022, 13: 1066954. doi: 10.3389/fpls.2022.1066954 URL
[10]	Gul Z, Tang ZH, Arif M, et al. An insight into abiotic stress and influx tolerance mechanisms in plants to cope in saline environments[J]. Biology, 2022, 11(4): 597. doi: 10.3390/biology11040597 URL
[11]	Gupta BK, Sahoo KK, Anwar K, et al. Silicon nutrition stimulates salt-overly sensitive(SOS)pathway to enhance salinity stress tolerance and yield in rice[J]. Plant Physiol Biochem, 2021, 166: 593-604. doi: 10.1016/j.plaphy.2021.06.010 URL
[12]	DeTar RA, Höhner R, Manavski N, et al. Loss of salt overly sensitive 1 prevents virescence in chloroplast K⁺/H⁺ efflux antiporter-deficient mutants[J]. Plant Physiol, 2022, 189(3): 1220-1225. doi: 10.1093/plphys/kiac142 URL
[13]	Wang YH, Pan CC, Chen QH, et al. Architecture and autoinhibitory mechanism of the plasma membrane Na⁺/H⁺ antiporter SOS1 in Arabidopsis[J]. Nat Commun, 2023, 14(1): 4487. doi: 10.1038/s41467-023-40215-y
[14]	Ponce KS, Meng LJ, Guo LB, et al. Advances in sensing, response and regulation mechanism of salt tolerance in rice[J]. Int J Mol Sci, 2021, 22(5): 2254. doi: 10.3390/ijms22052254 URL
[15]	Zhu JK, Liu J, Xiong L. Genetic analysis of salt tolerance in Arabidopsis. Evidence for a critical role of potassium nutrition[J]. Plant Cell, 1998, 10(7): 1181-1191. doi: 10.1105/tpc.10.7.1181 pmid: 9668136
[16]	Shi H, Ishitani M, Kim C, et al. The Arabidopsis thaliana salt tolerance gene SOS1 encodes a putative Na⁺/H⁺ antiporter[J]. Proc Natl Acad Sci USA, 2000, 97(12): 6896-6901. doi: 10.1073/pnas.120170197 pmid: 10823923
[17]	Wu SJ, Zhu JK. SOS1, a genetic locus essential for salt tolerance and potassium acquisition[J]. Plant Cell, 1996, 8(4): 617-627. doi: 10.2307/3870339 URL
[18]	Shi HZ, Quintero FJ, Pardo JM, et al. The putative plasma membrane Na⁺/H⁺ antiporter SOS1 controls long-distance Na⁺ transport in plants[J]. Plant Cell, 2002, 14(2): 465-477. doi: 10.1105/tpc.010371 URL
[19]	Che BN, Cheng C, Fang JJ, et al. The recretohalophyte Tamarix TrSOS1 gene confers enhanced salt tolerance to transgenic hairy root composite cotton seedlings exhibiting virus-induced gene silencing of GhSOS1[J]. Int J Mol Sci, 2019, 20(12): 2930. doi: 10.3390/ijms20122930 URL
[20]	时丕彪, 洪立洲, 王军, 等. 印度南瓜Na⁺/H⁺逆向转运蛋白基因CmaSOS1的克隆与表达分析[J]. 核农学报, 2020, 34(12): 2638-2646. doi: 10.11869/j.issn.100-8551.2020.12.2638
	Shi PB, Hong LZ, Wang J, et al. Cloning and expression analysis of a Na⁺/H⁺antiporter gene CmaSOS1 in Cucurbita maxima[J]. J Nucl Agric Sci, 2020, 34(12): 2638-2646.
[21]	Shahzad B, Shabala L, Zhou MX, et al. Comparing essentiality of SOS1-mediated Na⁺ exclusion in salinity tolerance between cultivated and wild rice species[J]. Int J Mol Sci, 2022, 23(17): 9900. doi: 10.3390/ijms23179900 URL
[22]	El Mahi H, Pérez-Hormaeche J, Luca AD, et al. A critical role of sodium flux via the plasma membrane Na⁺/H⁺ exchanger SOS1 in the salt tolerance of rice[J]. Plant Physiol, 2019, 180(2): 1046-1065. doi: 10.1104/pp.19.00324 URL
[23]	Tomita M, Yamashita M, Omichi A. Gene structure of three kinds of vacuolar-type Na⁺/H⁺ antiporters including TaNHX2 transcribed in bread wheat[J]. Genet Mol Biol, 2021, 44(1): e20200207. doi: 10.1590/1678-4685-gmb-2020-0207 URL
[24]	Jiang W, Pan R, Buitrago S, et al. Conservation and divergence of the TaSOS1 gene family in salt stress response in wheat(Triticum aestivum L.)[J]. Physiol Mol Biol Plants, 2021, 27(6): 1245-1260. doi: 10.1007/s12298-021-01009-y
[25]	Li SJ, Wu GQ, Lin LY. AKT1, HAK5, SKOR, HKT1;5, SOS1 and NHX1 synergistically control Na⁺ and K⁺ homeostasis in sugar beet(Beta vulgaris L.)seedlings under saline conditions[J]. J Plant Biochem Biotechnol, 2022, 31(1): 71-84. doi: 10.1007/s13562-021-00656-2
[26]	Zhang WT, Li JX, Dong JH, et al. RsSOS1 responding to salt stress might be involved in regulating salt tolerance by maintaining Na⁺ homeostasis in radish(Raphanus sativus L.)[J]. Horticulturae, 2021, 7(11): 458. doi: 10.3390/horticulturae7110458 URL
[27]	张婷婷, 康宇乾, 李雨欣, 等. 非生物胁迫下海马齿SpSOS1基因的表达及响应ABA模式分析[J]. 分子植物育种, 2021, 19(13): 4371-4377.
	Zhang TT, Kang YQ, Li YX, et al. Expression pattern analysis of SpSOS1 from Sesuvium portulacastrum under abiotic stresses and the response to ABA[J]. Mol Plant Breed, 2021, 19(13): 4371-4377.
