苜蓿抗旱性分子研究进展

doi:10.13560/j.cnki.biotech.bull.1985.2021-0057

摘要/Abstract

摘要：

干旱胁迫是严重限制全球农业生产的环境因素,造成了牧草大量减产。综述了4种重要苜蓿在干旱胁迫下的相关基因以及抗旱育种方面的主要进展,并对苜蓿耐旱性研究的前景及存在问题进行了讨论。提出可充分利用生物技术,挖掘抗旱相关基因资源,在豆科模式植物蒺藜苜蓿中验证基因功能,阐明苜蓿抗旱应答网络机制,进而从黄花苜蓿中克隆抗旱相关基因,最终通过分子育种手段培养高抗旱性紫花苜蓿新品种。培育耐旱紫花苜蓿新品种是改善其在干旱胁迫下生长、提高其产量的有效途径,通过生物技术获得耐旱紫花苜蓿种质可用于遗传改良,为我国粮改饲政策的实施提供饲草新材料,促进草牧业健康可持续发展。

关键词: 苜蓿属, 抗旱, 分子机制, 分子育种

Abstract:

Drought stress is one of the most serious environmental constraints for global agricultural production,and it causes severe losses in pasture production. In this paper,drought induction related genes and the main progresses on the breeding of 4 important Medicago plants with drought resistance were reviewed,and the prospects and issues in this field were also discussed. It is suggested that biological technology can be utilized to explore drought-resistance related gene resources,then to preliminarily verify the function of target genes in the model legume plant Medicago truncatula. Further the mechanism of drought resistance response network in M. truncatula was clarified,and then the drought-resistance related genes from Medicago falcata was cloned. Finally,cultivating new alfalfa varieties with high drought-resistance was proposed by using molecular breeding approaches. Cultivating new alfalfa varieties with drought-tolerant is an effective way to improve Medicago growth and yield under drought stress. It is of great significance to obtain drought-tolerant alfalfa germplasm through biotechnology approaches for genetic improvement,and to provide new cultivars during the implementation of the Grain to Feed Policy of China,and ultimately to promote the healthy and sustainable development of grass and animal husbandry.

Key words: Medicago, drought resistance, molecular mechanism, molecular breeding

李倩, 江文波, 王玉祥, 张博, 庞永珍. 苜蓿抗旱性分子研究进展[J]. 生物技术通报, 2021, 37(8): 243-252.

LI Qian, JIANG Wen-bo, WANG Yu-xiang, ZHANG Bo, PANG Yong-zhen. Research Progresses on the Drought Resistance of Medicago at Molecular Level[J]. Biotechnology Bulletin, 2021, 37(8): 243-252.

