[1]张中起,代名扬,李昆仑,等.大豆TOR家族全基因组鉴定及表达特征分析[J].江苏农业学报,2025,(10):1886-1898.[doi:doi:10.3969/j.issn.1000-4440.2025.10.002]
　ZHANG Zhongqi,DAI Mingyang,LI Kunlun,et al.Genome-wide identification and expression characteristics analysis of soybean TOR family[J].,2025,(10):1886-1898.[doi:doi:10.3969/j.issn.1000-4440.2025.10.002]
点击复制

大豆TOR家族全基因组鉴定及表达特征分析()

江苏农业学报[ISSN:1006-6977/CN:61-1281/TN]

卷:
期数:
2025年10期
页码:
1886-1898
栏目:
遗传育种·生理生化
出版日期:
2025-10-31

文章信息/Info

Title:
Genome-wide identification and expression characteristics analysis of soybean TOR family
作者:
张中起1; 代名扬2; 李昆仑1; 吕令华1; 刘磊1; 曹嵩晓3; 李品1; 张震彪4; 薛墨庸4
(1.菏泽市农业科学院,山东菏泽274000；2.江苏中烟工业有限责任公司技术中心,江苏南京210019；3.日照市东港区农业农村局,山东日照276800；4.中国农业科学院烟草研究所,山东青岛266101)
Author(s):
ZHANG Zhongqi1; DAI Mingyang2; LI Kunlun1; LYU Linghua1; LIU Lei1; CAO Songxiao3; LI Pin1; ZHANG Zhenbiao4; XUE Moyong4
(1.Heze Academy of Agricultural Sciences, Heze 274000, China;2.Technology Center, China Tobacco Jiangsu Industrial Co., Ltd., Nanjing 210019, China;3.Donggang District Bureau of Agriculture and Rural Affairs, Rizhao 276800, China;4.Tobacco Research Institute, Chinese Academy of Agricultural Sciences, Qingdao 266101, China)
关键词:
大豆; GmTOR1; GmTOR2; 生物信息学; 进化; 表达特征
Keywords:
soybean; GmTOR1; GmTOR2; bioinformatics; evolution; expression characteristics
分类号:
S565.1
DOI:
doi:10.3969/j.issn.1000-4440.2025.10.002
文献标志码:
A
摘要:
雷帕霉素靶(TOR)蛋白是植物生长、发育和应激反应中的主要调节因子。目前对植物TOR基因的研究仍十分有限,且在重要农作物大豆中的相关研究尤为缺乏。本研究利用生物信息学方法鉴定了大豆TOR基因家族,分析了其蛋白质理化性质、基因结构、蛋白质二级结构、磷酸化位点、染色体位置、系统发育关系、进化压力、启动子元件、蛋白质互作网络和表达模式等。结果表明,GmTOR1和GmTOR2蛋白序列长度均为2 468 aa,二级结构主要为α螺旋和无规则卷曲,分别有193个、191个潜力较高的磷酸化位点。GmTOR1和GmTOR2分别位于1号和11号染色体上,启动子区存在脱落酸(ABA)、茉莉酸甲酯(MeJA)、乙烯(ETH)、赤霉素(GA)和水杨酸(SA)等响应元件,表明GmTOR1和GmTOR2可能参与激素调控、干旱胁迫、盐胁迫、低温胁迫、大豆疫霉菌侵染等响应过程。转录组数据分析结果表明,GmTOR1和GmTOR2在茎尖、花、侧芽和豆荚中优势表达。GmTOR1和GmTOR2受淹水胁迫后在根中的表达先下降后上升,而在叶中的表达模式截然相反。GmTOR1和GmTOR2受低温胁迫、大豆疫霉菌侵染和乙烯诱导后下调表达。qRT-PCR验证结果表明,GmTOR1、GmTOR2基因在盐胁迫下的表达模式与转录组测序结果基本一致。本研究系统分析了GmTOR1和GmTOR2的序列特征、进化关系及表达模式,丰富了对大豆TOR基因家族的认知,并为它们的后续功能研究奠定了坚实的理论基础。
Abstract:
The target of rapamycin (TOR) protein serves as a central regulator in plant growth, development, and stress responses. Current research on plant TOR genes remains limited, with particularly scarce investigations in important crops like soybean. This study employed bioinformatics approaches to identify the soybean TOR gene family and analyzed various characteristics including protein physicochemical properties, gene structure, protein secondary structure, phosphorylation sites, chromosomal localization, phylogenetic relationships, evolutionary constraints, promoter elements, protein interaction networks, and expression patterns. Results revealed that both GmTOR1 and GmTOR2 proteins contain 2 468 amino acids, with secondary structures predominantly composed of α-helices and random coils, exhibiting 193 and 191 high-potential phosphorylation sites, respectively. GmTOR1 and GmTOR2 are located on chromosomes 1 and 11, respectively. Promoter regions contain response elements for abscisic acid (ABA), methyl jasmonate (MeJA), ethylene (ETH), gibberellin (GA), and salicylic acid (SA), suggesting GmTOR1 and GmTOR2 may participate in hormone regulation and responses to drought stress, salt stress, cold stress, and Phytophthora sojae infection. Transcriptomic data analysis demonstrated that GmTOR1 and GmTOR2 exhibited predominant expression in shoot apices, flowers, lateral buds, and pods. Under flooding stress, their expression in roots decreased initially and then increased, whereas the opposite pattern was observed in the leaves. GmTOR1 and GmTOR2 were down-regulated in response to cold stress, Phytophthora sojae infection, and ethylene induction. qRT-PCR validation results indicated that the expression patterns of GmTOR1 and GmTOR2 under salt stress were largely consistent with the transcriptome sequencing results. Here, we systematically analyzed the sequence characteristics, evolutionary relationships, and expression patterns of GmTOR1 and GmTOR2. These findings enrich our understanding of the soybean TOR gene family and provide a solid theoretical foundation for their subsequent functional studies.

