Advances in Studies on Identification Methods and Molecular Biology of Drought Resistance in Sorghum

doi:10.13560/j.cnki.biotech.bull.1985.2020-0941

Abstract

Abstract:

Sorghum[Sorghum bicolor（L.）Moench]is an important food,feed and energy crop in the world. Because of its developed root system,strong water absorption ability,high water utilization rate and strong drought tolerance,it is widely planted in arid and semi-arid areas,which has important research value in the field of crop drought resistance research. Drought is the main stress factor affecting the growth and development of sorghum. The drought resistances among different varieties of sorghum vary largely. Screening sorghum varieties with strong drought resistance is of importantly theoretical and practical significance for promoting the process of drought resistance breeding of sorghum. Identification methods and identification indexes of sorghum resistances during different growth and development periods,as well as the evaluation methods of drought resistant germplasm resources of sorghum are introduced in this paper. The progress of molecular genetic research on sorghum is reviewed from three aspects：transcriptome analysis,drought resistance QTL localization and drought resistance gene mining,and the development direction of drought resistance of sorghum is prospected,aiming to provide reference for further exploring the molecular mechanism of drought resistance of sorghum and speeding up molecular genetic breeding of drought resistance of sorghum.

Key words: sorghum, drought resistance identification, drought resistance indicators, molecular biology

JIANG Bai-yang, BAI Wen-bin, ZHANG Jiang-hua, FAN Na, SHI Li-juan. Advances in Studies on Identification Methods and Molecular Biology of Drought Resistance in Sorghum[J]. Biotechnology Bulletin, 2021, 37(4): 260-272.

References 91

[1]	屈春艳. 水旱条件下小麦产量性状和抗旱性的全基因组关联分析[D]. 泰安:山东农业大学, 2018.
	Qu CY. Genome-wide association study on yield traits and drought tolerance under irrigation and drought conditions in wheat[D]. Tai’an:Shandong Agricultural University, 2018.
[2]	李瑞雪, 孙任洁, 汪泰初, 等. 植物抗旱性鉴定评价方法及抗旱机制研究进展[J]. 生物技术通报, 2017,33(7):40-48.
	Li RX, Sun RJ, Wang TC, et al. Research progress on identification and evaluation methods, and mechanism of drought resistance in plants[J]. Biotechnology Bulletin, 2017,33(7):40-48.
[3]	Essien MA, Pistorius E. A review on physiological contrivances of drought tolerance in sorghum, genetic basis and breeding methods[J]. Global Journal of Plant Breeding and Genetics, 2016,3(8):248-259.
[4]	单长卷, 韩蕊莲, 梁宗锁. 干旱胁迫下黄土高原4种乡土禾草抗氧化特性[J]. 生态学报, 2012,32(4):1174-1184.
	Shan CJ, Han RL, Liang ZS. Antioxidant properties of four native grasses in Loess Plateau under drought stress[J]. Acta Ecologica Sinica, 2012,32(4):1174-1184.
[5]	Wang W, Vinocur B, Altman A. Plant responses to drought, salinityand extreme temperatures:towards genetic engineering for stress tolerance[J]. Planta, 2003,218:1-14. doi: 10.1007/s00425-003-1105-5 URL
[6]	刘晨阳, 张蕙杰, 辛翔飞. 世界高粱供需格局变动及趋势分析[J]. 中国食物与营养, 2020,26(3):42-46.
	Li CY, Zhang HJ, Xin XF. Analysis on changes and trends in the world’s sorghum supply and demand[J]. Food and Nutrition in China, 2020,26(3):42-46.
[7]	Hsiao TC, Fereres E, Acevedo E, Henderson DW. Water stress and dynamics of growth and yield of crop plants[M]. Berlin:Springer-Verlag Berlin Heidelberg, 1976: 281-305.
[8]	张锡梅, 徐勇. 谷子、糜子、高粱、玉米抗旱品种气孔扩散阻力、蒸腾速率、叶水势关系的研究[J]. 干旱地区农业研究, 1987,5(3):80-85.
	Zhang XM, Xu Y. A study of relation of leaf water potential with resistance of stomatic diffusion and evapotranspiration rate of millet, broom, corn millet, sorghum and corn[J]. Agricultural Research in the Arid Areas, 1987,5(3):80-85.
[9]	山仑, 徐炳成. 论高粱的抗旱性及在旱区农业中的地位[J]. 中国农业科学, 2009,42:2342-2348.
