转录组和代谢组联合解析水稻分蘖期响应淹涝胁迫的分子机制

doi:10.14088/j.cnki.issn0439-8114.2025.06.036

湖北农业科学 ›› 2025, Vol. 64 ›› Issue (6): 220-231.doi: 10.14088/j.cnki.issn0439-8114.2025.06.036

转录组和代谢组联合解析水稻分蘖期响应淹涝胁迫的分子机制

罗肖郧^1,2, 李培德^1,2, 郑兴飞^1,2, 殷得所^1,2, 王红波^1,2, 胡建林^1,2, 胡鹏^1,2, 刘丹^1,2, 文艺^1,2, 陈东攀^1,2, 雷添杰³, 徐得泽^1,2

1.湖北省农业科学院粮食作物研究所/粮食作物种质创新与遗传改良湖北省重点实验室/农业农村部作物分子育种重点实验室, 武汉 430064;
2.湖北洪山实验室,武汉 430070;
3.中国农业科学院农业环境与可持续发展研究所,北京 100081

收稿日期:2025-04-29 出版日期:2025-06-25 发布日期:2025-07-18
通讯作者: 徐得泽（1978-）,男,安徽庐江人,研究员,主要从事高档优质稻与专用稻育种研究,（电子信箱）dezexu@163.com。
作者简介:罗肖郧（1990-）,男,湖北十堰人,助理研究员,主要从事水稻遗传育种研究,（电话）18827046869（电子信箱）luoxy0916@hbaas.com;共同第一作者,李培德（1981-）,男,安徽亳州人,副研究员,主要从事水稻遗传育种及栽培研究,（电话）13477038996（电子信箱）lipeide2003@163.com
基金资助:
国家重点研发计划项目（2023YFD2300300）; 湖北省自然科学基金青年项目（2024AFB406）; 湖北省农业科学院青年科学基金项目（2024NKYJJ04）; 粮食作物种质创新与遗传改良湖北省重点实验室开放课题（2023lzjj03; 2022lzjj08; 2020lzjj05）

Research on molecular mechanisms of Oryza sativa L. response to waterlogging stress during tillering stage using integrated transcriptome-metabolome analysis

LUO Xiao-yun^1,2, LI Pei-de^1,2, ZHENG Xing-fei^1,2, YIN De-suo^1,2, WANG Hong-bo^1,2, HU Jian-lin^1,2, HU Peng^1,2, LIU Dan^1,2, WEN Yi^1,2, CHEN Dong-pan^1,2, LEI Tian-jie³, XU De-ze^1,2

1. Institute of Food Crops/Hubei Key Laboratory of Food Crop Germplasm and Genetic Improvement/Key Laboratory of Crop Molecular Breeding,Ministry of Agriculture and Rural Affairs,Hubei Academy of Agricultural Sciences,Wuhan 430064,China;
2. Hubei Hongshan Laboratory,Wuhan 430070,China;
3. Institute of Environment and Sustainable Development in Agriculture, Chinese Academy of Agricultural Sciences, Beijing 100081,China

Received:2025-04-29 Published:2025-06-25 Online:2025-07-18

摘要/Abstract

摘要： 以水稻（Oryza sativa L.）品种籼稻鄂中6号和粳稻长粒梗384为材料,在分蘖期进行2/3株高水深淹水处理(持续7 d),通过转录组和代谢组分析胁迫后的基因表达和代谢物变化。转录组分析显示,鄂中6号和长粒梗384中分别鉴定出3 080和1 624个差异表达基因（DEGs）,显著富集于碳水化合物生物合成过程、活性氧代谢过程、植物与病原体相互作用及植物激素信号转导等通路。代谢组分析发现,鄂中6号和长粒梗384中苯及其衍生物、有机酸和内酯类化合物等差异代谢物（DAMs）显著富集,涉及二萜类生物合成和类黄酮生物合成等途径。转录组和代谢组联合分析发现,鄂中6号和长粒梗384中有7条共同通路显著富集,分别为代谢途径、次级代谢产物的生物合成、多种植物次级代谢产物的生物合成、二萜类生物合成、亚油酸代谢、苯丙烷类生物合成、类胡萝卜素生物合成。水稻通过激活抗氧化系统、调控激素信号及积累防御性代谢物应对淹涝胁迫。