[28]	Zhao CY, William D, Sandhu D. Isolation and characterization of salt overly sensitive family genes in spinach[J]. Physiol Plant, 2021, 171(4): 520-532. doi: 10.1111/ppl.v171.4 URL
[29]	Brindha C, Vasantha S, Raja AK, et al. Characterization of the salt overly sensitive pathway genes in sugarcane under salinity stress[J]. Physiol Plant, 2021, 171(4): 677-687. doi: 10.1111/ppl.13245 pmid: 33063359
[30]	Yang YQ, Han XL, Ma L, et al. Dynamic changes of phosphatidylinositol and phosphatidylinositol 4-phosphate levels modulate H⁺-ATPase and Na⁺/H⁺ antiporter activities to maintain ion homeostasis in Arabidopsis under salt stress[J]. Mol Plant, 2021, 14(12): 2000-2014. doi: 10.1016/j.molp.2021.07.020 URL
[31]	Xie Q, Zhou Y, Jiang XY. Structure, function, and regulation of the plasma membrane Na⁺/H⁺ antiporter salt overly sensitive 1 in plants[J]. Front Plant Sci, 2022, 13: 866265. doi: 10.3389/fpls.2022.866265 URL
[32]	Zhang MH, Cao JF, Zhang TX, et al. A putative plasma membrane Na⁺/H⁺ antiporter GmSOS1 is critical for salt stress tolerance in Glycine max[J]. Front Plant Sci, 2022, 13: 870695. doi: 10.3389/fpls.2022.870695 URL
[33]	陈叶, 丁健, 阮成江, 等. 文冠果Na⁺/H⁺逆向转运蛋白基因XsSOS1的克隆与表达分析[J]. 分子植物育种, 2022. http://kns.cnki.net/kcms/detail/46.1068.S.20220803.1806.016.html.
	Chen Y, Ding J, Ruan CJ, et al. Cloning and expression analysis of a Na⁺/H⁺ antiporter gene XsSOS1 in Xanthoceras sorbifolium[J]. Mol Plant Breed, 2022. http://kns.cnki.net/kcms/detail/46.1068.S.20220803.1806.016.html.
[34]	Arciniegas Vega JP, Melino VJ. Uncovering natural genetic variants of the SOS pathway to improve salinity tolerance in maize[J]. New Phytol, 2022, 236(2): 313-315. doi: 10.1111/nph.18422 pmid: 35977055
[35]	Xu FC, Wang MJ, Guo YW, et al. The Na⁺/H⁺ antiporter GbSOS1 interacts with SIP5 and regulates salt tolerance in Gossypium barbadense[J]. Plant Sci, 2023, 330: 111658. doi: 10.1016/j.plantsci.2023.111658 URL
[36]	Liu ZY, Xie QJ, Tang FF, et al. The ThSOS3 gene improves the salt tolerance of transgenic Tamarix hispida and Arabidopsis thaliana[J]. Front Plant Sci, 2021, 11: 597480. doi: 10.3389/fpls.2020.597480 URL
[37]	Taji T, Komatsu K, Katori T, et al. Comparative genomic analysis of 1047 completely sequenced cDNAs from an Arabidopsis-related model halophyte, Thellungiella halophila[J]. BMC Plant Biol, 2010, 10: 261. doi: 10.1186/1471-2229-10-261 URL
[38]	Cosentino C, Fischer-Schliebs E, Bertl A, et al. Na⁺/H⁺ antiporters are differentially regulated in response to NaCl stress in leaves and roots of Mesembryanthemum crystallinum[J]. New Phytol, 2010, 186(3): 669-680. doi: 10.1111/j.1469-8137.2010.03208.x pmid: 20298477
[39]	Guo Q, Meng L, Han JW, et al. SOS1 is a key systemic regulator of salt secretion and K⁺/Na⁺ homeostasis in the recretohalophyte Karelinia caspia[J]. Environ Exp Bot, 2020, 177: 104098. doi: 10.1016/j.envexpbot.2020.104098 URL
[40]	Chen XG, Lu XK, Shu N, et al. GhSOS1, a plasma membrane Na⁺/H⁺ antiporter gene from upland cotton, enhances salt tolerance in transgenic Arabidopsis thaliana[J]. PLoS One, 2017, 12(7): e0181450. doi: 10.1371/journal.pone.0181450 URL
[41]	郑琳琳, 张慧荣, 贺龙梅, 等. 唐古特白刺质膜Na⁺/H⁺逆向转运蛋白基因的克隆与表达分析[J]. 草业学报, 2013, 22(4): 179-186. doi: 10.11686/cyxb20130422
	Zheng LL, Zhang HR, He LM, et al. Isolation and expression analysis of a plasma membrane Na⁺/H⁺ antiporter from Nitraria tangutorum[J]. Acta Prataculturae Sin, 2013, 22(4): 179-186.
[42]	黄坤勇, 李杉杉, 郭强, 等. 黄花草木樨MoSOS1基因克隆及表达分析[J]. 生物技术通报, 2017, 33(9): 120-130. doi: 10.13560/j.cnki.biotech.bull.1985.2017-0364
	Huang KY, Li SS, Guo Q, et al. Cloning and expression analysis of MoSOS1 gene in Melilotus officinalis[J]. Biotechnol Bull, 2017, 33(9): 120-130.
[43]	Wang JY, Li Q, Zhang M, et al. The high pH value of alkaline salt destroys the root membrane permeability of Reaumuria trigyna and leads to its serious physiological decline[J]. J Plant Res, 2022, 135(6): 785-798. doi: 10.1007/s10265-022-01410-y
[44]	荆海瑜, 周扬, 郭晓颖, 等. 木榄细胞膜Na⁺/H⁺逆向运输蛋白BgSOS1功能的初步验证[J]. 海南大学学报: 自然科学版, 2014, 32(3): 252-259, 269.
	Jing HY, Zhou Y, Guo XY, et al. Preliminary analysis of a Na⁺/H⁺ antiporter BgSOS1 at plasma membrane of Bruguiera gymnorrhiza[J]. Nat Sci J Hainan Univ, 2014, 32(3): 252-259, 269.
