作物学报 ›› 2026, Vol. 52 ›› Issue (11): 3264-3278.doi: 10.3724/SP.J.1006.2026.65034
袁荣1,**,杜媛媛1,**,洪美艳1,高界1,黄镇2,李可琪2,*,曾新华1,*
Yuan Rong1,**,Du Yuan-Yuan1,**,Hong Mei-Yan1,Gao Jie1,Huang Zhen2,Li Ke-Qi2,*,Zeng Xin-Hua1,*
摘要: 甘蓝型油菜是我国重要的油料作物,干旱胁迫严重影响其发芽率和幼苗建成,进而导致减产。为深入挖掘甘蓝型油菜种子萌发期耐旱基因,本研究以PEG-6000溶液模拟干旱胁迫处理,对183份甘蓝型油菜种质进行萌发期耐旱性鉴定和筛选,测定发芽率(GR)、发芽势(GP)、发芽指数(GI)、成苗率(SP)、活力指数(VI)、芽鲜重(SFW)、根鲜重(RFW)、根长(RL)等指标。通过全基因组关联分析(GWAS),共检测到9个显著关联的SNP位点,鉴定出213个显著性关联基因。为进一步探究这些遗传位点背后的分子机制,选取6个种子萌发期耐旱性存在显著差异的种质进行转录组分析,在0%和15%质量分数PEG-6000溶液处理下,6个种质分别检测出13,251、10,250、10,455、8968、9990、16,706个差异表达基因(DEGs)。这些DEGs主要富集于“植物激素信号转导”、“植物-病原体相互作用”、“MAPK信号通路-植物”、“苯丙素的生物合成”和“淀粉和蔗糖代谢”等与干旱响应密切相关的KEGG通路。通过整合GWAS和RNA-seq数据,最终筛选出7个与种子萌发期耐旱相关的候选基因。其中,ZS11A03G038720、ZS11A09G011050、ZS11C02G013040和ZS11C02G013790在PEG处理后均下调,ZS11A03G039060、ZS11C02G013640和ZS11C02G013660在PEG处理后均上调。研究结果不仅揭示了甘蓝型油菜种子萌发期耐旱性的关键遗传位点,还筛选出7个具有重要育种应用价值的核心候选基因,为研究油菜萌发期耐旱机理和通过分子辅助选育耐旱油菜品种提供了基因资源和理论参考。
| [1] Nguyen T C T, Abrams S R, Friedt W, et al. Quantitative trait locus analysis of seed germination, seedling vigour and seedling-regulated hormones in Brassica napus. Plant Breed, 2018, 137: 388–401. [2] El Balla M M A, Hamid A A, Abdelmageed A H A. Effects of time of water stress on flowering, seed yield and seed quality of common onion (Allium cepa L.) under the arid tropical conditions of Sudan. Agric Water Manag, 2013, 121: 149–157. [3] Liu Y, Xu H, Wen X X, et al. Effect of polyamine on seed germination of wheat under drought stress is related to changes in hormones and carbohydrates. J Integr Agric, 2016, 15: 2759–2774. [4] Salami M, Heidari B, Alizadeh B, et al. Dissection of quantitative trait nucleotides and candidate genes associated with agronomic and yield-related traits under drought stress in rapeseed varieties: integration of genome-wide association study and transcriptomic analysis. Front Plant Sci, 2024, 15: 1342359. [5] 刘婷婷, 庞进平, 徐一涌. 干旱胁迫对不同油菜品种苗期生物量和根系的影响. 甘肃农业科技, 2019, 50(6): 4–8. Liu T T, Pang J P, Xu Y Y. Effects of drought stress on seedling biomass and root system of different rapeseed cultivars. Gansu Agric Sci Technol, 2019, 50(6): 4–8 (in Chinese with English abstract). [6] Muscolo A, Sidari M, Anastasi U, et al. Effect of PEG-induced drought stress on seed germination of four lentil genotypes. J Plant Interact, 2014, 9: 354–363. [7] Hellal F A, El-Shabrawi H M, Abd El-Hady M, et al. Influence of PEG induced drought stress on molecular and biochemical constituents and seedling growth of Egyptian barley cultivars. J Genet Eng Biotechnol, 2018, 16: 203–212. [8] Khanzada H, Wassan G M, He H H, et al. Differentially evolved drought stress indices determine the genetic variation of Brassica napus at seedling traits by genome-wide association mapping. J Adv Res, 2020, 24: 447–461. [9] 张天瑶, 李鸿洋, 杜媛媛, 等. PEG模拟干旱下甘蓝型油菜萌发期耐旱性全基因组关联分析. 