关键词: 转录组, 代谢组, 淹涝胁迫, 水稻（Oryza sativa L.）, 分蘖期, 分子机制

Abstract: Using Oryza sativa L. cultivars Ezhong 6 (indica) and Changligeng 384 (japonica) as materials, plants at the tillering stage were subjected to 2/3 plant height water depth treatment (lasting 7 days). Transcriptome and metabolome analyses were performed to investigate post-stress gene expression and metabolite changes.Transcriptome analysis identified 3 080 and 1 624 differentially expressed genes (DEGs) in Ezhong 6 and Changligeng 384, respectively, which were significantly enriched in carbohydrate biosynthetic process, reactive oxygen species metabolic process, plant-pathogen interaction, and plant hormone signal transduction pathways.Metabolome analysis revealed significant enrichment of differentially accumulated metabolites (DAMs) such as benzene and substituted derivatives, organic acids, and lactones in Ezhong 6 and Changligeng 384, primarily involved in diterpenoid biosynthesis and flavonoid biosynthesis pathways.Integrated transcriptome-metabolome analysis identified seven co-enriched pathways in both cultivars: Metabolic pathways, biosynthesis of secondary metabolites, biosynthesis of various plant secondary metabolites, diterpenoid biosynthesis, linoleic acid metabolism, phenylpropanoid biosynthesis, and carotenoid biosynthesis.Oryza sativa L. responded to waterlogging stress by activating antioxidant systems, regulating hormone signaling, and accumulating defensive metabolites.

Key words: transcriptome, metabolome, waterlogging stress, rice (Oryza sativa L.), tillering stage, molecular mechanisms

中图分类号:

S332.4

罗肖郧, 李培德, 郑兴飞, 殷得所, 王红波, 胡建林, 胡鹏, 刘丹, 文艺, 陈东攀, 雷添杰, 徐得泽. 转录组和代谢组联合解析水稻分蘖期响应淹涝胁迫的分子机制[J]. 湖北农业科学, 2025, 64(6): 220-231.

LUO Xiao-yun, LI Pei-de, ZHENG Xing-fei, YIN De-suo, WANG Hong-bo, HU Jian-lin, HU Peng, LIU Dan, WEN Yi, CHEN Dong-pan, LEI Tian-jie, XU De-ze. Research on molecular mechanisms of Oryza sativa L. response to waterlogging stress during tillering stage using integrated transcriptome-metabolome analysis[J]. HUBEI AGRICULTURAL SCIENCES, 2025, 64(6): 220-231.