	Shan L, Xu BC. Discussion on drought resistance of sorghum and its status in agriculture in arid and semiarid regions[J]. Scientia Agricultura Sinica, 2009,42:2342-2348.
[10]	黄玉兰. PEG模拟干旱条件下烯效唑对薏苡幼苗生理生化的调控及转录组分析[D]. 大庆:黑龙江八一农垦大学, 2017.
	Huang YL. The physiological and biochemical regulation of uniconazole to Coix lachrymal-jobi L. seedlings and transcriptome analysis under PEG drought stress[D]. Daqing:Heilongjiang Bayi Agricultural University, 2017.
[11]	Bussis D, Kauder F, Eineke D. Acclimation of potato plants to polyethylene glycolinduced water deficit Ⅱ:contents and subcellular distribution of organic solutes[J]. J Exp Bot, 1998,49(325):1361-1370. doi: 10.1093/jxb/49.325.1361 URL
[12]	张立军, 樊金娟, 阮燕晔. 聚乙二醇在植物渗透胁迫生理研究中的应用[J]. 植物生理学通讯, 2004,40(3):361-364.
	Zhang LJ, Fan JJ, Ruan YY. Application of polyethylene glycol in the study of plant osmotic stress physiology[J]. Plant Physiology Communications, 2004,40(3); 361-364.
[13]	王平, 王春语, 丛玲, 等. PEG-6000模拟干旱胁迫下高粱种子萌发期抗旱、敏感材料筛选、鉴定[J]. 山西农业大学学报:自然科学版, 2020,40(3):30-36.
	Wang P, Wang CY, Cong L, et al. Screening and identification of drought-resistant and sensitive sorghum varieties under PEG-6000 simulated drought stress conditions at germination stage[J]. J Shanxi Agric Univ:Nature Science Edition, 2020,40(3):30-36.
[14]	陈冰嬬, 徐宁, 李淑杰. 高粱亲本系萌发期抗旱性鉴定[J]. 中国农业大学学报, 2018,23(8):17-29.
	Chen BR, Xu N, Li SJ. Identification of the drought resistance of sorghum parental lines at germination stage[J]. Journal of China Agricultural University, 2018,23(8):17-29.
[15]	王志恒, 魏玉清, 邹芳. PEG-6000和盐碱胁迫对甜高粱种子萌发影响研究[J]. 种子, 2019,38(5):39-43.
	Wang ZH, Wei YQ, Zhou F. Study on effects of PEG-6000 and saline-alkali stress on seed germination of sweet sorghum[J]. Seed, 2019,38(5):39-43.
[16]	吴奇, 周宇飞, 高悦. 不同高粱品种萌发期抗旱性筛选与鉴定[J]. 作物学报, 2016,42(8):1233-1246.
	Wu Q, Zhou YF, Gao Y. Screening and identification for drought resistance during germination in sorghum cultivars[J]. Acta Agronomica Sinica, 2016,42(8):1233-1246.
[17]	王艺陶, 周宇飞, 李丰先. 基于主成分和SOM聚类分析的高粱品种萌发期抗旱性鉴定与分类[J]. 作物学报, 2014,40(1):110-121.
	Wang YT, Zhou YF, Li FX. Identification and classification of sorghum cultivars for drought resistance during germination stage based on principal components analysis and self organizing map cluster analysis[J]. Acta Agronomica Sinica, 2014,40(1):110-121.
[18]	张丽霞, 王春语, 王平, 等. 种子萌发期高粱抗旱材料的筛选与鉴定[J]. 分子植物育种, 2018,16(17):5796-5803.
	Zhang LX, Wang CY, Wang P, et al. Screening and identification of drought-resistant materials of sorghum at seed germination stage[J]. Molecular Plant Breeding, 2018,16(17):5796-5803.
[19]	程量, 林良斌. 作物耐旱性生理生化指标研究进展[J]. 中国农学通报, 2014,30(3):27-31.
	Cheng L, Lin LB. The research progress of physiological and biochemical indexes about drought tolerance in crop[J]. Chinese Agricultural Science Bulletin, 2014,30(3):27-31.
[20]	Maribel LD, Satoshi T. Antioxidant responses of rice seedlings to salinity stress[J]. Plant Science, 1998,135(1):1-9. doi: 10.1016/S0168-9452(98)00025-9 URL
[21]	胡荣海, 昌小平. 反复干旱法的生理基础及其应用[J]. 华北农学报, 1996,11(3):51-56.