植物遗传资源学报, 2024, 25: 2058–2068. Zhang T Y, Li H Y, Du Y Y, et al. Genome-wide association analysis of drought tolerance in Brassica napus during germination under PEG simulated drought. J Plant Genet Resour, 2024, 25: 2058–2068 (in Chinese with English abstract). [10] Batool M, El-Badri A M, Wang Z K, et al. Rapeseed morpho-physio-biochemical responses to drought stress induced by PEG-6000. Agronomy, 2022, 12: 579. [11] Fang S, Zhao P M, Tan Z D, et al. Combining physio-biochemical characterization and transcriptome analysis reveal the responses to varying degrees of drought stress in Brassica napus L. Int J Mol Sci, 2022, 23: 8555. [12] Khan S U, Saeed S, Khan M H U, et al. Advances and challenges for QTL analysis and GWAS in the plant-breeding of high-yielding: a focus on rapeseed. Biomolecules, 2021, 11: 1516. [13] Lu G Y, Tian Z T, Chen P Y, et al. Comprehensive morphological and molecular insights into drought tolerance variation at germination stage in Brassica napus accessions. Plants, 2024, 13: 3296. [14] Ahmad N, Ibrahim S, Kuang L Q, et al. Integrating genome-wide association study with transcriptomic data to predict candidate genes influencing Brassica napus root and biomass-related traits under low phosphorus conditions. Biotechnol Biofuels Bioprod, 2023, 16: 149. [15] Xiang X R, Qiu P, Mei Z C, et al. Genome-wide association study and transcriptome analysis reveal natural variation of key genes regulation flowering time in rapeseed. Mol Breed, 2024, 44: 40. [16] 陈致富, 李勤菲, 张永晶, 等. 白菜型油菜品种萌发期的抗旱性鉴定与筛选. 植物遗传资源学报, 2015, 16: 15–22. Chen Z F, Li Q F, Zhang Y J, et al. Identification and screening of resources with tolerance against drought stress in Brassica rapa during germination stage. J Plant Genet Resour, 2015, 16: 15–22 (in Chinese with English abstract). [17] Clarke W E, Higgins E E, Plieske J, et al. A high-density SNP genotyping array for Brassica napus and its ancestral diploid species based on optimised selection of single-locus markers in the allotetraploid genome. Theor Appl Genet, 2016, 129: 1887–1899. [18] Pritchard J K, Stephens M, Donnelly P. Inference of population structure using multilocus genotype data. Genetics, 2000, 155: 945–959. [19] Zhang L Y, Yang B, Li X D, et al. Integrating GWAS, RNA-Seq and functional analysis revealed that BnaA02.SE mediates silique elongation by affecting cell proliferation and expansion in Brassica napus. Plant Biotechnol J, 2024, 22: 2907–2920. [20] 张超, 杨博, 张立源, 等. 