参考文献

[1] GREGORY P J, GEORGE T S.Feeding nine billion: The challenge to sustainable crop production[J]. Journal of experimental botany, 2011, 62(15): 5233-5239.
[2] GODFRAY H C J, BEDDINGTON J R, CRUTE I R, et al. Food security: The challenge of feeding 9 billion people[J]. Science, 2010, 327: 812-818.
[3] 李茂松, 李森, 李育慧. 中国近50年洪涝灾害灾情分析[J]. 中国农业气象, 2004(1): 40-43.
[4] 周建林, 周广洽, 陈良碧. 洪涝对水稻的危害及其抗灾减灾的栽培措施[J]. 自然灾害学报, 2001(1): 103-106.
[5] VOESENEK L A C J, BAILEY-SERRES J. Flood adaptive traits and processes: An overview[J]. New phytologist, 2015, 206(1): 57-73.
[6] FUKAO T, XIONG L.Genetic mechanisms conferring adaptation to submergence and drought in rice: Simple or complex?[J]. Current opinion in plant biology, 2013, 16(2): 196-204.
[7] HATTORI Y, NAGAI K, ASHIKARI M.Rice growth adapting to deepwater[J]. Current opinion in plant biology, 2011, 14(1): 100-105.
[8] KATO Y, COLLARD B C Y, SEPTININGSIH E M, et al. Physiological analyses of traits associated with tolerance of long-term partial submergence in rice[J]. AoB plants, 2014, 6: plu058.
[9] LORETI E, VAN VEEN H, PERATA P.Plant responses to flooding stress[J]. Current opinion in plant biology, 2016, 33: 64-71.
[10] ZHOU W, CHEN F, MENG Y, et al.Plant waterlogging/flooding stress responses: From seed germination to maturation[J]. Plant physiology and biochemistry, 2020, 148: 228-236.
[11] 聂功平,陈敏敏,杨柳燕,等.植物响应淹水胁迫的研究进展[J]. 中国农学通报, 2021, 37(18): 57-64.
[12] ISMAIL A M, SINGH U S, SINGH S, et al.The contribution of submergence-tolerant (Sub1) rice varieties to food security in flood-prone rainfed lowland areas in Asia[J]. Field crops research, 2013, 152: 83-93.
[13] 杨建莹, 霍治国, 吴立,等. 西南地区水稻洪涝等级评价指标构建及风险分析[J]. 农业工程学报, 2015, 31(16): 135-144.
[14] 张林. 水稻涝害的危害症状及防治措施[J]. 农业灾害研究, 2018, 8(2): 54-55,65.
[15] 杨龙树, 张从合, 严志,等. 长江流域水稻涝害的发生及应对措施[J]. 农业灾害研究, 2020, 10(4): 58-59,78.
[16] 宣守丽, 石春林, 张建华,等. 分蘖期淹水胁迫对水稻地上部物质分配及产量构成的影响[J]. 江苏农业学报, 2013, 29(6): 1199-1204.
[17] 邵长秀,潘学标,李家文,等. 不同生育阶段洪涝淹没时长对水稻生长发育及产量构成的影响[J]. 农业工程学报,2019,35(3):125-133.
[18] VELCULESCU V E, ZHANG L, ZHOU W, et al.Characterization of the yeast transcriptome[J]. Cell, 1997, 88(2): 243-251.
[19] COSTA V, ANGELINI C, DE FEIS I, et al.Uncovering the complexity of transcriptomes with RNA-Seq[J]. Journal of biomedicine and biotechnology, 2010,1: 853916.
[20] FIEHN O, KOPKA J, DÖRMANN P, et al. Metabolite profiling for plant functional genomics[J]. Nature biotechnology, 2000, 18(11): 1157-1161.
[21] FIEHN O.Combining genomics, metabolome analysis, and biochemical modelling to understand metabolic networks[J]. Comparative and functional genomics, 2001, 2(3): 155-168.
[22] NICHOLSON J K, LINDON J C, HOLMES E.Metabonomics: Understanding the metabolic responses of living systems to pathophysiological stimuli via multivariate statistical analysis of biological NMR spectroscopic data[J]. Xenobiotica, 2008, 29(11): 1181-1189.
[23] FIEHN O.Metabolomics-the link between genotypes and phenotypes[J]. Functional genomics, 2002, 48: 155-171.
[24] GUIJAS C, MONTENEGRO-BURKE J R, WARTH B, et al. Metabolomics activity screening for identifying metabolites that modulate phenotype[J]. Nature biotechnology, 2018, 36(4): 316-320.
[25] 李洁, 姚晓华. 多组学关联分析作物耐逆境胁迫研究进展[J]. 广东农业科学, 2019, 46(8): 22-28.
[26] 季元, 于冰, 陈偲学. 多组学技术在植物应答非生物胁迫中的研究进展[J]. 中国农学通报, 2023, 39(23): 1-7.
[27] 薛守宇, 朱涛, 李冰冰,等. 转录组和代谢组联合分析在植物中的应用研究[J]. 山西农业大学学报(自然科学版), 2022, 42(3): 1-13.
[28] YAMAKAWA H, HAKATA M.Atlas of rice grain filling-related metabolism under high temperature: joint analysis of metabolome and transcriptome demonstrated inhibition of starch accumulation and induction of amino acid accumulation[J]. Plant and cell physiology, 2010, 51(5): 795-809.