图/表 2

参考文献 67

[1]	Cook DR. Medicago truncatula-a model in the making[J]. Current Opinion in Plant Biology, 1999, 2(4):301-304. pmid: 10459004
[2]	Liu ZP, Chen TL, Ma LC, et al. Global transcriptome sequencing using the Illumina platform and the development of EST-SSR markers in autotetraploid alfalfa[J]. PLoS One, 2013, 8(12):e83549. doi: 10.1371/journal.pone.0083549 URL
[3]	O’Rourke JA, Fu F, Bucciarelli B, et al. The Medicago sativa gene index 1.2:a web-accessible gene expression atlas for investigating expression differences between Medicago sativa subspecies[J]. BMC Genomics, 2015, 16(1):502. doi: 10.1186/s12864-015-1718-7 URL
[4]	杨青川. 苜蓿种植区划及品种指南[M]. 北京: 中国农业大学出版社, 2012.
	Yang QC. Guide for alfalfa planting zone and cultivar[M]. Beijing: China Agricultural University Press, 2012.
[5]	王俊杰, 云锦凤, 吕世杰. 黄花苜蓿种质的优良特性与利用价值[J]. 内蒙古农业大学学报:自然科学版, 2008, 29(1):215-219.
	Wang JJ, Yun JF, Lv SJ. Characteristics and utilizing values on the germplasm of Medicago falcata[J]. Journal of Inner Mongolia Agricultural University:Natural Science Edition, 2008, 29(1):215-219.
[6]	杜凯青. 草原3号杂花苜蓿表型多样性研究[D]. 呼和浩特:内蒙古农业大学, 2020.
	Du KQ. Phenotypic diversity of Medicago varia Martin.cv. Caoyuan No3[D]. Hohhot:Inner Mongolia Agricultural University, 2020.
[7]	Maureira-Butler IJ, Pfeil BE, Muangprom A, et al. The reticulate history of Medicago(Fabaceae)[J]. Systematic Biology, 2008, 57(3):466-482. doi: 10.1080/10635150802172168 pmid: 18570039
[8]	Maghsoodi M, Razmjoo J. Identify physiological markers for drought tolerance in alfalfa[J]. Agron J, 2015, 107(1):149-157. doi: 10.2134/agronj14.0255 URL
[9]	Huang Z, Liu Y, Cui Z, et al. Soil water storage deficit of alfalfa(Medicago sativa)grasslands along ages in arid area(China)[J]. Field Crops Research, 2018, 221:1-6. doi: 10.1016/j.fcr.2018.02.013 URL
[10]	Sivakumar MVK, Das HP, Brunini O. Impacts of present and future climate variability and change on agriculture and forestry in the arid and semi-arid tropics[J]. Climatic Change, 2005, 70(70):31-72. doi: 10.1007/s10584-005-5937-9 URL
[11]	Chen HT, Zeng Y, Yang YZ, et al. Allele-aware chromosome-level genome assembly and efficient transgene-free genome editing for the autotetraploid cultivated alfalfa[J]. Nature Communications, 2020, 11(1):2494. doi: 10.1038/s41467-020-16338-x URL
[12]	Zhao Y, Wei X, Ji X, et al. Endogenous NO-mediated transcripts involved in photosynjournal and carbohydrate metabolism in alfalfa(Medicago sativa L.)seedlings under drought stress[J]. Plant Physiology and Biochemistry, 2019, 141:456-465. doi: 10.1016/j.plaphy.2019.06.023 URL
[13]	Luo D, Zhou Q, Wu YQ, et al. Full-length transcript sequencing and comparative transcriptomic analysis to evaluate the contribution of osmotic and ionic stress components towards salinity tolerance in the roots of cultivated alfalfa(Medicago sativa L.)[J]. BMC Plant Biology, 2019, 19(1):32. doi: 10.1186/s12870-019-1630-4 URL
[14]	Shen C, Du H, Chen Z, et al. The Chromosome-Level genome sequence of the autotetraploid alfalfa and resequencing of core germplasms provide genomic resources for alfalfa research[J]. Mol Plant, 2020, 13(9):1250-1261. doi: S1674-2052(20)30216-1 pmid: 32673760
[15]	Zhang T, Yu LX, Zheng P, et al. Identification of loci associated with drought resistance traits in heterozygous autotetraploid alfalfa(Medicago sativa L.)using genome-wide association studies with genotyping by sequencing[J]. PLoS One, 2015, 10(9):e0138931. doi: 10.1371/journal.pone.0138931 URL
[16]	Ray IM, Han Y, Lei E, et al. Identification of QTL for alfalfa forage biomass productivity during drought[J]. Crop Science, 2015. DOI: 10.2135/cropsci2014.12.0840 doi: 10.2135/cropsci2014.12.0840
[17]	Wang Z, Wang X, Zhang H, et al. A genome-wide association study approach to the identification of candidate genes underlying agronomic traits in alfalfa(Medicago sativa L.)[J]. Plant Biotechnology Journal, 2020, 18(3):611-613. doi: 10.1111/pbi.13251 pmid: 31487419
[18]	Miao Z, Xu W, Li D, et al. De novo transcriptome analysis of Medicago falcata reveals novel insights about the mechanisms underlying abiotic stress-responsive pathway[J]. BMC Genomics, 2015, 16(1):818-836. doi: 10.1186/s12864-015-2019-x URL
[19]	Duan M, Zhang R, Zhu F, et al. A lipid-anchored NAC transcription factor is translocated into the nucleus and activates glyoxalase I expression during drought stress[J]. The Plant Cell, 2017, 29(7):1748-1772. doi: 10.1105/tpc.17.00044 pmid: 28684428
[20]	苗震龑. 黄花苜蓿非生物胁迫数据挖掘与系统分析[D]. 北京:中国农业大学, 2014.
	Miao ZY. Data mining and systemic analysis of Medicago falcata under abiotic stress[D]. Beijing:China Agricultural University, 2014.
[21]	Zhang JY, Cruz De Carvalho MH, Torres Jerez I, et al. Global reprogramming of transcription and metabolism in Medicago truncatula during progressive drought and after rewatering[J]. Plant Cell & Environment, 2014, 37(11):2553-2576.
[22]	刘赢. 蒺藜苜蓿R2R3-MYB转录因子的发掘及其对非生物胁迫反应研究[D]. 哈尔滨:哈尔滨师范大学, 2017.
	Liu Y. Identification and characterization of R2R3-MYB transcription factor and their response to the abiotic stress in Medicago truncatula[D]. Harbin:Harbin Normal University, 2017.