基于QTL定位和全基因组关联分析挖掘甘蓝型油菜收获指数相关位点. 作物学报, 2022, 48: 2180–2195. Zhang C, Yang B, Zhang L Y, et al. Mining harvest index loci based on QTL mapping and genome-wide association study in rapeseed (Brassica napus L.). Acta Agron Sin, 2022, 48: 2180–2195 (in Chinese with English abstract). [21] 蒙姜宇, 梁光伟, 贺亚军, 等. 甘蓝型油菜耐盐和耐旱相关性状的QTL分析. 作物学报, 2021, 47: 462–471. Meng J Y, Liang G W, He Y J, et al. QTL mapping of salt and drought tolerance related traits in Brassica napus L. Acta Agron Sin, 2021, 47: 462–471 (in Chinese with English abstract). [22] 李阳阳, 吴丹, 许军红, 等. 基于QTL和转录组测序鉴定甘蓝型油菜耐旱候选基因. 作物学报, 2024, 50: 820–835. Li Y Y, Wu D, Xu J H, et al. Identification of candidate genes associated with drought tolerance based on QTL and transcriptome sequencing in Brassica napus L. Acta Agron Sin, 2024, 50: 820–835 (in Chinese with English abstract). [23] Dubos C, Stracke R, Grotewold E, et al. MYB transcription factors in Arabidopsis. Trends Plant Sci, 2010, 15: 573–581. [24] Ding Z H, Li S M, An X L, et al. Transgenic expression of MYB15 confers enhanced sensitivity to abscisic acid and improved drought tolerance in Arabidopsis thaliana. J Genet Genom, 2009, 36: 17–29. [25] Kim H, Hwang H, Hong J W, et al. A rice orthologue of the ABA receptor, OsPYL/RCAR5, is a positive regulator of the ABA signal transduction pathway in seed germination and early seedling growth. J Exp Bot, 2012, 63: 1013–1024. [26] Andrási N, Pettkó-Szandtner A, Szabados L. Diversity of plant heat shock factors: regulation, interactions, and functions. J Exp Bot, 2021, 72: 1558–1575. [27] Zhang F, Pan Z J, Han C Y, et al. Pyrus betulaefolia ERF3 interacts with HsfC1a to coordinately regulate aquaporin PIP1;4 and NCED4 for drought tolerance. Hortic Res, 2024, 11: uhae090. [28] Kumar A, Partap M, Warghat A R. From growth to survival: Aux/IAA genes in plant development and stress management. Plant Sci, 2026, 362: 112750. [29] Su X, Zhang X L, Luo J, et al. The IAA7-ARF7-ARF19 auxin signaling module plays diverse roles in Arabidopsis growth and development. Planta, 2025, 262: 12. [30] Safi H, Saibi W, Alaoui M M, et al. A wheat lipid transfer protein (TdLTP4) promotes tolerance to abiotic and biotic stress in Arabidopsis thaliana. Plant Physiol Biochem, 2015, 89: 64–75. [31] Duan Y J, Shang X G, He Q F, et al. LIPID TRANSFER PROTEIN4 regulates cotton ceramide content and activates fiber cell elongation. Plant Physiol, 2023, 193: 1816–1833. [32] Feng H, Wang S, Dong D F, et al. Arabidopsis Ubiquitin-conjugating enzymes UBC7, UBC13, and UBC14 are required in plant responses to multiple stress conditions. Plants, 2020, 9: 723. [33] Eastmond P J, Van Dijken A J H, Spielman M, et