[45]	Gao JJ, Sun J, Cao PP, et al. Variation in tissue Na⁺ content and the activity of SOS1 genes among two species and two related Genera of Chrysanthemum[J]. BMC Plant Biol, 2016, 16: 98. doi: 10.1186/s12870-016-0781-9 URL
[46]	Venturini L, Ferrarini A, Zenoni S, et al. De novo transcriptome characterization of Vitis vinifera cv. Corvina unveils varietal diversity[J]. BMC Genomics, 2013, 14: 41. doi: 10.1186/1471-2164-14-41 pmid: 23331995
[47]	Wang S, Li Z, Rui R, et al. Cloning and characterization of a plasma membrane Na⁺/H⁺ antiporter gene from Cucumis sativus[J]. Russ J Plant Physiol, 2013, 60(3): 330-336. doi: 10.1134/S102144371303014X URL
[48]	Hu J, Hu XK, Zhang HW, et al. Moderate NaCl alleviates osmotic stress in Lycium ruthenicum[J]. Plant Growth Regul, 2022, 96(1): 25-35. doi: 10.1007/s10725-021-00754-0
[49]	Meng KB, Wu YX. Footprints of divergent evolution in two Na⁺/H⁺ type antiporter gene families(NHX and SOS1)in the genus Populus[J]. Tree Physiol, 2018, 38(6): 813-824. doi: 10.1093/treephys/tpx173 URL
[50]	李平, 王晓宇, 徐惠, 等. 蓖麻质膜型Na⁺/H⁺逆向转运蛋白基因(RcSOS1)克隆及表达载体构建[J]. 分子植物育种, 2018, 16(10): 3182-3189.
	Li P, Wang XY, Xu H, et al. Cloning and expression vectors establishment of plasma membrane Na⁺/H⁺ antiporter gene(RcSOS1)in castor(Ricinus communis L.)[J]. Mol Plant Breed, 2018, 16(10): 3182-3189.
[51]	Li Q, Tang Z, Hu YB, et al. Functional analyses of a putative plasma membrane Na⁺/H⁺ antiporter gene isolated from salt tolerant Helianthus tuberosus[J]. Mol Biol Rep, 2014, 41(8): 5097-5108. doi: 10.1007/s11033-014-3375-3 URL
[52]	Guo JR, Dong XX, Han GL, et al. Salt-enhanced reproductive development of Suaeda salsa L. coincided with ion transporter gene upregulation in flowers and increased pollen K⁺ content[J]. Front Plant Sci, 2019, 10: 333. doi: 10.3389/fpls.2019.00333 URL
[53]	Wang WY, Liu YQ, Duan HR, et al. SsHKT1;1 is coordinated with SsSOS1 and SsNHX1 to regulate Na⁺ homeostasis in Suaeda salsa under saline conditions[J]. Plant Soil, 2020, 449(1/2): 117-131. doi: 10.1007/s11104-020-04463-x
[54]	Wang HY, Tang XL, Shao CY, et al. Molecular cloning and bioinformatics analysis of a new plasma membrane Na⁺/H⁺ antiporter gene from the halophyte Kosteletzkya virginica[J]. Sci World J, 2014, 2014: 141675.
[55]	Jarvis DE, Ryu CH, Beilstein MA, et al. Distinct roles for SOS1 in the convergent evolution of salt tolerance in Eutrema salsugineum and Schrenkiella parvula[J]. Mol Biol Evol, 2014, 31(8): 2094-2107. doi: 10.1093/molbev/msu152 pmid: 24803640
[56]	马苏勇, 祝建波, 王爱英, 等. 大叶补血草质膜Na⁺/H⁺逆向转运蛋白基因SOS1的克隆及对番茄的遗传转化[J]. 生物技术通报, 2010(9): 111-115.
	Ma SY, Zhu JB, Wang AY, et al. Cloning and genetic transformation of tomato with SOS1 gene from Limonium gmelinii[J]. Biotechnol Bull, 2010(9): 111-115.
[57]	Abd El-Moneim D, ELsarag EIS, Aloufi S, et al. Quinoa(Chenopodium quinoa Willd.): genetic diversity according to ISSR and SCoT markers, relative gene expression, and morpho-physiological variation under salinity stress[J]. Plants, 2021, 10(12): 2802. doi: 10.3390/plants10122802 URL
[58]	Ma Q, Li YX, Yuan HJ, et al. ZxSOS1 is essential for long-distance transport and spatial distribution of Na⁺ and K⁺ in the xerophyte Zygophyllum xanthoxylum[J]. Plant Soil, 2014, 374(1): 661-676. doi: 10.1007/s11104-013-1891-x URL
[59]	Wang Z, Hong YC, Li YM, et al. Natural variations in SlSOS1 contribute to the loss of salt tolerance during tomato domestication[J]. Plant Biotechnol J, 2021, 19(1): 20-22. doi: 10.1111/pbi.v19.1 URL
[60]	Lu QH, Wang YQ, Xu JP, et al. Effect of ABA on physiological characteristics and expression of salt tolerance-related genes in Tartary buckwheat[J]. Acta Physiol Plant, 2021, 43(5): 1-11. doi: 10.1007/s11738-020-03172-3
[61]	Yadav NS, Shukla PS, Jha A, et al. The SbSOS1 gene from the extreme halophyte Salicornia brachiata enhances Na⁺ loading in xylem and confers salt tolerance in transgenic tobacco[J]. BMC Plant Biol, 2012, 12: 188. doi: 10.1186/1471-2229-12-188
[62]	Yang Y, Xu LF, Li WX, et al. A Na⁺/H⁺ antiporter-encoding salt overly sensitive 1 gene, LpSOS1, involved in positively regulating the salt tolerance in Lilium pumilum[J]. Gene, 2023, 874: 147485. doi: 10.1016/j.gene.2023.147485 URL
[63]	Kaewjiw N, Laksana C, Chanprame S. Cloning and identification of salt overly sensitive(SOS1)gene of sugarcane[J]. Int J Agric Biol, 2018, 20(7): 1569-1574.