[23]	Kang Y, Sakiroglu M, et al. Genome-wide association of drought-related and biomass traits with HapMap SNPs in Medicago truncatula[J]. Plant Cell Environ, 2015, 38(10):1997-2011. doi: 10.1111/pce.12520 URL
[24]	Wang T, Chen L, Zhao M, et al. Identification of drought-responsive microRNAs in Medicago truncatula by genome-wide high-throughput sequencing[J]. BMC Genomics, 2011, 12(1):367-378. doi: 10.1186/1471-2164-12-367 URL
[25]	杨青川, 孙彦, 康俊梅. 紫花苜蓿耐盐相关基因克隆研究进展[J]. 草地学报, 2005, 13(3):253-256.
	Yang QC, Sun Y, Kang JM. Research on the advancement of salt tolerant gene in alfalfa[J]. Acta Agrestia Sinica, 2005, 13(3):253-256.
[26]	申玉华, 徐振军, 唐立红, 等. 紫花苜蓿NAC类转录因子基因MsNAC2的克隆及其功能分析[J]. 中国农业科学, 2015, 48(15):2925-2938.
	Shen YH, Xu ZJ, Tang LH, et al. Cloning and function analysis of the MsNAC2 gene with NAC transcription factor from alfalfa[J]. Scientia Agricultura Sinica, 2015, 48(15):2925-2938.
[27]	常娜. 黄花苜蓿MfNAC35基因的功能分析[D]. 呼和浩特:内蒙古大学, 2018.
	Chang N. Funtional analysis of MFNAC35 gene in Medicago falcata[D]. Hohhot:Inner Mongolia University, 2018.
[28]	Song Y, Lv J, Qiu N, et al. The constitutive expression of alfalfa MsMYB2L enhances salinity and drought tolerance of Arabidopsis thaliana[J]. Plant Physiol Biochem, 2019, 141:300-305. doi: 10.1016/j.plaphy.2019.06.007 URL
[29]	张春霄. AP2/EREBP家族MfERF049基因对提高豆科植物生物胁迫和非生物胁迫抗性的功能分析[D]. 呼和浩特:内蒙古大学, 2019.
	Zhang CX. The functional analysis of MfERF049 gene in AP2/EREBP Family with improve biological and abiotic stress resistance about leguminous plant[D]. Hohhot:Inner Mongolia University, 2019.
[30]	付佳宾. 野生黄花苜蓿ERF家族成员MfERF026和MfERF086基因的克隆及初步功能研究[D]. 呼和浩特:内蒙古大学, 2019.
	Fu JB. Identification of MfERF026 and MfERF086 genes from ERF family of wild Medicago falcata[D]. Hohhot:Inner Mongolia University, 2019.
[31]	Chen T, Yang Q, Zhang X, et al. An alfalfa(Medicago sativa L.)ethylene response factor gene, MsERF11, enhances salt tolerance in transgenic Arabidopsis[J]. Plant Cell Reports, 2012, 31(9):1737-1746. doi: 10.1007/s00299-012-1287-z URL
[32]	Li Y, Sun Y, Yang Q, et al. Isolation and characterization of a gene from Medicago sativa L. encoding a bZIP transcription factor[J]. Molecular Biology Reports, 2013, 40(2):1227-1239. doi: 10.1007/s11033-012-2165-z pmid: 23096087
[33]	王菊萍, 王珍, 张铁军, 等. 蒺藜苜蓿MtbHLH148转录因子的克隆与转化及其功能分析[J]. 西北植物学报, 2019, 39(6):963-973.
	Wang JP, Wang Z, Zhang TJ, et al. Cloning and analysis of a basic helix-loop-helix(bHLH)transcription factor MtbHLH148 from Medicago truncatula L.[J]. Acta Botanica Boreali-Occidentalia Sinica, 2019, 39(6):963-973.
[34]	Ma L, Wang Y, Liu W, et al. Overexpression of an alfalfa GDP-mannose 3, 5-epimerase gene enhances acid, drought and salt tolerance in transgenic Arabidopsis by increasing ascorbate accumulation[J]. Biotechnol Lett, 2014, 36(11):2331-2341. doi: 10.1007/s10529-014-1598-y URL
[35]	Zhang Z, Wang Y, Chang L, et al. MsZEP, a novel zeaxanthin epoxidase gene from alfalfa(Medicago sativa), confers drought and salt tolerance in transgenic tobacco[J]. Plant Cell Reports, 2016, 35(2):439-453. doi: 10.1007/s00299-015-1895-5 URL
[36]	Wang Z, et al. Overexpression of alfalfa Orange gene in tobacco enhances carotenoid accumulation and tolerance to multiple abiotic stresses[J]. Plant Physiol Biochem, 2018, 130:613-622. doi: 10.1016/j.plaphy.2018.08.017 URL
[37]	Tan J, Zhuo C, Guo Z. Nitric oxide mediates cold and dehydration induced expression of a novel MfHyPRP that confers tolerance to abiotic stress[J]. Physiol Plant, 2013, 149(3):310-320.
[38]	黄思源. 干旱胁迫下蒺藜苜蓿的转录组分析及醛脱氢酶基因MtALDH7A1初步功能研究[D]. 杨凌:西北农林科技大学, 2019.
	Huang SY. Transcriptome analysis of Medicago truncatula and functional analysis of MtALDH7A1 gene under drought stress[D]. Yangling:North West Agriculture and Forestry University, 2019.
[39]	张业猛, 沈迎芳, 王英芳, 等. 蒺藜苜蓿MtLEA5B的克隆和功能分析[J]. 生物技术通报, 2018, 34(7):101-107.
	Zhang YM, Shen YF, Wang YF, et al. Cloning and Functional Analysis of the MtLEA5B Gene from Medicago truncatula[J]. Biotechnology Bulletin, 2018, 34(7):101-107.
[40]	Xie C, Zhang R, Qu Y, et al. Overexpression of MtCAS31 enhances drought tolerance in transgenic Arabidopsis by reducing stomatal density[J]. New Phytologist, 2012, 195(1):124-135. doi: 10.1111/j.1469-8137.2012.04136.x URL
[41]	师尚礼, 南丽丽, 郭全恩. 中国苜蓿育种取得的成就及展望[J]. 植物遗传资源学报, 2010, 11(1):46-51.
	Shi SL, Nan LL, Guo QE. Achievements and prospect of alfalfa breeding in china[J]. Journal of Plant Genetic Resources, 2010, 11(1):46-51.
[42]	Hubbard K, Hassanein N. Confronting coexistence in the United States:organic agriculture, genetic engineering, and the case of roundup ready alfalfa[J]. Agriculture and Human Values, 2013, 30(3):325-335. doi: 10.1007/s10460-012-9394-6 URL
[43]	Marita JM, Ralph J, Hatfield RD, et al. Structural and compositional modifications in lignin of transgenic alfalfa down-regulated in caffeic acid 3-O-methyltransferase and caffeoyl coenzyme A 3-O-methyltransferase[J]. Phytochemistry, 2003, 1:53-65.
[44]	Ashraf M, Akram NA. Improving salinity tolerance of plants through conventional breeding and genetic engineering:An analytical comparison[J]. Biotechnol Adv, 2009, 27(6):744-752. doi: 10.1016/j.biotechadv.2009.05.026 URL
[45]	Webb KJ. Transformation of forage legumes using Agrobacterium tumefaciens[J]. Theor Appl Genet, 1986, 72(1):53-58. doi: 10.1007/BF00261454 pmid: 24247771