	Hu RH, Chang XP. The physiological base and utilization of repeated drought method[J]. Acta Agriculturae Boreali-Sinica, 1996,11(3):51-56.
[22]	Reddy BV, Ramaiah B, Kumar AA, et al. Evaluation of sorghum genotypes for stay-green trait and grain yield[J]. SAT Agric Res, 2007,3:1-4.
[23]	刘婷婷, 陈道钳, 王仕稳. 不同品种高粱幼苗在干旱复水过程中的生理生态响应[J]. 草业学报, 2018,27(6):100-110.
	Liu TT, Chen DQ, Wang SW. Physio-ecological responses to drought and subsequent re-watering in sorghum seeding[J]. Acta Prataculturae Sinica, 2018,27(6):100-110.
[24]	荣少英. 甜高粱和普通高粱对水分适应性的研究[D]. 郑州:河南大学, 2011.
	Rong SY. Study on water adaptation to sweet sorghum and grain sorghum[D]. Zhengzhou:Henan University, 2011.
[25]	Azarinasrabad A, Mousavinik SM, Galavi M, et al. Evaluation of water stress on yield, its components and some physiological traits at different growth stages in grain sorghum genotypes[J]. Notulae Sci Biol, 2016,8:204-210.
[26]	解芳, 翟国伟, 邹桂花. 干旱胁迫对高粱苗期抗旱生理特性的影响[J]. 浙江农业学报, 2012,24(5):53-757.
	Xie F, Zhai GW, Zou GH. Effect of drought stress on physiological characteristics of sorghumm at seedling stage[J]. Acta Agriculturae Zhejiangensis, 2012,24(5):53-757.
[27]	Bibi A, Sadaqat HA, Tahir MFN, et al. Screening of sorghum(Sorghum bicolor var Moench)for drought tolerance at seedling stage in polyethylene glycol[J]. The Journal of Animal & Plant Sciences, 2012,22:671-678.
[28]	刘方明, 高玉山, 孙云云. 高粱抗旱性鉴定研究进展[J]. 东北农业科学, 2016,41(3):5-7.
	Liu FM, Gao YS, Sun YY. Advances of researches on identification of drought resistance of sorghum[J]. Journal of Northeast Agricultural Sciences, 2016,41(3):5-7.
[29]	张文英, 智慧, 柳斌辉. 谷子全生育期抗旱性鉴定及抗旱指标筛选[J]. 植物遗传资源学报, 2010,11(5):560-565.
	Zhang WY, Zhi H, Liu BH. Indexes screening for drought resistance test of foxtail millet[J]. Journal of Plant Genetic Resources, 2010,11(5):560-565.
[30]	李龙. 普通菜豆种质资源抗旱性鉴定与抗旱生理特性研究[D]. 北京:中国农业科学院, 2014.
	Li L. Drought resistance identification of germplasm resources and study on physiological characteristics related to drought resistance in common bean(Phaseolus vulgaris L.)[D]. Beijing:Chinese Academy of Agricultural Sciences, 2014.
[31]	杨彬, 张一中, 周福平. 高粱抗旱性鉴定方法及相关QTL定位研究进展[J]. 植物遗传资源学报, 2019,20(3):496-507.
	Yang B, Zhang YZ, Zhou FP. Advances in studies in identification methods and QTL mapping of drought resistance in sorghum[J]. Journal of Plant Genetic Resources, 2019,20(3):496-507.
[32]	袁闯, 许兴, 毛桂莲. 甜高粱成熟期抗旱性指标筛选与评价方法[J]. 干旱地区农业研究, 2020,38(1):280-289.
	Yuan C, Xu X, Mao GL. Screening and evaluation methods of drought resistance index of sweet sorghum at maturity[J]. Agricultural Research in the Arid Areas, 2020,38(1):280-289.
[33]	王玉斌, 平俊爱, 牛皓. 粒用高粱种质中后期抗旱性鉴定筛选与分类指标评价[J]. 中国农业科学, 2019,52(22):4039-4049.
	Wang YB, Ping JA, Niu H. Evaluation of identification and classification index for drought resistance at middle and late growth stage in grain sorghum germplasms[J]. Scientia Agricultura Sinica, 2019,52(22):4039-4049.
[34]	吕鑫, 张福耀, 平俊爱, 等. 25个饲草高粱恢复系主要农艺性状及其抗旱性的相关分析[J]. 中国农学通报, 2013,29(29):6-13.