[64]	Han QQ, Wang YP, Li J, et al. The mechanistic basis of sodium exclusion in Puccinellia tenuiflora under conditions of salinity and potassium deprivation[J]. Plant J, 2022, 112(2): 322-338. doi: 10.1111/tpj.v112.2 URL
[65]	Núñez-Ramírez R, Sánchez-Barrena MJ, Villalta I, et al. Structural insights on the plant salt-overly-sensitive 1(SOS1)Na⁺/H⁺ antiporter[J]. J Mol Biol, 2012, 424(5): 283-294. doi: 10.1016/j.jmb.2012.09.015 pmid: 23022605
[66]	Quintero FJ, Martinez-Atienza J, Villalta I, et al. Activation of the plasma membrane Na/H antiporter salt-overly-sensitive 1(SOS1)by phosphorylation of an auto-inhibitory C-terminal domain[J]. Proc Natl Acad Sci USA, 2011, 108(6): 2611-2616. doi: 10.1073/pnas.1018921108 URL
[67]	Goswami P, Paulino C, Hizlan D, et al. Structure of the archaeal Na⁺/H⁺ antiporter NhaP1 and functional role of transmembrane helix 1[J]. EMBO J, 2011, 30(2): 439-449. doi: 10.1038/emboj.2010.321 URL
[68]	Zheng M, Li JP, Zeng CW, et al. Subgenome-biased expression and functional diversification of a Na⁺/H⁺ antiporter homoeologs in salt tolerance of polyploid wheat[J]. Front Plant Sci, 2022, 13: 1072009. doi: 10.3389/fpls.2022.1072009 URL
[69]	Ma X, Li QH, Yu YN, et al. The CBL-CIPK pathway in plant response to stress signals[J]. Int J Mol Sci, 2020, 21(16): 5668. doi: 10.3390/ijms21165668 URL
[70]	Tang RJ, Wang C, Li KL, et al. The CBL-CIPK calcium signaling network: unified paradigm from 20 years of discoveries[J]. Trends Plant Sci, 2020, 25(6): 604-617. doi: 10.1016/j.tplants.2020.01.009 URL
[71]	Wu GQ, Wang JL, Li SJ. 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 URL
[72]	Hao SH, Wang YR, Yan YX, et al. A review on plant responses to salt stress and their mechanisms of salt resistance[J]. Horticulturae, 2021, 7(6): 132. doi: 10.3390/horticulturae7060132 URL
[73]	Park HJ, Qiang Z, Kim WY, et al. Diurnal and circadian regulation of salt tolerance in Arabidopsis[J]. J Plant Biol, 2016, 59(6): 569-578. doi: 10.1007/s12374-016-0317-8 URL
[74]	Chen MX, She ZY, Aslam M, et al. Genomic insights of the WRKY genes in kenaf(Hibiscus cannabinus L.) reveal that HcWRKY44 improves the plant's tolerance to the salinity stress[J]. Front Plant Sci, 2022, 13: 984233. doi: 10.3389/fpls.2022.984233 URL
[75]	Wu X, Xu JN, Meng XN, et al. Linker histone variant HIS1-3 and WRKY1 oppositely regulate salt stress tolerance in Arabidopsis[J]. Plant Physiol, 2022, 189(3): 1833-1847. doi: 10.1093/plphys/kiac174 URL
[76]	Zhang HF, Guo JB, Chen XQ, et al. Pepper bHLH transcription factor CabHLH035 contributes to salt tolerance by modulating ion homeostasis and proline biosynthesis[J]. Hortic Res, 2022, 9: uhac203. doi: 10.1093/hr/uhac203 URL
[77]	Fu HQ, Yu X, Jiang YY, et al. SALT OVERLY SENSITIVE 1 is inhibited by clade D protein phosphatase 2C D6 and D7 in Arabidopsis thaliana[J]. Plant Cell, 2023, 35(1): 279-297. doi: 10.1093/plcell/koac283 URL
[78]	Duscha K, Martins Rodrigues C, Müller M, et al. 14-3-3 proteins and other candidates form protein-protein interactions with the cytosolic C-terminal end of SOS1 affecting its transport activity[J]. Int J Mol Sci, 2020, 21(9): 3334. doi: 10.3390/ijms21093334 URL
[79]	Kim WY, Ali Z, Park HJ, et al. Release of SOS2 kinase from sequestration with GIGANTEA determines salt tolerance in Arabidopsis[J]. Nat Commun, 2013, 4: 1352. doi: 10.1038/ncomms2357
[80]	Li JF, Zhou HP, Zhang Y, et al. The GSK3-like kinase BIN2 is a molecular switch between the salt stress response and growth recovery in Arabidopsis thaliana[J]. Dev Cell, 2020, 55(3): 367-380.e6. doi: 10.1016/j.devcel.2020.08.005 URL
[81]	Amirbakhtiar N, Ismaili A, Ghaffari MR, et al. Transcriptome analysis of bread wheat leaves in response to salt stress[J]. PLoS One, 2021, 16(7): e0254189. doi: 10.1371/journal.pone.0254189 URL
[82]	Zhou M, Wang W. SOS1 safeguards plant circadian rhythm against daily salt fluctuations[J]. Proc Natl Acad Sci USA, 2022, 119(36): e2212950119. doi: 10.1073/pnas.2212950119 URL
[83]	Pei SY, Liu YT, Li WK, et al. OSCA1 is an osmotic specific sensor: a method to distinguish Ca²⁺-mediated osmotic and ionic perception[J]. New Phytol, 2022, 235(4): 1665-1678. doi: 10.1111/nph.v235.4 URL
[84]	Yuan F, Yang HM, Xue Y, et al. OSCA1 mediates osmotic-stress-evoked Ca²⁺ increases vital for osmosensing in Arabidopsis[J]. Nature, 2014, 514(7522): 367-371. doi: 10.1038/nature13593
[85]	Jiang ZH, Zhou XP, Tao M, et al. Plant cell-surface GIPC sphingolipids sense salt to trigger Ca²⁺ influx[J]. Nature, 2019, 572(7769): 341-346. doi: 10.1038/s41586-019-1449-z