[29] ZHANG J, LUO W, ZHAO Y, et al.Comparative metabolomic analysis reveals a reactive oxygen species‐dominated dynamic model underlying chilling environment adaptation and tolerance in rice[J]. New phytologist, 2016, 211(4): 1295-1310.
[30] SHU L, LOU Q, MA C, et al.Genetic, proteomic and metabolic analysis of the regulation of energy storage in rice seedlings in response to drought[J]. Proteomics, 2011, 11(21): 4122-4138.
[31] DUBEY S, MISRA P, DWIVEDI S, et al.Transcriptomic and metabolomic shifts in rice roots in response to Cr(VI) stress[J]. BMC genomics, 2010, 11: 1-19.
[32] WANG W S, ZHAO X Q, LI M, et al.Complex molecular mechanisms underlying seedling salt tolerance in rice revealed by comparative transcriptome and metabolomic profiling[J]. Journal of experimental botany, 2016, 67(1): 405-419.
[33] WANG L,STEGEMANN J P.Extraction of high quality RNA from polysaccharide matrices using cetlytrimethylammonium bromide[J]. Biomaterials, 2010, 31(7): 1612-1618.
[34] CHEN S, ZHOU Y, CHEN Y, et al.Fastp: An ultra-fast all-in-one FASTQ preprocessor[J]. Bioinformatics, 2018, 34(17): i884-i890.
[35] KIM D, LANGMEAD B, SALZBERG S L.HISAT: A fast spliced aligner with low memory requirements[J]. Nature methods, 2015, 12(4): 357-360.
[36] LOVE M I, HUBER W, ANDERS S.Moderated estimation of fold change and dispersion for RNA-seq data with DESeq2[J]. Genome biology, 2014, 15(12): 1-21.
[37] VARET H, BRILLET-GUÉGUEN L, COPPÉE J-Y, et al. SARTools: A DESeq2- and EdgeR-Based R pipeline for comprehensive differential analysis of RNA-Seq data[J]. PLoS one, 2016, 11(6): e0157022.
[38] ASHBURNER M, BALL C A, BLAKE J A, et al.Gene ontology: Tool for the unification of biology[J]. Nature genetics,2000,25(1):25-29.
[39] KANEHISA M, ARAKI M. GOTO S, et al.KEGG for linking genomes to life and the environment[J]. Nucleic acids research, 2007, 36: 480-484.
[40] KANEHISA M, GOTO S.KEGG: Kyoto encyclopedia of genes and genomes[J]. Nucleic acids research, 2000, 28(1): 27-30.
[41] SAUTER M.Rice in deep water: How to take heed against a sea of troubles[J]. Naturwissenschaften, 2000, 87(7): 289-303.
[42] PERATA P, VOESENEK L A C J. Submergence tolerance in rice requires Sub1A,an ethylene-response-factor-like gene[J]. Trends in plant science, 2007, 12(2): 43-46.
[43] PANDA D, SHARMA S G, SARKAR R K.Chlorophyll fluorescence parameters, CO₂ photosynthetic rate and regeneration capacity as a result of complete submergence and subsequent re-emergence in rice (Oryza sativa L.)[J]. Aquatic botany, 2008,88(2): 127-133.
[44] HATTORI Y, NAGAI K, FURUKAWA S, et al.The ethylene response factors SNORKEL1 and SNORKEL2 allow rice to adapt to deep water[J]. Nature, 2009, 460: 1026-1030.
[45] ADAK M K, GHOSH N, DASGUPTA D K, et al.Impeded carbohydrate metabolism in rice plants under submergence stress[J]. Rice science, 2011, 18(2): 116-126.
[46] PUCCIARIELLO C, PERATA P.Quiescence in rice submergence tolerance: An evolutionary hypothesis[J]. Trends in plant science, 2013, 18(7): 377-381.
[47] DAS K K, SARKAR R K, ISMAIL A M.Elongation ability and non-structural carbohydrate levels in relation to submergence tolerance in rice[J]. Plant science, 2005, 168(1): 131-136.
[48] PANDA D, SARKAR R K.Non-structural carbohydrate metabolism associated with submergence tolerance in rice[J]. Genetics and plant physiology, 2011, 1(3-4): 155-162.
[49] BLOKHINA O, VIROLAINEN E, FAGERSTEDT K V.Antioxidants, oxidative damage and oxygen deprivation stress: A review[J]. Annals of botany, 2003, 91(2): 179-194.
[50] MITTLER R,VANDERAUWERA S,GOLLERY M, et al.Reactive oxygen gene network of plants[J]. Trends in plant science, 2004, 9(10): 490-498.