参考文献/References:

[1]万志前,刘政,俞秦峰. 我国大豆新品种保护的现状分析与思考[J]. 江苏农业科学,2024,52(11):35-44.
[2]鲍洁,张小允,许世卫. 我国大豆消费影响因素分析及趋势预测[J]. 江苏农业科学,2023,51(8):240-248.
[3]李艺阳,吴冕,顾和平,等. 江苏省大豆豆荚真菌病害的病原菌分离与鉴定[J]. 江苏农业学报,2024,40(10):1801-1809.
[4]曹帅,杜仲阳,刘鹏,等. 碱胁迫对大豆光合特性及内源激素含量的影响[J]. 江苏农业学报,2020,36(2):284-291.
[5]CALDANA C, MARTINS M C M, MUBEEN U, et al. The magic ‘hammer’ of TOR:the multiple faces of a single pathway in the metabolic regulation of plant growth and development[J]. Journal of Experimental Botany,2019,70(8):2217-2225.
[6]QUILICHINI T D, GAO P, PANDEY P K, et al. A role for TOR signaling at every stage of plant life[J]. Journal of Experimental Botany,2019,70(8):2285-2296.
[7]SHI L, WU Y, SHEEN J. TOR signaling in plants:conservation and innovation[J]. Development,2018,145(13):dev160887.
[8]BURKART G M, BRANDIZZI F. A tour of TOR complex signaling in plants[J]. Trends in Biochemical Sciences,2021,46(5):417-428.
[9]ZHANG N, MENG Y Y, LI X, et al. Metabolite-mediated TOR signaling regulates the circadian clock in Arabidopsis[J]. Proceedings of the National Academy of Sciences of the United States of America,2019,116(51):25395-25397.
[10]YE R Q, WANG M Y, DU H, et al. Glucose-driven TOR-FIE-PRC2 signalling controls plant development[J]. Nature,2022,609(7929):986-993.
[11]王臻,吴春霞,扈玉婷,等. 盐芥TOR基因的克隆及对盐芥生长发育的影响[J]. 华南理工大学学报(自然科学版),2011,39(5):143-148.
[12]AGREDANO-MORENO L T, REYES DE LA CRUZ H, MARTNEZ-CASTILLA L P, et al. Distinctive expression and functional regulation of the maize (Zea mays L. ) TOR kinase ortholog[J]. Molecular BioSystems,2007,3(11):794-802.
[13]孙林啸. 水稻核糖体蛋白S6激酶受TOR/RAPTOR2介导调控类囊体膜形成和植物生长[D]. 武汉:华中农业大学,2016.
[14]FU L W, LIU Y L, QIN G C, et al. The TOR-EIN2 axis mediates nuclear signalling to modulate plant growth[J]. Nature,2021,591(7849):288-292.
[15]WANG Z Z, LI H, WENG Y X. OsFKBP12 transduces the sucrose signal from OsNIN8 to the OsTOR pathway in a loosely binding manner for cell division[J]. iScience,2025,28(1):111555.
[16]LIU Y L, HU J, DUAN X L, et al. Target of rapamycin (TOR):a master regulator in plant growth,development,and stress responses[J]. Annual Review of Plant Biology,2025,76(1):341-371.
[17]LI K L, XUE H, TANG R J, et al. TORC pathway intersects with a calcium sensor kinase network to regulate potassium sensing in Arabidopsis[J]. Proceedings of the National Academy of Sciences of the United States of America,2023,120(47):e2316011120.
[18]李丹阳. 雷帕霉素靶蛋白通过自噬和茉莉酸途径调节平邑甜茶对低氮胁迫的反应[D]. 杨凌:西北农林科技大学,2022.
[19]ZHUO F P, XIONG F J, DENG K X, et al. Target of rapamycin (TOR) negatively regulates ethylene signals in Arabidopsis[J]. International Journal of Molecular Sciences,2020,21(8):2680.