[86]	Yin XC, Xia YQ, Xie Q, et al. The protein kinase complex CBL10-CIPK8-SOS1 functions in Arabidopsis to regulate salt tolerance[J]. J Exp Bot, 2020, 71(6): 1801-1814. doi: 10.1093/jxb/erz549 URL
[87]	Zhu JK. Abiotic stress signaling and responses in plants[J]. Cell, 2016, 167(2): 313-324. doi: 10.1016/j.cell.2016.08.029 URL
[88]	Guo Y, Liu Y, Zhang Y, et al. Effects of exogenous calcium on adaptive growth, photosynthesis, ion homeostasis and phenolics of Gleditsia sinensis Lam. plants under salt stress[J]. Agriculture, 2021, 11(10): 978. doi: 10.3390/agriculture11100978 URL
[89]	Fraile-Escanciano A, Kamisugi Y, Cuming AC, et al. The SOS1 transporter of Physcomitrella patens mediates sodium efflux in planta[J]. New Phytol, 2010, 188(3): 750-761. doi: 10.1111/j.1469-8137.2010.03405.x pmid: 20696009
[90]	Gupta A, Shaw BP, Sahu BB. Post-translational regulation of the membrane transporters contributing to salt tolerance in plants[J]. Funct Plant Biol, 2021, 48(12): 1199-1212. doi: 10.1071/FP21153 pmid: 34665998
[91]	Quintero FJ, Ohta M, Shi HZ, et al. Reconstitution in yeast of the Arabidopsis SOS signaling pathway for Na⁺ homeostasis[J]. Proc Natl Acad Sci USA, 2002, 99(13): 9061-9066. doi: 10.1073/pnas.132092099 URL
[92]	Zhou Y, Lai ZS, Yin XC, et al. Hyperactive mutant of a wheat plasma membrane Na⁺/H⁺ antiporter improves the growth and salt tolerance of transgenic tobacco[J]. Plant Sci, 2016, 253: 176-186. doi: 10.1016/j.plantsci.2016.09.016 URL
[93]	Xie Q, Yang Y, Wang Y, et al. The calcium sensor CBL10 negatively regulates plasma membrane H⁺-ATPase activity and alkaline stress response in Arabidopsis[J]. Environ Exp Bot, 2022, 194: 104752. doi: 10.1016/j.envexpbot.2021.104752 URL
[94]	Park HJ, Kim WY, Yun DJ. A new insight of salt stress signaling in plant[J]. Mol Cells, 2016, 39(6): 447-459. doi: 10.14348/molcells.2016.0083 pmid: 27239814
[95]	Du WM, Lin HX, Chen S, et al. Phosphorylation of SOS3-like calcium-binding proteins by their interacting SOS2-like protein kinases is a common regulatory mechanism in Arabidopsis[J]. Plant Physiol, 2011, 156(4): 2235-2243. doi: 10.1104/pp.111.173377 URL
[96]	谢玲玲, 王金龙, 伍国强. 植物CBL-CIPK信号系统响应非生物胁迫的调控机制[J]. 植物学报, 2021, 56(5): 614-626. doi: 10.11983/CBB21024
	Xie LL, Wang JL, Wu GQ. Regulatory mechanisms of the plant CBL-CIPK signaling system in response to abiotic stress[J]. Chin Bull Bot, 2021, 56(5): 614-626.
[97]	Wang Q, Guan C, Wang P, et al. The effect of AtHKT1;1 or AtSOS1 mutation on the expressions of Na⁺ or K⁺ transporter genes and ion homeostasis in Arabidopsis thaliana under salt stress[J]. Int J Mol Sci, 2019, 20(5): 1085. doi: 10.3390/ijms20051085 URL
[98]	Venkataraman G, Shabala S, Véry AA, et al. To exclude or to accumulate? Revealing the role of the sodium HKT1;5 transporter in plant adaptive responses to varying soil salinity[J]. Plant Physiol Biochem, 2021, 169: 333-342. doi: 10.1016/j.plaphy.2021.11.030 URL
[99]	Riedelsberger J, Miller JK, Valdebenito-Maturana B, et al. Plant HKT channels: an updated view on structure, function and gene regulation[J]. Int J Mol Sci, 2021, 22(4): 1892. doi: 10.3390/ijms22041892 URL
[100]	Gu WT, Zhou LB, Liu RY, et al. Synergistic responses of NHX, AKT1, and SOS1 in the control of Na⁺ homeostasis in sweet sorghum mutants induced by ¹²C⁶⁺-ion irradiation[J]. Nucl Sci Tech, 2018, 29(1): 10.
[101]	Sun QA, Yamada T, Han YL, et al. Differential responses of NHX1 and SOS1 gene expressions to salinity in two Miscanthus sinensis anderss. accessions with different salt tolerance[J]. Phyton, 2021, 90(3): 827-836. doi: 10.32604/phyton.2021.013805 URL
[102]	Olías R, Eljakaoui Z, Li J, et al. The plasma membrane Na⁺/H⁺ antiporter SOS1 is essential for salt tolerance in tomato and affects the partitioning of Na⁺ between plant organs[J]. Plant Cell Environ, 2009, 32(7): 904-916. doi: 10.1111/pce.2009.32.issue-7 URL
[103]	Ishikawa T, Shabala L, Zhou MX, et al. Comparative analysis of root Na⁺ relation under salinity between Oryza sativa and Oryza coarctata[J]. Plants, 2022, 11(5): 656. doi: 10.3390/plants11050656 URL
[104]	Fan YF, Yin XC, Xie Q, et al. Co-expression of SpSOS1 and SpAHA1 in transgenic Arabidopsis plants improves salinity tolerance[J]. BMC Plant Biol, 2019, 19(1): 74. doi: 10.1186/s12870-019-1680-7
[105]	Shabala S, Munns R. Salinity stress: physiological constraints and adaptive mechanisms[M]// Plant Stress Physiology. UK: CABI, 2017: 24-63.