[46]	Zhang JY, Broeckling CD, Blancaflor EB, et al. Overexpression of WXP1, a putative Medicago truncatula AP2 domain-containing transcription factor gene, increases cuticular wax accumulation and enhances drought tolerance in transgenic alfalfa(Medicago sativa L.)[J]. Plant Journal, 2005, 42(5):689-707. doi: 10.1111/tpj.2005.42.issue-5 URL
[47]	Tang LL, Cai H, Ji W, et al. Overexpression of GsZFP1 enhances salt and drought tolerance in transgenic alfalfa(Medicago sativa L.)[J]. Plant Physiology and Biochemistry, 2013, 71:22-30. doi: 10.1016/j.plaphy.2013.06.024 URL
[48]	Tang L, Cai H, Zhai H, et al. Overexpression of Glycine soja WRKY20 enhances both drought and salt tolerance in transgenic alfalfa(Medicago sativa L.)[J]. Plant Cell, Tissue and Organ Culture, 2014, 118(1):77-86. doi: 10.1007/s11240-014-0463-y URL
[49]	李静. 玉米ABP9基因转化紫花苜蓿及其抗旱性分析[D]. 兰州:兰州大学, 2012.
	Li J. Expression of ABP9 enhances tolerance to drought stress in transgenic alfalfa[D]. Lanzhou:Lanzhou University, 2012.
[50]	Wang Z, Su G, Li M, et al. Overexpressing Arabidopsis ABF3 increases tolerance to multiple abiotic stresses and reduces leaf size in alfalfa[J]. Plant Physiol Biochem, 2016, 109:199-208. doi: 10.1016/j.plaphy.2016.09.020 URL
[51]	Mckersie BD, Bowley SR, Harjanto E, et al. Water-deficit tolerance and field performance of transgenic alfalfa overexpressing superoxide dismutase[J]. Plant Physiology, 1996, 111(4):1177-1181. doi: 10.1104/pp.111.4.1177 URL
[52]	韩利芳, 张玉发. 烟草MnSOD基因在保定苜蓿中的转化[J]. 生物技术通报, 2004(1):39-42.
	Han LF, Zhang YF. The transformation of tobacco MnSOD gene into Baoding alfalfa[J]. Biotechnology Bulletin, 2004(1):39-46.
[53]	Bao A, Wang S, Wu G, et al. Overexpression of the Arabidopsis H⁺-PPase enhanced resistance to salt and drought stress in transgenic alfalfa(Medicago sativa L.)[J]. Plant Science, 2009, 176(2):232-240. doi: 10.1016/j.plantsci.2008.10.009 URL
[54]	Bao AK, Du BQ, Touil L, et al. Co-expression of tonoplast Cation/H⁺ antiporter and H⁺-pyrophosphatase from xerophyte Zygophyllum xanthoxylum improves alfalfa plant growth under salinity, drought and field conditions[J]. Plant Biotechnol J, 2016, 14(3):964-975. doi: 10.1111/pbi.2016.14.issue-3 URL
[55]	Li H, Wang Z, Ke Q, et al. Overexpression of codA gene confers enhanced tolerance to abiotic stresses in alfalfa[J]. Plant Physiology and Biochemistry, 2014, 85:31-40. doi: 10.1016/j.plaphy.2014.10.010 URL
[56]	Duan Z, Zhang D, Zhang J, et al. Co-transforming bar and CsALDH genes enhanced resistance to herbicide and drought and salt stress in transgenic alfalfa(Medicago sativa L.)[J]. Frontiers in Plant Science, 2015, 6:1115.
[57]	Wang Z, Ke Q, Kim MD, et al. Transgenic alfalfa plants expressing the sweet potato Orange gene exhibit enhanced abiotic stress tolerance[J]. PLoS One, 2015, 10(5):e126050.
[58]	Zhang J, Duan Z, Zhang DP, et al. Co-transforming bar and CsLEA enhanced tolerance to drought and salt stress in transgenic alfalfa(Medicago sativa L.)[J]. Biochemical and Biophysical Research Communications, 2016, 472(1):75-82. doi: 10.1016/j.bbrc.2016.02.067 pmid: 26906624
[59]	聂利珍, 刘红葵, 房永雨. 转AmDHN基因紫花苜蓿耐旱性的研究[J]. 北方农业学报, 2017, 45(1):6-9.
	Nie LZ, Liu HK, Fang YY. Study on enhancing tolerance to drought stress of transgenic alfalfa(Medicago sativa L.)with the AmDHN gene[J]. Journal of Northern Agriculture, 2017, 45(1):6-9.
[60]	Zheng GS, Fan C, Di S, et al. Over expression of Arabidopsis EDT1 gene confers drought tolerance in alfalfa(Medicago sativa L.)[J]. Frontiers in Plant Science, 2017, 8:2125. doi: 10.3389/fpls.2017.02125 URL
[61]	田野. 霸王ZxABCG11的功能验证及其对紫花苜蓿(Medicago sativa L.)的遗传转化[D]. 兰州:兰州大学, 2017.
	Tian Y. functional verification of ZxABCG11 from Zygophyllum xanthoxylum and genetic transformation to alfalfa（Medicago sativa L.）[D]. Lanzhou:Lanzhou University, 2017.
[62]	刘媛, 夏阳, 杨克强, 等. 渗透胁迫下转BADH基因苜蓿组培苗的抗性响应[J]. 中国农学通报, 2009, 25(4):133-136.
	Liu Y, Xia Y, Yang KQ, et al. Physiological responses to osmotic stress in Transgenic(BADH)Medicago Sativa[J]. Chinese Agricultural Science Bulletin, 2009, 25(4):133-136.
[63]	Suárez R, Calderón C, Iturriaga G. Enhanced tolerance to multiple abiotic stresses in transgenic alfalfa accumulating trehalose[J]. Crop Science, 2009, 49(5):1791-1799. doi: 10.2135/cropsci2008.09.0573 URL
[64]	Wang Z, Li H, Ke Q, et al. Transgenic alfalfa plants expressing AtNDPK2 exhibit increased growth and tolerance to abiotic stresses[J]. Plant Physiol Biochem, 2014, 84:67-77. doi: 10.1016/j.plaphy.2014.08.025 URL
[65]	Zhou C, Ma Z Y, Zhu L, et al. Overexpression of EsMcsu1 from the halophytic plant Eutrema salsugineum promotes abscisic acid biosynjournal and increases drought resistance in alfalfa(Medicago sativa L.)[J]. Genet Mol Res, 2015, 14(4):17204-17218. doi: 10.4238/2015.December.16.20 pmid: 26681214
[66]	Zhu J. Salt and drought stress signal transduction in plants[J]. Annual Review of Plant Biology, 2002, 53(1):247-273. doi: 10.1146/annurev.arplant.53.091401.143329 URL
[67]	马金星, 张吉宇, 单丽燕, 等. 中国草品种审定登记工作进展[J]. 草业学报, 2011, 20(1):206-213.
	Ma JX, Zhang JY, Shan LY, et al. Progress of the herbage variety approval and registration in China[J]. Acta Prataculturae Sinica, 2011, 20(1):206-213.