[20]邹佩锦. 扰动TOR信号途径对丹参生长及其主要活性物质生物合成途径的影响[D]. 成都:成都中医药大学,2023.
[21]杨惠. 转PnTOR基因烟草的黑胫病抗性研究[D]. 重庆:西南大学,2020.
[22]赵珺玥. CsTCTPs介导TOR信号调控黄瓜抗病性的研究[D]. 沈阳:沈阳农业大学,2019.
[23]张彤彤. TOR抑制剂雷帕霉素对番茄幼苗生长发育及低温响应的影响[D]. 沈阳:沈阳农业大学,2019.
[24]孙丽园. 干旱胁迫下TOR与JA信号相互作用对棉苗生长的影响[D]. 南京:南京农业大学,2020.
[25]CHEN C J, WU Y, LI J W, et al. TBtools-II:a “one for all,all for one” bioinformatics platform for biological big-data mining[J]. Molecular Plant,2023,16(11):1733-1742.
[26]ZHANG Z. KaKs_Calculator 3. 0:calculating selective pressure on coding and non-coding sequences[J]. Genomics,Proteomics & Bioinformatics,2022,20(3):536-540.
[27]TAMURA K, STECHER G, KUMAR S. MEGA11:molecular evolutionary genetics analysis version 11[J]. Molecular Biology and Evolution,2021,38(7):3022-3027.
[28]陈文臻,刘佳琦,都浩. 植物雷帕霉素靶蛋白激酶研究进展[J]. 浙江大学学报(农业与生命科学版),2023,49(5):591-606.
[29]赵珺玥,范海延,崔娜,等. 植物雷帕霉素靶蛋白(TOR)信号通路研究进展[J]. 植物生理学报,2018,54(4):549-556.
[30]MENAND B, DESNOS T, NUSSAUME L, et al. Expression and disruption of the Arabidopsis TOR (target of rapamycin) gene[J]. Proceedings of the National Academy of Sciences of the United States of America,2002,99(9):6422-6427.
[31]李亚超,刘静,周梦岩,等. 植物响应非生物胁迫的磷酸化修饰组学研究进展[J]. 西北植物学报,2020,40(8):1436-1446.
[32]宋想,王璐瑶,富博晓,等. 高等植物启动子元件鉴定与合成研究进展[J]. 植物学报,2024,59(5):691-708.
[33]AGARWAL M, HAO Y J, KAPOOR A, et al. A R2R3 type MYB transcription factor is involved in the cold regulation of CBF genes and in acquired freezing tolerance[J]. Journal of Biological Chemistry,2006,281(49):37636-37645.
[34]NARUSAKA Y, NAKASHIMA K, SHINWARI Z K, et al. Interaction between two cis-acting elements,ABRE and DRE in ABA-dependent expression of Arabidopsis rd29A gene in response to dehydration and high-salinity stresses[J]. The Plant Journal,2003,34(2):137-148.
[35]DOLFERUS R, JACOBS M, PEACOCK W J, et al. Differential interactions of promoter elements in stress responses of the Arabidopsis Adh gene[J]. Plant Physiology,1994,105(4):1075-1087.
[36]张钰蛟. SlMYC2-TOR介导JA信号调控番茄生长发育的机制[D]. 沈阳:沈阳农业大学,2023.
[37]DANECEK P, AUTON A, ABECASIS G, et al. The variant call format and VCFtools[J]. Bioinformatics,2011,27(15):2156-2158.
[38]KORNELIUSSEN T S, MOLTKE I, ALBRECHTSEN A, et al. Calculation of Tajima’s D and other neutrality test statistics from low depth next-generation sequencing data[J]. BMC Bioinformatics,2013,14:289.
[39]XU K, ZHAO Y, ZHAO S H, et al. Genome-wide identification and low temperature responsive pattern of actin depolymerizing factor (ADF) gene family in wheat (Triticum aestivum L.)[J]. Frontiers in Plant Science,2021,12:618984.
[40]CAPITAO C, SHUKLA N, WANDROLOVA A, et al. Functional characterization of SMG7 paralogs in Arabidopsis thaliana[J]. Frontiers in Plant Science,2018,9:1602.