[106]	Katiyar-Agarwal S, Zhu JH, Kim K, et al. The plasma membrane Na⁺/H⁺ antiporter SOS1 interacts with RCD1 and functions in oxidative stress tolerance in Arabidopsis[J]. Proc Natl Acad Sci USA, 2006, 103(49): 18816-18821. pmid: 17023541
[107]	Feki K, Tounsi S, Masmoudi K, et al. The durum wheat plasma membrane Na⁺/H⁺ antiporter SOS1 is involved in oxidative stress response[J]. Protoplasma, 2017, 254(4): 1725-1734. doi: 10.1007/s00709-016-1066-8 URL
[108]	Zhou Y, Yin XC, Wan SM, et al. The Sesuvium portulacastrum plasma membrane Na⁺/H⁺ antiporter SpSOS1 complemented the salt sensitivity of transgenic Arabidopsis sos1 mutant plants[J]. Plant Mol Biol Rep, 2018, 36(4): 553-563. doi: 10.1007/s11105-018-1099-6
[109]	Xu XD, Yuan L, Xie QG. The circadian clock ticks in plant stress responses[J]. Stress Biol, 2022, 2(1): 15. doi: 10.1007/s44154-022-00040-7 pmid: 37676516
[110]	Yang YQ, Wu YJ, Ma L, et al. The Ca²⁺ sensor SCaBP3/CBL7 modulates plasma membrane H⁺-ATPase activity and promotes alkali tolerance in Arabidopsis[J]. Plant Cell, 2019, 31(6): 1367-1384. doi: 10.1105/tpc.18.00568 URL
[111]	Oh DH, Lee SY, Bressan RA, et al. Intracellular consequences of SOS1 deficiency during salt stress[J]. J Exp Bot, 2010, 61(4): 1205-1213. doi: 10.1093/jxb/erp391 URL
[112]	Zhou JY, Hao DL, Yang GZ. Regulation of cytosolic pH: the contributions of plant plasma membrane H⁺-ATPases and multiple transporters[J]. Int J Mol Sci, 2021, 22(23): 12998. doi: 10.3390/ijms222312998 URL
[113]	Guo KM, Babourina O, Rengel Z. Na⁺/H⁺ antiporter activity of the SOS1 gene: lifetime imaging analysis and electrophysiological studies on Arabidopsis seedlings[J]. Physiol Plant, 2009, 137(2): 155-165.
[114]	Cha JY, Kim J, Jeong SY, et al. The Na⁺/H⁺ antiporter salt overly sensitive 1 regulates salt compensation of circadian rhythms by stabilizing gigantea in arabidopsis[J]. Proc Natl Acad Sci USA, 2022, 119(33): e2207275119. doi: 10.1073/pnas.2207275119 URL
[115]	Ma YC, Wang L, Wang JY, et al. Isolation and expression analysis of salt overly sensitive gene family in grapevine(Vitis vinifera)in response to salt and PEG stress[J]. PLoS One, 2019, 14(3): e0212666. doi: 10.1371/journal.pone.0212666 URL
[116]	Shahzad B, Yun P, Rasouli F, et al. Root K⁺ homeostasis and signalling as a determinant of salinity stress tolerance in cultivated and wild rice species[J]. Environ Exp Bot, 2022, 201: 104944. doi: 10.1016/j.envexpbot.2022.104944 URL
[117]	Chai HX, Guo JF, Zhong YL, et al. The plasma-membrane polyamine transporter PUT3 is regulated by the Na⁺/H⁺ antiporter SOS1 and protein kinase SOS2[J]. New Phytol, 2020, 226(3): 785-797. doi: 10.1111/nph.v226.3 URL
[118]	Farooq M, Park JR, Jang YH, et al. Rice cultivars under salt stress show differential expression of genes related to the regulation of Na⁺/K⁺ balance[J]. Front Plant Sci, 2021, 12: 680131. doi: 10.3389/fpls.2021.680131 URL
[119]	Yue YS, Zhang MC, Zhang JC, et al. SOS1 gene overexpression increased salt tolerance in transgenic tobacco by maintaining a higher K⁺/Na⁺ ratio[J]. J Plant Physiol, 2012, 169(3): 255-261. doi: 10.1016/j.jplph.2011.10.007 URL
[120]	Akrimi R, Hajlaoui H, Batelli G, et al. Electromagnetic water enhanced metabolism and agro-physiological responses of potato(Solanum tuberosum L)under saline conditions[J]. J Agron Crop Sci, 2021, 207(1): 44-58. doi: 10.1111/jac.v207.1 URL
[121]	Awaji SM, Hanjagi PS, Pushpa BN, et al. Overexpression of plasma membrane Na⁺/H⁺ antiporter OsSOS1 gene improves salt tolerance in transgenic rice plants[J]. Oryza, 2020, 57(4): 277-287. doi: 10.35709/ory.2019.56.2 URL
[122]	Rao YR, Ansari MW, Sahoo RK, et al. Salicylic acid modulates ACS, NHX1, sos1 and HKT1;2 expression to regulate ethylene overproduction and Na⁺ ions toxicity that leads to improved physiological status and enhanced salinity stress tolerance in tomato plants cv. Pusa Ruby[J]. Plant Signal Behav, 2021, 16(11): 1950888. doi: 10.1080/15592324.2021.1950888 URL