序号 No.	基因 Genes	基因来源 Gene sources	转基因物种 Genetically modified species	功能 Functions	参考文献 Reference
1	MsNAC2	紫花苜蓿（Medicago sativa）	烟草（Nicotiana tabacum）	丙二醛含量降低,脯氨酸含量、 SOD 和 POD 的活性提高 Reduce MDA content,increase proline content,SOD and POD activity	[26]
2	MfNAC35	黄花苜蓿（Medicago falcata）	烟草（N. tabacum）	抗逆相关基因 CAT、SOD、P5CS表达的上调 Up-regulation of stress-resistance related genes CAT,SOD,and P5CS	[27]
3	MfNACsa	黄花苜蓿（M. falcata）	蒺藜苜蓿（Medicago truncatula）	MfNACsa在脱水胁迫下直接结合Glyl启动子并激活其转录 MfNACsa directly binds to the Glyl promoter and activates its transcription under dehydration stress	[19]
4	MsMYB2L	紫花苜蓿（M. sativa）	拟南芥（Arabidopsis）	脯氨酸、可溶性糖增加,降低了脂质过氧化 Increase proline and soluble sugars and reduce lipid peroxidation	[28]
5	MfERF049	黄花苜蓿（M. falcata）	蒺藜苜蓿（M. truncatula）	干旱胁迫诱导MfERF049在根中的表达 Drought stress induces the expression of MfERF049 in roots	[29]
6	MfERF026	黄花苜蓿（M. falcata）	蒺藜苜蓿（M. truncatula）	干旱胁迫诱导MfERF026在根、茎和叶均表达 Drought stress induces the expression of MfERF026 in roots,stems and leaves	[30]
7	MfERF086	黄花苜蓿（M. falcata）	蒺藜苜蓿（M. truncatula）	干旱胁迫诱导MfERF086在根、茎和叶均表达 Drought stress induces the expression of MfERF086 in roots,stems and leaves	[30]
8	MsERF11	紫花苜蓿（M. sativa）	拟南芥（Arabidopsis）	干旱胁迫诱导MsERF11表达 Drought stress induces the expression of MsERF11	[31]
9	MsZIP	紫花苜蓿（M. sativa）	烟草（N. tabacum）	丙二醛含量、相对含水量、可溶性糖含量、可溶性蛋白含量和脯氨酸含量提高 Increase MDA content,relative water content（RWC）,soluble sugar content,soluble protein content and proline content	[32]
10	MtbHLH148	蒺藜苜蓿（M. truncatula ）	拟南芥（Arabidopsis）	根长增加 Increase root length	[33]
11	MsGME	紫花苜蓿（M. sativa）	拟南芥（Arabidopsis）	更低的MDA含量、较少膜损伤 Lower MDA content,and less membrane damage	[34]
12	MsZEP	紫花苜蓿（M. sativa）	烟草（N. tabacum）	相对含水量和ABA 含量提高,调节气孔导度及气孔开度 Increase RWC and ABA content,and adjust stomata conductance and stomata opening	[35]
13	MsOr	紫花苜蓿（M. sativa）	烟草（N. tabacum）	类胡萝卜素增加,丙二醛含量降低 increase carotenoids,decrease MDA content	[36]
14	MfHyPRP	黄花苜蓿（M. falcata）	烟草（N. tabacum）	转基因株系的鲜重显著高于野生型 The fresh weight of the transgenic line is significantly higher than that of the wild type	[37]
15	MtALDH7A1	蒺藜苜蓿（M. truncatula）	拟南芥（Arabidopsis）	影响根的生长及侧根数量的增多 Affect the growth of roots and increase the number of lateral roots	[38]
16	MtLEA5B	蒺藜苜蓿（M. truncatula）	大肠杆菌（Escherichia. coli）	胁迫后LEA蛋白仍能保持一定的结构 The LEA protein can still maintain a certain structure after stress	[39]
17	MtCAS31	蒺藜苜蓿（M. truncatula）	拟南芥（Arabidopsis）	降低气孔密度 Decrease stomatal density	[40]