[51] SANTOSA I E, RAM P C, BOAMFA E I, et al.Patterns of peroxidative ethane emission from submerged rice seedlings indicate that damage from reactive oxygen species takes place during submergence and is not necessarily a post-anoxic phenomenon[J]. Planta, 2007, 226(1): 193-202.
[52] USHIMARO T, SHIBASAKA M, TSUJI H.Development of O₂--detoxification system during adaptation to air of submerged rice seedlings[J]. Plant and cell physiology, 1992, 33(8): 1065-1071.
[53] ALSCHER R G, ERTURK N, HEATH L S.Role of superoxide dismutases (SODs) in controlling oxidative stress in plants[J]. Journal of experimental botany, 2002, 53(372): 1331-1341.
[54] ELLA E S, KAWANO N, ITO O.Importance of active oxygen-scavenging system in the recovery of rice seedlings after submergence[J]. Plant science, 2003, 165(1): 85-93.
[55] 夏石头, 彭克勤, 曾可. 水稻涝害生理及其与水稻生产的关系[J]. 植物生理学通讯, 2000, 36(6): 581-588.
[56] FUKAO T, BAILEY-SERRES J.Ethylene-a key regulator of submergence responses in rice[J]. Plant science, 2008, 175(1-2): 43-51.
[57] SCHMITZ A J, FOLSOM J J, JIKAMARU Y, et al.SUB1A-mediated submergence tolerance response in rice involves differential regulation of the brassinosteroid pathway[J]. New phytologist, 2013, 198(4): 1060-1070.
[58] AYANO M, KANI T, KOJIMA M, et al.Gibberellin biosynthesis and signal transduction is essential for internode elongation in deepwater rice[J]. Plant, cell & environment, 2014, 37(10): 2313-2324.
[59] 张凤,陈伟. 代谢组学在植物逆境生物学中的研究进展[J]. 生物技术通报, 2021, 37(8): 1-11.
[60] UPCHURCH R G.Fatty acid unsaturation, mobilization, and regulation in the response of plants to stress[J]. Biotechnology letters, 2008, 30(6): 967-977.
[61] KACHROO A, KACHROO P.Fatty acid-derived signals in plant defense[J]. Annual review of phytopathology, 2009, 47: 153-176.
[62] 王玉, 杨雪, 杨蕊菁,等. 调控苯丙烷类生物合成的MYB类转录因子研究进展[J]. 安徽农业大学学报, 2019, 46(5): 859-864.
[63] 葛文佳, 辛建攀, 田如男. 植物苯丙烷代谢及其对重金属胁迫的响应研究进展[J]. 生物工程学报, 2023, 39(2): 425-445.
[64] 汪胜勇,陈宇航,陈会丽,等. 水稻减数分裂期高温对苯丙烷类代谢及下游分支代谢途径的影响[J]. 中国水稻科学,2023,37(4):368-378.
[65] 谢凤, 郝乐, 王振杰,等. 苯丙烷代谢途径分支对生物胁迫响应的研究进展[J]. 中国植保导刊, 2023, 43(2): 23-30.
[66] 尚军, 吴旺泽, 马永贵. 植物苯丙烷代谢途径[J]. 中国生物化学与分子生物学报, 2022, 38(11): 1467-1476.
[67] CARDOSO A A, GORI A, DA-SILVA C J, et al. Abscisic acid biosynthesis and signaling in plants: Key targets to improve water use efficiency and drought tolerance[J]. Applied sciences, 2020, 10(18): 6322.
[68] CHEN K, LI G J, BRESSAN R A, et al.Abscisic acid dynamics, signaling, and functions in plants[J]. Journal of integrative plant biology, 2020, 62(1): 25-54.
[69] BROOKBANK B P, PATEL J, GAZZARRINI S, et al.Role of basal ABA in plant growth and development[J]. Genes (Basel), 2021, 12(12): 1936.
[70] KUMAR S, SHAH S H, VIMALA Y, et al.Abscisic acid: Metabolism, transport, crosstalk with other plant growth regulators, and its role in heavy metal stress mitigation[J]. Frontiers in plant science, 2022, 13: 972856.
[71] LIU H, SONG S, ZHANG H, et al.Signaling transduction of ABA, ROS, and Ca²+ in plant stomatal closure in response to drought[J]. International journal of molecular sciences, 2022, 23(23): 142824.
[72] 刘晓龙, 徐晨, 邵勤,等. 脱落酸提高水稻抗逆性的研究进展[J]. 东北农业科学, 2022, 47(6): 29-33.
[73] RAMA KRISHNAYYA G, SETTER T L, SARKAR R K, et al.Influence of phosphorus application to floodwater on oxygen concentrations and survival of rice during complete submergence[J]. Experimental agriculture, 1999, 35(2): 167-180.
[74] GAUTAM P, NAYAK A K, LAL B, et al.Submergence tolerance in relation to application time of nitrogen and phosphorus in rice (Oryza sativa L.)[J]. Environmental and experimental botany, 2014, 99: 159-166.