[123]	Chang PJ, Wu Z, Song NN, et al. Identification MdeSOS1 in Magnolia denudata and its function in response to salt stress[J]. J Plant Interact, 2020, 15(1): 417-426. doi: 10.1080/17429145.2020.1843721 URL
[124]	Hichri I, Muhovski Y, Clippe A, et al. SlDREB2, a tomato dehydration-responsive element-binding 2 transcription factor, mediates salt stress tolerance in tomato and Arabidopsis[J]. Plant Cell Environ, 2016, 39(1): 62-79. doi: 10.1111/pce.v39.1 URL

物种名 Species name	基因名称 Gene name	基因登录号 GenBank accession No.	氨基酸 Amino acid	亚细胞定位 Sub-cellular localization	跨膜结构域TMD	亲水性平均值 GRAVY	等电点 pI	分子量 Mw/kD	参考文献 Reference
甜菜 Beta vulgaris	BvSOS1	/	1 162	PM	12	0.085	6.34	128.46	Unpublished
小盐芥thellungiella halophila	ThSOS1	EF207775	1 146	PM	11	0.090	6.40	126.35	[36]
小盐芥thellungiella halophila	ThSOS1S	AB562331	473	PM	11	0.722	6.15	51.19	[37]
冰叶日中花 Mesembryanthemum crystallinum	McSOS1	EF207776	1 115	PM	12	0.110	5.97	127.36	[38]
花花柴 Karelinia caspia	KcSOS1	MN481368	1 136	PM	12	0.070	5.97	126.02	[39]
水稻Oryza sativa	OsSOS1-1	AY785147	1 148	PM	12	0.046	6.77	127.92	[11,21 -22]
水稻Oryza sativa	OsSOS1-2	MG602252	1 148	PM	13	0.047	6.77	127.90	[11,21 -22]
小麦 Triticum aestivum	TaSOS1-1	KJ563230	1 142	PM	12	0.110	7.00	126.22	[24]
小麦 Triticum aestivum	TaSOS1-2	AY326952	1 142	PM	13	0.106	7.40	126.18	[24]
大豆 Glycine max	GmSOS1	JQ287499	1 143	PM	12	0.089	6.30	126.43	[32]
文冠果Xanthoceras sorbifolium	XsSOS1	/	915	PM	6	/	6.19	102.50	[33]
萝卜 Raphanus sativus	RsSOS1	MZ484950	1 137	PM	12	0.093	6.71	125.62	[26]
海马齿Sesuvium portulacastrum	SpSOS1	JX674067	1 155	PM	12	0.063	6.08	127.95	[27]
印度南瓜 Cucurbita pepo	CmaSOS1	NW_019272028	1 142	PM	12	0.079	5.92	126.70	[20]
陆地棉Gossypium hirsutum	GhSOS1	KU994886	1 152	PM	12	0.061	6.32	128.18	[40]
拟南芥Arabidopsis thaliana	AtSOS1	AT2G01980	1 146	PM	13	0.098	7.62	127.19	[16]
荞麦Fagopyrum esculentum	FeSOS1	MH064255	1 132	PM	12	0.087	6.36	126.20	[31]
白刺 Nitraria tangutorum	NtSOS1	KC292267	1 163	PM	11	0.107	6.43	127.59	[41]
黄花草木樨Melilotus officinalis	MoSOS1	MG680455	957	PM	8	0.000	6.10	107.21	[42]
黄花红砂Reaumuria trigyna	RtSOS1	KC292265	1 145	PM	11	0.114	6.10	126.79	[43]
木榄Bruguiera gymnorhiza	BgSOS1	HM054521	1 153	PM	12	0.136	6.19	126.91	[44]
盐生草Halogeton glomeratus	HgSOS1	KT759142	1 165	PM	11	0.086	6.21	128.97	/
牡蒿 Artemisia japonica	AjSOS1	KP896475	1 147	PM	12	0.061	6.08	127.04	[45]
菠菜 Spinacia oleracea	SoSOS1	HG799055	1 102	/	12	0.164	6.38	121.97	[28]
葡萄 Vitis vinifera	VvSOS1	NM_001281211	1 141	/	12	0.122	6.32	126.30	[46]
黄瓜 Cucumis sativus	CsSOS1	JQ655747	1 144	PM	11	0.078	6.30	127.27	[47]
黑果枸杞Lycium ruthenicum	LrSOS1	MH454108	1 160	/	12	0.062	5.89	128.61	[48]
欧洲油菜 Brassica napus	BnSOS1	EU487184	1 142	PM	11	0.109	6.45	126.01	[26]
胡杨 Populus euphratica	PeSOS1	KX132789	1 145	PM	12	0.141	6.43	126.84	[49]
篦麻 Ricinus communis	RcSOS1	KX943304	1 143	PM	12	0.069	6.62	126.49	[50]
菊芋 Helianthus tuberosus	HtSOS1	KC410809	1 130	PM	12	0.091	6.00	125.00	[51]
盐地碱蓬 Suaeda salsa	SsSOS1	KF914414.1	1 169	PM	12	0.023	6.52	129.20	[52-53]
绿豆 Vigna radiata	VrSOS1	KC855193	1 046	/	11	0.119	6.20	116.10	/
节节麦 Aegilops tauschii	AtASOS1	FN356231	1 142	/	12	0.10	6.85	126.18	/
短柄草Brachypodium sylvaticum	BsSOS1	KF671967	1 140	/	12	0.078	7.63	126.77	/
海滨锦葵Kosteletzkya virginica	KvSOS1	KJ577576	1 147	PM	12	0.073	6.18	127.56	[54]
山萮菜Eutrema salsugineum	EsSOS1	KF671959	1 127	/	12	0.064	6.47	124.51	[55]
一粒小麦Triticum monococcum	TmSOS1	FN356229	1 142	/	12	0.108	6.87	126.33	/
大叶补血草Limonium gmelinii	LgSOS1	EU780458	1 151	PM	13	0.163	6.15	126.52	[56]
藜麦Chenopodium quinoa	CqSOS1	EU024570	1 158	PM	11	0.094	6.27	127.90	[57]
霸王Zygophyllum xanthoxylon	ZxSOS1	GU177864	1 153	PM	12	0.083	6.20	127.88	[58]
番茄Solanum lycopersicum	SlSOS1-1	AB675690	1 151	PM	12	0.069	5.85	127.54	[59]
番茄Solanum lycopersicum	SlSOS1-2	NM_001247769	1 151	PM	13	0.083	5.89	127.50	[59]
苦荞麦Fagopyrum tataricum	FtSOS1	KY659584	1 131	PM	12	0.090	6.13	125.88	[60]
盐角草Salicornia brachiata	SbSOS1	EU879059	1 159	/	11	0.037	5.92	128.55	[61]
山丹 Lilium pumilum	LpSOS1	OM650676	1 161	PM	12	0.107	6.59	129.14	[62]
甘蔗Saccharum officinarum	ScSOS1	KT003285	423	Nucleus	/	-0.396	9.12	47.60	[63]