序号 No.	基因 Genes	基因来源 Gene sources	转基因物种 Genetically modified species	功能 Functions	参考文献 Reference
1	MsNAC2	紫花苜蓿（Medicago sativa）	烟草（Nicotiana tabacum）	丙二醛含量降低,脯氨酸含量、 SOD 和 POD 的活性提高 Reduce MDA content,increase proline content,SOD and POD activity	[26]
2	MfNAC35	黄花苜蓿（Medicago falcata）	烟草（N. tabacum）	抗逆相关基因 CAT、SOD、P5CS表达的上调 Up-regulation of stress-resistance related genes CAT,SOD,and P5CS	[27]
3	MfNACsa	黄花苜蓿（M. falcata）	蒺藜苜蓿（Medicago truncatula）	MfNACsa在脱水胁迫下直接结合Glyl启动子并激活其转录 MfNACsa directly binds to the Glyl promoter and activates its transcription under dehydration stress	[19]
4	MsMYB2L	紫花苜蓿（M. sativa）	拟南芥（Arabidopsis）	脯氨酸、可溶性糖增加,降低了脂质过氧化 Increase proline and soluble sugars and reduce lipid peroxidation	[28]
5	MfERF049	黄花苜蓿（M. falcata）	蒺藜苜蓿（M. truncatula）	干旱胁迫诱导MfERF049在根中的表达 Drought stress induces the expression of MfERF049 in roots	[29]
6	MfERF026	黄花苜蓿（M. falcata）	蒺藜苜蓿（M. truncatula）	干旱胁迫诱导MfERF026在根、茎和叶均表达 Drought stress induces the expression of MfERF026 in roots,stems and leaves	[30]
7	MfERF086	黄花苜蓿（M. falcata）	蒺藜苜蓿（M. truncatula）	干旱胁迫诱导MfERF086在根、茎和叶均表达 Drought stress induces the expression of MfERF086 in roots,stems and leaves	[30]
8	MsERF11	紫花苜蓿（M. sativa）	拟南芥（Arabidopsis）	干旱胁迫诱导MsERF11表达 Drought stress induces the expression of MsERF11	[31]
9	MsZIP	紫花苜蓿（M. sativa）	烟草（N. tabacum）	丙二醛含量、相对含水量、可溶性糖含量、可溶性蛋白含量和脯氨酸含量提高 Increase MDA content,relative water content（RWC）,soluble sugar content,soluble protein content and proline content	[32]
10	MtbHLH148	蒺藜苜蓿（M. truncatula ）	拟南芥（Arabidopsis）	根长增加 Increase root length	[33]
11	MsGME	紫花苜蓿（M. sativa）	拟南芥（Arabidopsis）	更低的MDA含量、较少膜损伤 Lower MDA content,and less membrane damage	[34]
12	MsZEP	紫花苜蓿（M. sativa）	烟草（N. tabacum）	相对含水量和ABA 含量提高,调节气孔导度及气孔开度 Increase RWC and ABA content,and adjust stomata conductance and stomata opening	[35]
13	MsOr	紫花苜蓿（M. sativa）	烟草（N. tabacum）	类胡萝卜素增加,丙二醛含量降低 increase carotenoids,decrease MDA content	[36]
14	MfHyPRP	黄花苜蓿（M. falcata）	烟草（N. tabacum）	转基因株系的鲜重显著高于野生型 The fresh weight of the transgenic line is significantly higher than that of the wild type	[37]
15	MtALDH7A1	蒺藜苜蓿（M. truncatula）	拟南芥（Arabidopsis）	影响根的生长及侧根数量的增多 Affect the growth of roots and increase the number of lateral roots	[38]
16	MtLEA5B	蒺藜苜蓿（M. truncatula）	大肠杆菌（Escherichia. coli）	胁迫后LEA蛋白仍能保持一定的结构 The LEA protein can still maintain a certain structure after stress	[39]
17	MtCAS31	蒺藜苜蓿（M. truncatula）	拟南芥（Arabidopsis）	降低气孔密度 Decrease stomatal density	[40]