转录组和代谢组联合解析水稻分蘖期响应淹涝胁迫的分子机制

Research on molecular mechanisms of Oryza sativa L. response to waterlogging stress during tillering stage using integrated transcriptome-metabolome analysis

PDF

可视化

摘要/Abstract

引用本文

使用本文

参考文献

相关文章 15

编辑推荐

Metrics

本文评价

[1]	孟刚, 王瑞娴, 楚渠, 彭云武, 杨金宏, 陈安利, 张笙源, 凌君. 野桑蚕全长转录组测序及生物信息学分析[J]. 湖北农业科学, 2025, 64(6): 197-206.
[2]	刘芬, 屈成, 洪超怡, 肖建平, 朱世军, 郝明. 水稻硅突变体的研究进展[J]. 湖北农业科学, 2025, 64(5): 5-9.
[3]	张慧, 王谊, 杭晓宁, 张健, 廖敦秀, 唐荣莉. 受旱对水稻和土壤镉含量及镉抗性相关微生物的影响[J]. 湖北农业科学, 2025, 64(5): 10-16.
[4]	朱思艺, 邓龙君, 李天才, 张扬, 郭勇垚, 罗伟, 杜宗君. 金沙鲈鲤全长转录组测序分析与抗菌肽基因的鉴定[J]. 湖北农业科学, 2025, 64(3): 167-175.
[5]	葛杰, 何玉池, 展帆, 李之龙. 多倍体水稻DHF-CBS基因的抗旱性鉴定[J]. 湖北农业科学, 2024, 63(9): 216-220.
[6]	康兵, 顾祝禹, 皮杰, 黄博阳, 李文超, 唐东海. 硅基叶面阻控剂对水稻的控镉效应[J]. 湖北农业科学, 2024, 63(6): 22-26.
[7]	张应胡, 秦绪光, 马丽, 王皓杰, 胡玲, 田光明. 基于非靶向代谢组学挖掘两种日粮模式下蛋鸡回肠代谢物的差异[J]. 湖北农业科学, 2024, 63(6): 213-217.
[8]	毛楠楠, 孙勇胜, 陈辉, 周荣艳, 籍颖. 不同产蛋数岩鸽卵巢转录组的SNP分析[J]. 湖北农业科学, 2024, 63(4): 185-190.
[9]	李鑫奎, 罗学刚. 纳米二氧化硅对水稻种子萌发及分蘖期生长和光合特性的影响[J]. 湖北农业科学, 2024, 63(11): 6-12.
[10]	赵源, 李秋剑, 张力, 王卫民, 郭进, 芦伟龙, 许嘉阳, 贾玮, 韩丹, 史久长, 许自成. 四川凉山烟区不同烤烟品种的适应性研究[J]. 湖北农业科学, 2024, 63(11): 100-107.
[11]	郭晓亮, 穆森, 段媛媛, 胡昌强, 黄浩. 川牛膝片产地趁鲜加工及其可行性[J]. 湖北农业科学, 2024, 63(11): 114-121.
[12]	郝玲, 杨莹, 张佩, 邱航, 钱沈旸. 基于大气环流特征量和海温的水稻产量结构因素预报模型研究——以江苏省为例[J]. 湖北农业科学, 2023, 62(8): 27-36.
[13]	肖婉钰, 孙艺嘉, 周贤玉, 任海龙, 张晶, 黄江华, 许东林. 水稻发芽率测定中污染性微生物的鉴定及多样性[J]. 湖北农业科学, 2023, 62(7): 51-56.
[14]	李兴华, 陈展鹏, 蔡正军, 邹彩琼, 张中南, 张文超, 丁凤菊. 稻田套养不同密度黑斑蛙对水稻产量和土壤肥力的影响初探[J]. 湖北农业科学, 2023, 62(5): 5-7.
[15]	毛可欣, 王海荣, 安淼, 吕巍, 李健, 李国田. 猕猴桃‘脐红'与‘徐香'越冬期转录组的比较分析[J]. 湖北农业科学, 2023, 62(5): 165-171.