	Lü X, Zhang FY, Ping JA, et al. Correlation analysis of 25 main agronomic characters and drought resistance in forage sorghum restorer lines[J]. Chinese Agricultural Science Bulletin, 2013,29(29):6-13.
[35]	Winter SR, Musick JT, Porter KB. Evaluation of screening techniques for breeding drought-resistance winter wheat[J]. Crop Sci, 1988,28(3):512-516. doi: 10.2135/cropsci1988.0011183X002800030018x URL
[36]	Clarke JM, DePauw RM, Townley-Smith TF. Evaluation of methods for quantification of drought tolerance in wheat[J]. Crop Sci, 1992,32:723-728. doi: 10.2135/cropsci1992.0011183X003200030029x URL
[37]	兰巨生, 胡福顺, 张景瑞. 作物抗旱指数的概念和统计方法[J]. 华北农学报, 1990,5(2):20-25.
	Lan JS, Hu FS, Zhang JR. The concept and statistical method of drought resistance index in crops[J]. Acta Agriculturae Boreali-Sinica, 1990,5(2):20-25.
[38]	王瑞, 张福耀, 王花云, 等. 高粱抗旱种质筛选及遗传多样性的SSR分析[J]. 植物遗传资源学报, 2014,15(4):871-876.
	Wang R, Zhang FY, Wang HY, et al. Drought resistance screening and genetic diversity by SSR markers in sorghum[J]. Journal of Plant Genetic Resources, 2014,15(4):871-876.
[39]	Tuinstra MR, Grote EM, Goldsbrough PB, et al. Genetic analysis of post-flowering drought tolerance and components of grain development in Sorghum bicolor(L.)Moench[J]. Molecular Breeding, 1997,3(6):439-448. doi: 10.1023/A:1009673126345 URL
[40]	Abraha T, Nyende AB, Mwangi SG, et al. Identification of Sorghum(Sorghum bicolor L. Moench)landraces tolerant to post flowering drought stress using drought tolerance indices[J]. Journal of Plant Breeding and Crop Science, 2015,7(7):211-218. doi: 10.5897/JPBCS URL
[41]	汪灿, 周棱波, 张国兵. 酒用糯高粱资源成株期抗旱性鉴定及抗旱指标筛选[J]. 中国农业科学, 2017,50(8):1388-1402.
	Wang C, Zhou LB, Zhang GB. Drought resistance identification and drought resistance indices screening of liquor-making waxy sorghum resources at adult plant stage[J]. Scientia Agricultura Sinica, 2017,50(8):1388-1402.
[42]	高雪. 不同品系(种)甜高粱对盐胁迫和干旱胁迫的生理响应研究[D]. 银川:宁夏大学, 2018.
	Gao X. Study on physiological responses of different strains of sweet sorghum to salinity stress and drought stress[D]. Yinchuan:Ningxia University, 2018.
[43]	栗雨勤, 张文英, 王有增. 作物抗旱性鉴定指标研究及进展[J]. 河北农业科学, 2004,8(1):58-61.
	Li YQ, Zhang WY, Wang YZ. Advances of studies on crop drought resistance identification[J]. Journal of Hebei Agricultural Sciences, 2004,8(1):58-61.
[44]	Hasan SA, Rabei SH, Nada RM, et al. Water use efficiency in the drought-stressed sorghum and maize in relation to expression of aqua-porin genes[J]. Biologia Plantarum, 2017,61(9):127-137. doi: 10.1007/s10535-016-0656-9 URL
[45]	Guha A, Chhajed SS, Choudhary S, et al. Hydraulic anatomy affects genotypicvariation in plant water use and shows differential organ specific plasticity to drought in Sorghum bicolor[J]. Environmental and Experimental Botany, 2018,15(7):25-37.
[46]	谭静, 刘帆, 李自卫. 玉米品种耐旱性鉴定及耐旱指标筛选[J]. 西南农业学报, 2013,26(1):26-31.
	Tan J, Liu F, Li ZW. Screening of drought tolerant Maize hybrids and identifying for drought indicators[J]. Southwest China Journal of Agricultural Sciences, 2013,26(1):26-31.
[47]	周华, 张新, 刘腾云. 高通量转录组测序的数据分析与基因发掘[J]. 江西科学, 2012,30(5):607-611.