序号 No.	基因 Genes	功能 Functions	基因来源 Gene sources	转基因苜蓿品种Transgenic alfalfa varieties	抗旱性评价 Drought resistance evaluation	参考文献Reference
1	WXP1	乙烯反应元件结合转录因子 Ethylene response element binding transcription factor	蒺藜苜蓿 Medicago truncatula	Regen SY-4D	叶片表皮蜡质改变;提高净光合速率,蒸腾速率和气孔导度 Change wax in leaf cuticle; improve net photosynthetic rate,transpiration rate and stoma conductance	[46]
2	ZFP1	编码 Cys2/His2 型锌指蛋白 Encoding Cys2/His2 zinc finger protein	野生大豆 Glycine soja	肇东苜蓿 Zhandong	MtCOR47,MtRAB18,MtP5CS 和MtRD2表达量提高 Increased expression level of MtCOR47,MtRAB18,MtP5CS and MtRD2	[47]
3	WRKY20	转录因子 Transcription factor	野生大豆 G. soja	肇东苜蓿 Zhandong	脯氨酸和可溶性糖增加,角质层增厚 Increase content of proline and soluble sugar,and thicken stratum corneum	[48]
4	ABP9	bZIP家族的转录因子 Transcription factors of the bZIP family	玉米 Zea mays	保定苜蓿 Baoding alfalfa	较高相对含水量、叶绿素含量和脯氨酸;较低丙二醛含量 Higher RWC,chlorophyll content and proline; lower malondialdehyde content	[49]
5	ABF3	bZIP转录因子 Transcription factors of the bZIP	拟南芥 Arabidopsis thaliana	新疆大叶Xinjiang Daye	降低蒸腾速率和活性氧含量 Reduce transpiration rate and active oxygen content	[50]
6	Mn-SOD	锰超氧化物歧化酶Mn-SOD	烟草 Nicotiana plumbaginifolia	保定苜蓿 Baoding	MnSOD 活性高,SOD 活性增加 Higher MnSOD activity,increase SOD activity	[51-52]
7	AVP1	液泡H⁺-焦磷酶H⁺-PPase Vacuolar H⁺-PPase gene	拟南芥 A. thaliana	新疆大叶 Xinjiang Daye	根系中积累较多的Na⁺、K⁺和 Ca²⁺ Accumulate in the root system more Na⁺、K⁺ and Ca²⁺	[53]
8	NHX 和 VP1-1	Na⁺逆向转运蛋白和植物液泡膜H⁺-PPase基因 Na⁺ antiporter and plant vacuolar membrane H⁺-PPase gene	霸王 Zygophyllum xanthoxylum	新疆大叶Xinjiang Daye	叶片和根系中积累较多的Na⁺、K⁺和 Ca²⁺ Accumulate more Na⁺、K⁺ and Ca²⁺in the leaf and root	[54]
9	codA	胆碱氧化酶Choline oxidase	细菌 Bacteria	新疆大叶Xinjiang Daye	保持较高的相对水分含量,提高甘氨酸甜菜碱和脯氨酸的含量 Maintain a high RWC and increase the content of glycine betaine and proline	[55]
10	Bar和ALDH	醛脱氢酶 Aldehyde dehydrogenase	无芒隐子草 Cleistogenes songorica	金皇后The Golden Queen	Na⁺含量低,K⁺含量高,相对含水量较高 Low Na⁺ content,high K⁺ content,high relative water content	[56]
11	IbOr	橙色基因 Orange gene	甘薯 Ipomoea batatas	新疆大叶Xinjiang Daye	较高含水量,较低MDA含量 high RWC,low MDA content	[57]
12	bar和LEA	干旱胁迫应答因子 Drought stress response factor	无芒隐子草 C. songorica	金皇后 The Golden Quee	较高的相对含水量、较高的地上部生物量、膜损伤和渗透胁迫低 High RWC,high above-ground biomass,low membrane damage and osmotic stress	[58]
13	DHN	脱水素 Dehydrin	沙冬青 Ammopiptanthus mongolicus	中苜2 号 Zhongmu No. 2	Pro 的积累量增加,进一步诱导 Pro 合成途径相关基因的表达 The accumulation of Pro increases,which further induces the expression level of genes related to the pro synthesis pathway	[59]
14	EDT1	编码同域亮氨酸拉链家族的转录因子 Transcription factor of leucine zipper family	拟南芥 A. thaliana	中苜1号 Zhongmu No.1	降低膜透性和丙二醛含量,较高的可溶性糖和脯氨酸含量、超氧化物歧化酶活性和叶绿素含量 Reduce membrane permeability and MDA content,higher soluble sugar and proline content,superoxide dismutase activity and chlorophyll content	[60]
15	ABCG11	角质层脂质转运蛋白编码基因 Encoding lipid transport protein of stratum corneum	霸王 Z. xanthoxylum	新疆大叶 Xinjiang Daye	改变了角质和蜡质的成分、厚度,调节气孔开合程度来减少水分的蒸发量 Change the composition and thickness of the cuticle and wax,and adjust the degree of stomata opening and closing to reduce water evaporation	[61]
16	BADH	甜菜碱醛脱氢酶 Betaine Aldehyde Dehydrogenase	山菠菜 Atriplex hortensis	中苜1号 Zhongmu No.1	可溶性糖含量增加,脯氨酸含量减少 Increase soluble sugar content,and decrease proline content	[62]
17	TPS	6-磷酸海藻糖合酶（TPS） Trehalose synthase gene	酵母 Yeast	Regen SY27x	较高含水量 Higher water content	[63]
18	NDPK	核苷二磷酸激酶 Nucleoside diphosphate kinase	拟南芥 A. thaliana	新疆大叶 Xinjiang Daye	较高相对含水量,较高脯氨酸含量和较低MDA含量 Higher RWC,and higher proline content and lower MDA content	[64]
19	Mcsu1	编码钼辅因子硫化酶 Encoding molybdenum cofactor sulfoxygenase	山萮菜 Eutrema salsugineum	新疆大叶Xinjiang Daye	较低的ROS 含量、离子渗透和 MDA 含量 Lower reactive oxygen species（ROS）,ion leakage,and MDA content	[65]