小花碱茅Puccinellia tenuiflora	PtSOS1-1	EF440291	1 137	PM	11	0.107	6.48	125.50	[64]
小花碱茅Puccinellia tenuiflora	PtSOS1-2	GQ452778	1 143	PM	13	0.103	6.82	126.16	[64]
海岛棉Gossypium barbadense	GbSOS1	/	1 152	PM	/	/	6.42	128.04	[35]

物种名 Species name	基因名称 Gene name	基因登录号 GenBank accession No.	氨基酸 Amino acid	亚细胞定位 Sub-cellular localization	跨膜结构域TMD	亲水性平均值 GRAVY	等电点 pI	分子量 Mw/kD	参考文献 Reference
甜菜 Beta vulgaris	BvSOS1	/	1 162	PM	12	0.085	6.34	128.46	Unpublished
小盐芥thellungiella halophila	ThSOS1	EF207775	1 146	PM	11	0.090	6.40	126.35	[36]
小盐芥thellungiella halophila	ThSOS1S	AB562331	473	PM	11	0.722	6.15	51.19	[37]
冰叶日中花 Mesembryanthemum crystallinum	McSOS1	EF207776	1 115	PM	12	0.110	5.97	127.36	[38]
花花柴 Karelinia caspia	KcSOS1	MN481368	1 136	PM	12	0.070	5.97	126.02	[39]
水稻Oryza sativa	OsSOS1-1	AY785147	1 148	PM	12	0.046	6.77	127.92	[11,21 -22]
水稻Oryza sativa	OsSOS1-2	MG602252	1 148	PM	13	0.047	6.77	127.90	[11,21 -22]
小麦 Triticum aestivum	TaSOS1-1	KJ563230	1 142	PM	12	0.110	7.00	126.22	[24]
小麦 Triticum aestivum	TaSOS1-2	AY326952	1 142	PM	13	0.106	7.40	126.18	[24]
大豆 Glycine max	GmSOS1	JQ287499	1 143	PM	12	0.089	6.30	126.43	[32]
文冠果Xanthoceras sorbifolium	XsSOS1	/	915	PM	6	/	6.19	102.50	[33]
萝卜 Raphanus sativus	RsSOS1	MZ484950	1 137	PM	12	0.093	6.71	125.62	[26]
海马齿Sesuvium portulacastrum	SpSOS1	JX674067	1 155	PM	12	0.063	6.08	127.95	[27]
印度南瓜 Cucurbita pepo	CmaSOS1	NW_019272028	1 142	PM	12	0.079	5.92	126.70	[20]
陆地棉Gossypium hirsutum	GhSOS1	KU994886	1 152	PM	12	0.061	6.32	128.18	[40]
拟南芥Arabidopsis thaliana	AtSOS1	AT2G01980	1 146	PM	13	0.098	7.62	127.19	[16]
荞麦Fagopyrum esculentum	FeSOS1	MH064255	1 132	PM	12	0.087	6.36	126.20	[31]
白刺 Nitraria tangutorum	NtSOS1	KC292267	1 163	PM	11	0.107	6.43	127.59	[41]
黄花草木樨Melilotus officinalis	MoSOS1	MG680455	957	PM	8	0.000	6.10	107.21	[42]
黄花红砂Reaumuria trigyna	RtSOS1	KC292265	1 145	PM	11	0.114	6.10	126.79	[43]
木榄Bruguiera gymnorhiza	BgSOS1	HM054521	1 153	PM	12	0.136	6.19	126.91	[44]
盐生草Halogeton glomeratus	HgSOS1	KT759142	1 165	PM	11	0.086	6.21	128.97	/
牡蒿 Artemisia japonica	AjSOS1	KP896475	1 147	PM	12	0.061	6.08	127.04	[45]
菠菜 Spinacia oleracea	SoSOS1	HG799055	1 102	/	12	0.164	6.38	121.97	[28]
葡萄 Vitis vinifera	VvSOS1	NM_001281211	1 141	/	12	0.122	6.32	126.30	[46]
黄瓜 Cucumis sativus	CsSOS1	JQ655747	1 144	PM	11	0.078	6.30	127.27	[47]
黑果枸杞Lycium ruthenicum	LrSOS1	MH454108	1 160	/	12	0.062	5.89	128.61	[48]
欧洲油菜 Brassica napus	BnSOS1	EU487184	1 142	PM	11	0.109	6.45	126.01	[26]
胡杨 Populus euphratica	PeSOS1	KX132789	1 145	PM	12	0.141	6.43	126.84	[49]
篦麻 Ricinus communis	RcSOS1	KX943304	1 143	PM	12	0.069	6.62	126.49	[50]
菊芋 Helianthus tuberosus	HtSOS1	KC410809	1 130	PM	12	0.091	6.00	125.00	[51]
盐地碱蓬 Suaeda salsa	SsSOS1	KF914414.1	1 169	PM	12	0.023	6.52	129.20	[52-53]
绿豆 Vigna radiata	VrSOS1	KC855193	1 046	/	11	0.119	6.20	116.10	/
节节麦 Aegilops tauschii	AtASOS1	FN356231	1 142	/	12	0.10	6.85	126.18	/
短柄草Brachypodium sylvaticum	BsSOS1	KF671967	1 140	/	12	0.078	7.63	126.77	/
海滨锦葵Kosteletzkya virginica	KvSOS1	KJ577576	1 147	PM	12	0.073	6.18	127.56	[54]
山萮菜Eutrema salsugineum	EsSOS1	KF671959	1 127	/	12	0.064	6.47	124.51	[55]
一粒小麦Triticum monococcum	TmSOS1	FN356229	1 142	/	12	0.108	6.87	126.33	/
大叶补血草Limonium gmelinii	LgSOS1	EU780458	1 151	PM	13	0.163	6.15	126.52	[56]
藜麦Chenopodium quinoa	CqSOS1	EU024570	1 158	PM	11	0.094	6.27	127.90	[57]
霸王Zygophyllum xanthoxylon	ZxSOS1	GU177864	1 153	PM	12	0.083	6.20	127.88	[58]
番茄Solanum lycopersicum	SlSOS1-1	AB675690	1 151	PM	12	0.069	5.85	127.54	[59]
番茄Solanum lycopersicum	SlSOS1-2	NM_001247769	1 151	PM	13	0.083	5.89	127.50	[59]
苦荞麦Fagopyrum tataricum	FtSOS1	KY659584	1 131	PM	12	0.090	6.13	125.88	[60]
盐角草Salicornia brachiata	SbSOS1	EU879059	1 159	/	11	0.037	5.92	128.55	[61]
山丹 Lilium pumilum	LpSOS1	OM650676	1 161	PM	12	0.107	6.59	129.14	[62]
甘蔗Saccharum officinarum	ScSOS1	KT003285	423	Nucleus	/	-0.396	9.12	47.60	[63]
小花碱茅Puccinellia tenuiflora	PtSOS1-1	EF440291	1 137	PM	11	0.107	6.48	125.50	[64]
小花碱茅Puccinellia tenuiflora	PtSOS1-2	GQ452778	1 143	PM	13	0.103	6.82	126.16	[64]
海岛棉Gossypium barbadense	GbSOS1	/	1 152	PM	/	/	6.42	128.04	[35]