	Zhou H, Zhang X, Liu TY. Data processing and gene discovery of high-throughput transcriptome sequencing[J]. Jiangxi Science, 2012,30(5):607-611.
[48]	Velculescu VE, Zhang L, Vogelstein B, et al. Serialanalysis of gene expression[J]. Science, 1995,270(5235):484-487. doi: 10.1126/science.270.5235.484 URL
[49]	岳桂东, 高强, 罗龙海. 高通量测序技术在动植物研究领域中的应用[J]. 中国科学:生命科学, 2012,42(2):107-124.
	Yue GD, Gao Q, Luo LH. The application of high-throughput sequencing technology in plant and animal research[J]. Scientia Sinica Vitae, 2012,42(2):107-124.
[50]	Dugas DV, Monaco MK, Olsen A, et al. Functional annotation of thetranscriptome of Sorghum bicolor in response to osmoticstress and abscisic acid[J]. BMC Genomics, 2011,12(1):514-514. doi: 10.1186/1471-2164-12-514 URL
[51]	Fracasso A, Trindade LM, Amaducci S. Droughtstress tolerance strategies revealed by RNA-Seq in twosorghum genotypes with contrasting WUE[J]. BMC Plant Biology, 2016,16(1):115-115. doi: 10.1186/s12870-016-0800-x URL
[52]	Wang B, Regulski M, Tseng E, et al. A comparative transcriptional landscape of Maize and sorghum obtained by single-molecule sequencing[J]. Genome Research, 2018,28:921-928. doi: 10.1101/gr.227462.117 pmid: 29712755
[53]	邵丹阳. 干旱胁迫下甜高粱(辽甜一)转录组分析[D]. 郑州:河南大学, 2018.
	Shao DY. Transcriptome analysis of Sorghum dochna(LT-1)under water stress[D]. Zhengzhou:Henan University, 2018.
[54]	Shimada S, Makita Y, Kuriyama-Kondou T, et al. Functional and expression analyses of transcriptsbased on full-length cDNAs of sorghum bicolor[J]. DNA Research, 2015,22(6):485-493. doi: 10.1093/dnares/dsv030 URL
[55]	倪先林, 赵甘霖, 刘天朋, 等. 高粱重要抗性性状的基因定位研究进展[J]. 福建农业学报, 2012,27(6):652-660.
	Ni XL, Zhao GL, Liu TP, et al. Advances in sorghum resistance gene mapping[J]. Fujian Journal of Agricultural Sciences, 2012,27(6):652-660.
[56]	Badigannavar A, Teme N, Oliveira CA, et al. Physiological, genetic and molecular basis of drought resilience in sorghum[Sorghum bicolor(L.)Moench][J]. Indian Journal of Plant Physiology, 2018,23:670-688. doi: 10.1007/s40502-018-0416-2 URL
[57]	Kebede H, Subudhi PK, Rosenow DT, et al. Quantitative trait loci influencing drought tolerance in grain sorghum(Sorghum bicolor L. Moench)[J]. Theoretical and Applied Genetics, 2001,103:266-276. doi: 10.1007/s001220100541 URL
[58]	Tuinstra MR, Grote EM, Goldsbrough PB, et al. Identification of quantitative trait loci associated with pre-flowering drought tolerance in sorghum[J]. Crop Science, 1996,36:1337-1344. doi: 10.2135/cropsci1996.0011183X003600050043x URL
[59]	赵辉. 高粱的抗旱性比较及抗旱相关性状的QTL定位[D]. 杭州:浙江师范大学, 2014.
	Zhao H. Investigation of the variations of drought resistance and identification of QTLs for drought-related traitsin sorghum(Sorghum bicolor L. Moench)[D]. Hangzhou:Zhejiang Normal University, 2014.
[60]	Thomas H, Howarth CJ. Five ways to stay green[J]. Journal of Experimental Botany, 2000,51:329-337. doi: 10.1093/jexbot/51.suppl_1.329 URL
[61]	Thomas H, Rogers LJ. Turning over an old leaf[J]. University of Wales Review Science and Technology, 1989(6):29-38.