相似文献/References:

[1]刘朝茂,李成云.玉米与大豆间作对玉米叶片衰老的影响[J].江苏农业学报,2017,(02):322.[doi:doi:10.3969/j.issn.1000-4440.2017.02.013]
　LIU Chao-mao,LI Cheng-yun.Effects of maize/soybean intercropping on maize leaf senescence[J].,2017,(10):322.[doi:doi:10.3969/j.issn.1000-4440.2017.02.013]
[2]张令瑄,谢婷婷,王瑾,等.大田条件下ＵＶ-Ｂ 辐射增强对大豆根际土壤相关指标的影响[J].江苏农业学报,2016,(01):118.[doi:10.3969/j.issn.1000-4440.2016.01.018]
　ZHANG Ling-xuan,XIE Ting-ting,WANG Jin,et al.Soybean rhizosphere soil parameters in response to enhanced UV-B radiation under field condition[J].,2016,(10):118.[doi:10.3969/j.issn.1000-4440.2016.01.018]
[3]宁丽华,何晓兰,张大勇.大豆耐盐相关基因GmNcl1功能标记的开发及验证[J].江苏农业学报,2017,(06):1227.[doi:doi:10.3969/j.issn.1000-4440.2017.06.005]
　NING Li-hua,HE Xiao-lan,ZHANG Da-yong.Development and validation of the function marker of soybean salt tolerance gene GmNcl1[J].,2017,(10):1227.[doi:doi:10.3969/j.issn.1000-4440.2017.06.005]
[4]杨艳丽,杨勇,李大红,等.转桃PpCuZnSOD基因大豆的耐旱性[J].江苏农业学报,2018,(05):978.[doi:doi:10.3969/j.issn.1000-4440.2018.05.003]
　YANG Yan-li,YANG Yong,LI Da-hong,et al.Drought tolerance of transgenic soybean with PpCuZnSOD gene[J].,2018,(10):978.[doi:doi:10.3969/j.issn.1000-4440.2018.05.003]
[5]孙彦坤,陈睿,李静,等.不同降雨年型下反枝苋和大豆光合特征的比较[J].江苏农业学报,2019,(03):554.[doi:doi:10.3969/j.issn.1000-4440.2019.03.008]
　SUN Yan-kun,CHEN Rui,LI Jing,et al.Comparison of photosynthetic characteristics between Amaranthus retroexus and Glycine max under different annual rainfall pattern[J].,2019,(10):554.[doi:doi:10.3969/j.issn.1000-4440.2019.03.008]
[6]曹媛媛,陈春,郭婷婷,等.亲和性促生菌DW12-L的定殖及其对大豆生长的影响[J].江苏农业学报,2019,(04):776.[doi:doi:10.3969/j.issn.1000-4440.2019.04.004]
　CAO Yuan yuan,CHEN Chun,GUO Ting ting,et al.Colonization of soybean affinity rhizobacteria strain DW12-L and its effects on soybean growth[J].,2019,(10):776.[doi:doi:10.3969/j.issn.1000-4440.2019.04.004]
[7]丁俊男,于少鹏,李鑫,等.生物炭对大豆生理指标和农艺性状的影响[J].江苏农业学报,2019,(04):784.[doi:doi:10.3969/j.issn.1000-4440.2019.04.005]
　DING Jun nan,YU Shao peng,LI Xin,et al.Effects of biochar application on soybean physiological indices and agronomic traits[J].,2019,(10):784.[doi:doi:10.3969/j.issn.1000-4440.2019.04.005]
[8]曹帅,杜仲阳,刘鹏,等.碱胁迫对大豆光合特性及内源激素含量的影响[J].江苏农业学报,2020,(02):284.[doi:doi:10.3969/j.issn.1000-4440.2020.02.005]
　CAO Shuai,DU Zhong-yang,LIU Peng,et al.Effects of alkaline stress on photosynthetic characteristics and endogenous hormone contents of soybean[J].,2020,(10):284.[doi:doi:10.3969/j.issn.1000-4440.2020.02.005]
[9]邱爽,张军,何佳琦,等.大豆GmGolS2-1基因高温胁迫诱导表达及转基因烟草鉴定[J].江苏农业学报,2021,(01):38.[doi:doi:10.3969/j.issn.1000-4440.2021.01.005]
　QIU Shuang,ZHANG Jun,HE Jia-qi,et al.Expression of soybean GmGolS2-1 induced by heat stress and identification of GmGolS2-1 transgenic tobacco[J].,2021,(10):38.[doi:doi:10.3969/j.issn.1000-4440.2021.01.005]
[10]张斌,陈丽娟,李其华,等.栽培大豆GRAS转录因子家族基因鉴定及其盐胁迫下表达模式分析[J].江苏农业学报,2021,(02):296.[doi:doi:10.3969/j.issn.1000-4440.2021.02.004]
　ZHANG Bin,CHEN Li-juan,LI Qi-hua,et al.Identification of gene of GRAS transcription factor family in cultivated soybean（Glycine max L.） and expression pattern analysis under salt stress[J].,2021,(10):296.[doi:doi:10.3969/j.issn.1000-4440.2021.02.004]

备注/Memo

备注/Memo:
收稿日期：2025-02-08基金项目：山东省蔬菜产业技术体系菏泽综合试验站项目(SDAIT-05-26)；山东省自然科学基金项目(ZR2024QC237)作者简介：张中起(1991-),男,山东菏泽人,硕士,农艺师,主要从事豆类育种和基因组学研究。(E-mail)soybean2021@163.com通讯作者：薛墨庸,(E-mail)moyongxue@163.com
更新日期/Last Update: 2025-11-17