al. Trehalose-6-phosphate synthase 1, which catalyses the first step in trehalose synthesis, is essential for Arabidopsis embryo maturation. Plant J, 2002, 29: 225–235. [34] Lu X G, Zhang F Z, Zhang C L, et al. TaTPS11 enhances wheat cold resistance by regulating source-sink factor. Plant Physiol Biochem, 2024, 211: 108695. [35] Singh V, Shah J. Tomato responds to green peach aphid infestation with the activation of trehalose metabolism and starch accumulation. Plant Signal Behav, 2012, 7: 605–607. [36] Paneque A, Fortus H, Zheng J L, et al. The hexosamine biosynthesis pathway: regulation and function. Genes, 2023, 14: 933. [37] Van Vu K, Jeong C Y, Nguyen T T, et al. Deficiency of AtGFAT1 activity impairs growth, pollen germination and tolerance to tunicamycin in Arabidopsis. J Exp Bot, 2019, 70: 1775–1787. [38] Lunn D, Gaddipati S R, Tucker G A, et al. Null mutants of individual RABA genes impact the proportion of different cell wall components in stem tissue of Arabidopsis thaliana. PLoS One, 2013, 8: e75724. |
| [1] | 张为为, 任姣姣, 魏略, 吴若彤, 陶庭余, 吴传鹏, 阿布力克木·阿布力米提, 徐晓明, 吴鹏昊. 干旱胁迫下玉米雄穗结构性状的GWAS及全基因组选择[J]. 作物学报, 2026, 52(9): 2641-2659. |
| [2] | 黄迎, 刘沛琦, 解森, 许洛, 任元, 刘旭洋, 何冠华, 李永祥, 王天宇, 黎裕, 周涛, 廖江林, 李春辉, 张登峰. 玉米氮利用效率全基因组关联分析及候选基因挖掘[J]. 作物学报, 2026, 52(9): 2558-2572. |
| [3] | 刘淳, 许忆葳, 张莹莹, 李瑞, 燕永亮, 邹华文, 王逸茹, 郑军. 耐辐射异常球菌drfE基因提高玉米的耐旱性[J]. 作物学报, 2026, 52(8): 2257-2267. |
| [4] | 金海润, 欧阳青静, 龙旭, 晏中彬, 鲁海瑞, 古开烨, 田恩堂. 甘蓝型油菜种质资源菌核病抗性鉴定与候选基因筛选[J]. 作物学报, 2026, 52(7): 2027-2044. |
| [5] | 赵辉, 黄义文, 买春艳, 景鹏飞, 孙海艳, 吴培培, 于立强, 李辉利, 周阳, 郭宪瑞, 张宏军. 小麦抗倒伏相关性状全基因组关联分析[J]. 作物学报, 2026, 52(7): 1943-1953. |
| [6] | 习千辉, 徐梓瑗, 刘梦梦, 王宏艺, 郎凯琳, 井震海, 陈锋, 赵磊. 小麦籽粒铜含量的全基因组关联分析及候选基因预测[J]. 作物学报, 2026, 52(6): 1604-1617. |
| [7] | 毛嘉琦, 黄朋雨, 赵佳佳, 郑兴卫, 武棒棒, 郝宇琼, 屈非, 刘成, 马朋涛, 郑军. 山西小麦品种白粉病抗性评价及抗病基因分子检测[J]. 作物学报, 2026, 52(6): 1669-1681. |
| [8] | 唐宽强, 李公允, 宋美毅, 赵雪, 常春玲. 大豆株高性状全基因组关联分析及预测模型构建[J]. 作物学报, 2026, 52(6): 1743-1756. |
| [9] | 杨欣雨, 崔文涛, 迪力尼格尔·阿力木, 汪凯翔, 吴鹏昊, 任姣姣. 玉米穗上叶片数全基因组关联分析和全基因组选择[J]. 作物学报, 2026, 52(5): 1573-1590. |
| [10] | 韩亚鑫, 何冠华, 张小琼, 张登峰, 李永祥, 刘旭洋, 王天宇, 黎裕, 邹华文, 李春辉. 基于RNA-Seq和BSA-Seq联合分析挖掘玉米侧根密度基因资源[J]. 作物学报, 2026, 52(5): 1341-1352. |
| [11] | 闫安, 蒋昆炜, 王蓉圆, 田林, 张璐, 王韵, 徐建龙. 水稻剑叶小维管束数基因SVN7的鉴定与克隆[J]. 作物学报, 2026, 52(5): 1364-1372. |
| [12] | 杨亚莉, 徐明睿, 马越飞, 海艺蕊, 刘凯栋, 刘万茂, 孙颖. 玉米根尖及整根响应缺铁的转录组比较研究[J]. 作物学报, 2026, 52(4): 1006-1021. |
| [13] | 张力岚, 杨军, 王让剑. 基于WGCNA发掘茶树糖苷类香气前体含量性状相关的候选基因[J]. 作物学报, 2026, 52(2): 494-513. |
| [14] | 马毅娜, 吴晓明玉, 李藕琪, 王圆, 陈丽, 张盈川, 赵伦, 文静, 傅廷栋, 沈金雄. Bna-miR1040-EIF3A模块调控油菜开花时间的功能研究[J]. 作物学报, 2026, 52(2): 349-362. |
| [15] | 鲁雅妮, 丁超杰, 张煜, 杜习军, 齐学礼, 胡琳, 许为钢. 河南省200份小麦品种苗期茎基腐病抗性鉴定与全基因组关联分析[J]. 作物学报, 2026, 52(2): 363-375. |
|