[62]	Thomas H, Smart CM. Crops that stay green[J]. Ann Appl Biol, 1993,123:193-219. doi: 10.1111/aab.1993.123.issue-1 URL
[63]	Haussmann BIG, Mahalakshmi V, Reddy BVS, et al. QTL mapping of stay-green in two sorghum recombinantinbred populations[J]. Theoretical and Applied Genetics, 2002,106(1):133-142. pmid: 12582881
[64]	Tao YZ, Henzell RG, Jordan DR, et al. Identification of genomic regions associated with stay green in sorghum by testing RIL in multiple environments[J]. Theor Appl Genet, 2000,100(8):1225-1232. doi: 10.1007/s001220051428 URL
[65]	王立东. 高粱开花前后叶片持绿性状QTL分析[D]. 兰州:兰州大学, 2016.
	Wang LD. QTL analysis for leaf stay-green traits at flowering stage in Sorghum bicolor[D]. Lanzhou:Lanzhou University, 2016.
[66]	Xu W, Rosenow DT, Nguyen HT. Stay green trait in grain sorghum:relationship between visual rating and leafchlorophyll concentration[J]. Plant Breeding, 2000,119(4):365-367. doi: 10.1046/j.1439-0523.2000.00506.x URL
[67]	Subudhi PK, Rosenow DT, Nguyen HT. Quantitative trait loci for the stay green trait insorghum(Sorghum bicolor L. moench): consistency across genetic backgrounds and environments[J]. Theoretical and Applied Genetics, 2000,101(5-6):733-741. doi: 10.1007/s001220051538 URL
[68]	Harris K, Subudhi PK, Borrell A, et al. Sorghum stay-green QTL individually reduce post-flowering drought-inducedleaf senescence[J]. Journal of Experimental Botany, 2007,58(2):327-338. doi: 10.1093/jxb/erl225 URL
[69]	Subedi KD, Ma BL. Nitrogen uptake and partitioning in stay-green and leafy maize hybrids[J]. Crop Sci, 2005,45(2):740-747. doi: 10.2135/cropsci2005.0740 URL
[70]	Crasta RR, Xu W, Rosenow DT, et al. Mapping of Post-flowering drought resistancetraits in grain sorghum:association of QTLs influencing premature senescence and maturity[J]. Mol Gen Genet, 1999,262:579-588. doi: 10.1007/s004380051120 URL
[71]	Xu W, Subudhi PK, Crasta OR, et al. Molecular mapping of QTLs conferring stay-green in grain sorghum(Sorghum bicolor L. Moench)[J]. Genome, 2000,43:461-469. pmid: 10902709
[72]	梁文君, 孙超, 李鹏程, 等. 作物根系发育相关基因研究进展及在马铃薯抗旱育种中的应用展望[J]. 植物生理学报, 2020,56(4):613-624.
	Liang WJ, Sun C, Li PC, et al. Research advances in genes related to crop root development and its application prospects in drought-resistant potato breeding[J]. Plant Physiology Journal, 2020,56(4):613-624.
[73]	Olsen AN, Ernst HA, Leggio LL, et al. NAC transcription factors:structurally distinct, functionally diverse[J]. Trends Plant Sci, 2005,10(2):79-87. doi: 10.1016/j.tplants.2004.12.010 URL
[74]	Gao F, Xiong AS, Peng RH, et al. OsNAC52a rice NAC transcription factor, potentially responds to ABA andconfers drought tolerance in transgenic plants[J]. Plant Cell Tiss Organ Cult, 2010,100(3):255-262. doi: 10.1007/s11240-009-9640-9 URL
[75]	Hu HH, You J, Fang YJ, et al. Characterization of transcription factor gene SNAC2 conferring cold and salt tolerancein rice[J]. Plant Mol Biol, 2008,67(1-2):169-181. doi: 10.1007/s11103-008-9309-5 URL
[76]	Song SY, Chen Y, Chen J, et al. Physiological mechanisms underlying OsNAC5-dependent tolerance of rice plants toabiotic stress[J]. Planta, 2011,234(2):331-345. doi: 10.1007/s00425-011-1403-2 URL
[77]	祖祎祎, Ibadeti K, 孙清鹏, 高粱SbNAC0584基因克隆与表达分析[J]. 山东农业大学学报:自然科学版, 2015,46(4):497-502.
	Zu YY, Ibadeti K, Sun QP. The analysis on the cloning and expression of SbNAC0584 gene in Sorghum bicolor L.[J]. Journal of Shandong Agricultural University:Natural Science Edition, 2015,46(4):497-502.
[78]	卢敏. 玉米ZmSNAC1和高粱SbSNAC1基因的克隆与功能分析[D]. 北京:中国农业科学院, 2013.