序号 No.	基因 Genes	功能 Functions	基因来源 Gene sources	转基因苜蓿品种Transgenic alfalfa varieties	抗旱性评价 Drought resistance evaluation	参考文献Reference
1	WXP1	乙烯反应元件结合转录因子 Ethylene response element binding transcription factor	蒺藜苜蓿 Medicago truncatula	Regen SY-4D	叶片表皮蜡质改变;提高净光合速率,蒸腾速率和气孔导度 Change wax in leaf cuticle; improve net photosynthetic rate,transpiration rate and stoma conductance	[46]
2	ZFP1	编码 Cys2/His2 型锌指蛋白 Encoding Cys2/His2 zinc finger protein	野生大豆 Glycine soja	肇东苜蓿 Zhandong	MtCOR47,MtRAB18,MtP5CS 和MtRD2表达量提高 Increased expression level of MtCOR47,MtRAB18,MtP5CS and MtRD2	[47]
3	WRKY20	转录因子 Transcription factor	野生大豆 G. soja	肇东苜蓿 Zhandong	脯氨酸和可溶性糖增加,角质层增厚 Increase content of proline and soluble sugar,and thicken stratum corneum	[48]
4	ABP9	bZIP家族的转录因子 Transcription factors of the bZIP family	玉米 Zea mays	保定苜蓿 Baoding alfalfa	较高相对含水量、叶绿素含量和脯氨酸;较低丙二醛含量 Higher RWC,chlorophyll content and proline; lower malondialdehyde content	[49]
5	ABF3	bZIP转录因子 Transcription factors of the bZIP	拟南芥 Arabidopsis thaliana	新疆大叶Xinjiang Daye	降低蒸腾速率和活性氧含量 Reduce transpiration rate and active oxygen content	[50]
6	Mn-SOD	锰超氧化物歧化酶Mn-SOD	烟草 Nicotiana plumbaginifolia	保定苜蓿 Baoding	MnSOD 活性高,SOD 活性增加 Higher MnSOD activity,increase SOD activity	[51-52]
7	AVP1	液泡H⁺-焦磷酶H⁺-PPase Vacuolar H⁺-PPase gene	拟南芥 A. thaliana	新疆大叶 Xinjiang Daye	根系中积累较多的Na⁺、K⁺和 Ca²⁺ Accumulate in the root system more Na⁺、K⁺ and Ca²⁺	[53]
8	NHX 和 VP1-1	Na⁺逆向转运蛋白和植物液泡膜H⁺-PPase基因 Na⁺ antiporter and plant vacuolar membrane H⁺-PPase gene	霸王 Zygophyllum xanthoxylum	新疆大叶Xinjiang Daye	叶片和根系中积累较多的Na⁺、K⁺和 Ca²⁺ Accumulate more Na⁺、K⁺ and Ca²⁺in the leaf and root	[54]
9	codA	胆碱氧化酶Choline oxidase	细菌 Bacteria	新疆大叶Xinjiang Daye	保持较高的相对水分含量,提高甘氨酸甜菜碱和脯氨酸的含量 Maintain a high RWC and increase the content of glycine betaine and proline	[55]
10	Bar和ALDH	醛脱氢酶 Aldehyde dehydrogenase	无芒隐子草 Cleistogenes songorica	金皇后The Golden Queen	Na⁺含量低,K⁺含量高,相对含水量较高 Low Na⁺ content,high K⁺ content,high relative water content	[56]
11	IbOr	橙色基因 Orange gene	甘薯 Ipomoea batatas	新疆大叶Xinjiang Daye	较高含水量,较低MDA含量 high RWC,low MDA content	[57]
12	bar和LEA	干旱胁迫应答因子 Drought stress response factor	无芒隐子草 C. songorica	金皇后 The Golden Quee	较高的相对含水量、较高的地上部生物量、膜损伤和渗透胁迫低 High RWC,high above-ground biomass,low membrane damage and osmotic stress	[58]
13	DHN	脱水素 Dehydrin	沙冬青 Ammopiptanthus mongolicus	中苜2 号 Zhongmu No. 2	Pro 的积累量增加,进一步诱导 Pro 合成途径相关基因的表达 The accumulation of Pro increases,which further induces the expression level of genes related to the pro synthesis pathway	[59]
14	EDT1	编码同域亮氨酸拉链家族的转录因子 Transcription factor of leucine zipper family	拟南芥 A. thaliana	中苜1号 Zhongmu No.1	降低膜透性和丙二醛含量,较高的可溶性糖和脯氨酸含量、超氧化物歧化酶活性和叶绿素含量 Reduce membrane permeability and MDA content,higher soluble sugar and proline content,superoxide dismutase activity and chlorophyll content	[60]
15	ABCG11	角质层脂质转运蛋白编码基因 Encoding lipid transport protein of stratum corneum	霸王 Z. xanthoxylum	新疆大叶 Xinjiang Daye	改变了角质和蜡质的成分、厚度,调节气孔开合程度来减少水分的蒸发量 Change the composition and thickness of the cuticle and wax,and adjust the degree of stomata opening and closing to reduce water evaporation	[61]
16	BADH	甜菜碱醛脱氢酶 Betaine Aldehyde Dehydrogenase	山菠菜 Atriplex hortensis	中苜1号 Zhongmu No.1	可溶性糖含量增加,脯氨酸含量减少 Increase soluble sugar content,and decrease proline content	[62]
17	TPS	6-磷酸海藻糖合酶（TPS） Trehalose synthase gene	酵母 Yeast	Regen SY27x	较高含水量 Higher water content	[63]
18	NDPK	核苷二磷酸激酶 Nucleoside diphosphate kinase	拟南芥 A. thaliana	新疆大叶 Xinjiang Daye	较高相对含水量,较高脯氨酸含量和较低MDA含量 Higher RWC,and higher proline content and lower MDA content	[64]
19	Mcsu1	编码钼辅因子硫化酶 Encoding molybdenum cofactor sulfoxygenase	山萮菜 Eutrema salsugineum	新疆大叶Xinjiang Daye	较低的ROS 含量、离子渗透和 MDA 含量 Lower reactive oxygen species（ROS）,ion leakage,and MDA content	[65]