	Lu M. Cloning and functional analysis of ZmSNAC1 in maize(Zea mays L.)and SbSNAC1gene in sorghum[Sorghum bicolor(L.)Moench][D]. Beijing:Chinese Academy of Agricultural Sciences, 2013.
[79]	Xing HT, Guo P, Xia XL, et al. Pd ERECTA, a leucine-rich repeat receptor-like kinase of poplar, confers enhanced water use efficiency in Arabidopsis[J]. Planta, 2011,234(2):229-241. doi: 10.1007/s00425-011-1389-9 pmid: 21399949
[80]	Liu J, Zhang F, Zhou J, et al. Phytochrome B control of total leaf area andstomatal density affects drought tolerance in rice[J]. Plant Molecular Biology, 2012,78(3):289-300. doi: 10.1007/s11103-011-9860-3 URL
[81]	韩小东. 高粱ERECTA家族基因的克隆及其干旱胁迫相对表达水平的分析[D]. 南昌:江西农业大学, 2015.
	Han SD. The clone of ERECTA family gene from Sorghum and relative expression levels analysis of drought stress[D]. Nanchang:Jiangxi Agricultural University, 2015.
[82]	李韩帅. 高粱SbER2基因的抗旱生物学功能解析[D]. 太原:山西大学, 2018.
	Li HS. Biological functional analysis of SbER2 in response to drought stress in sorghum[D]. Taiyuan:Shanxi University, 2018.
[83]	刘洋. 高粱SbSKIP基因的克隆及其在烟草中的抗旱功能分析[J]. 农业生物技术学报, 2016,24(10):1500-1511.
	Liu Y. Cloning of SbSKIP gene from Sorghum(Sorghum bicolor)and analysis of drought-resistant function in tobacco(Nicotiana tabacum)[J]. Journal of Agricultural Biotechnology, 2016,24(10):1500-1511.
[84]	刁现民. 禾谷类杂粮作物耐逆和栽培技术研究新进展[J]. 中国农业科学, 2019,52(22):3943-3949.
	Diao XM. Progresses in stress tolerance and field cultivation studies of qrphan cereals in China[J]. Scientia Agricultura Sinica, 2019,52(22):3943-3949.
[85]	Li HS, Han XD, Liu XX, et al. A leucine-rich repeat-receptor-like kinase gene SbER2-1 from sorghum(Sorghum bicolor L.)confers drought tolerance in maize[J]. BMC Genomics, 2019,20(1): 1-15. doi: 10.1186/s12864-018-5379-1 URL
[86]	Mallory A, Vaucheret H. Form function and regulation of ARGONAUTE proteins[J]. Plant Cell, 2010,22(12):3879-3889. doi: 10.1105/tpc.110.080671 URL
[87]	Cao JY, Xu YP, Zhao L, et al. Tight regulation of the interaction between Brassica napus and Sclerotinia sclerotiorum at the microRNA level[J]. Plant Molecular Biology, 2016,92(1/2):39-55. doi: 10.1007/s11103-016-0494-3 URL
[88]	徐磊, 胡小文, 姚艳丽. 甜高粱WRKY转录因子基因的克隆与表达分析[J]. 西南农业学报, 2017,30(11):2429-2435.
	Xu L, Hu XW, Yao YL. Cloning and expression analysis of WRKY transcription factor gene in sweet sorghum[J]. Southwest China Journal of Agricultural Sciences, 2017, 30(11):2429-2435.
[89]	万庆. 甜高粱中SSWRKY28、SSWRKY76基因的克隆和功能分析[D]. 郑州:河南大学, 2018.
	Wan Q. Cloning and expression analysis of SSWRKY28 and SSWRKY76 genes in Sorghum dochna[D]. Zhengzhou:Henan University, 2018.
[90]	林俊俊, 郭怀刚, 董洁静. 高粱AGO蛋白家族基因鉴定及表达分析[J]. 核农学报, 2019,33(7):1291-1302.
	Lin JJ, Guo HG, Dong JJ. Identification and expression analysis of AGO protein family genes in Sorghum bicolor[J]. Journal of Nuclear Agricultural Sciences, 2019,33(7):1291-1302.
[91]	吉林省质量技术监督局. DB22/T2388-2015高粱抗旱性鉴定技术规程[S]. 北京: 中国标准出版社, 2015.
	Jilin provincial bureau of quality and technical supervision. DB22/T2388-2015 technical specification for drought resistance identification of sorghum[S]. Beijing: Standards Press of China, 2015.