宁夏枸杞LbWaxy基因家族鉴定及高浓度CO2处理下的基因表达特征分析

doi:10.16420/j.issn.0513-353x.2024-0814

摘要/Abstract

摘要：

Waxy是调控直链淀粉合成的关键基因，通过生物信息方法，鉴定了宁夏枸杞中7个LbWaxy基因家族成员，系统分析了其在CO₂浓度倍增处理下不同发育期器官组织中的表达特征。理化性质分析表明，LbWaxy基因家族主要分布于4条染色体上，且均含有Glycos_transf保守结构域；编码氨基酸数为613 ~ 1 408，蛋白分子量为67.19 ~ 158.23 kD，等电点介于5.06 ~ 8.29之间，均为亲水性蛋白，亚细胞定位预测主要位于细胞质中。进化树分析表明，LbWaxy与茄科植物及拟南芥的Waxy基因进化关系保守，主要聚类为3个大组和6个亚组。共线性分析发现LbWaxy基因与番茄、辣椒、茄子及拟南芥之间存在共线性基因对。蛋白互作网络分析表明LbWaxy2分别与LbWaxy1、LbWaxy4、LbWaxy5、LbWaxy7间存在互作关系。基于转录组和RT-qPCR分析表明，CO₂浓度倍增处理下，LbWaxy家族成员在不同发育时期的器官组织中差异表达，其中，7个成员在青果期的茎和转色期的叶中均显著上调；LbWaxy1、bWaxy2和LbWaxy7在不同发育时期茎、根及果中显著上调，表明其为响应CO₂浓度倍增参与宁夏枸杞淀粉合成的关键调控基因。

关键词: 宁夏枸杞, LbWaxy家族基因, 淀粉酶, 表达分析, 大气CO₂浓度

Abstract:

The Waxy gene，a key regulator of amylose synthesis，is central to starch biosynthesis. In this study，bioinformatic analysis identified seven LbWaxy genes in Goji berr（Lycium barbarum L.）. Systematically characterized their expression patterns across multiple tissues at different stages under elevated CO₂ concentrations. Genomic mapping showed that the LbWaxy genes are distributed across four chromosomes，and all encode proteins containing a Glycos_transf conserved domain. The encoded proteins range from 613 to 1 408 aa，with predicted molecular weights of 67.19 to 158.23 kDa and isoelectric points ranging from 5.06 to 8.29. Subcellular localization prediction indicated that all proteins are hydrophilic and primarily cytoplasmic. Phylogenetic analysis demonstrated a conserved evolutionary relationship between LbWaxy and Waxy genes from Solanaceae species and Arabidopsis thaliana，with the proteins clustering into three main groups and six subgroups. Collinearity analysis indicated syntenic relationships between the LbWaxy genes and homologs in tomato（Solanum lycopersicum），pepper（Capsicum annuum），eggplant（Solanum melongena），and Arabidopsis thaliana. Protein-protein interaction network predictions suggested potential interactions between LbWaxy2 and LbWaxy1，LbWaxy4，LbWaxy5，and LbWaxy7. Combined transcriptomic and RT-qPCR analyses revealed differential expression of LbWaxy genes under elevated CO₂ concentrations across tissues and stages. Notably，all seven members showed significant up-regulated in stems at the green fruit stage and in leaves at the different stage. Furthermore，LbWaxy1，LbWaxy2，and LbWaxy7 exhibited significant up-regulated in stems，roots，and fruits at multiple stages，suggesting their role as key regulatory genes in starch synthesis under elevated CO₂ concentrations.

Key words: Lycium barbarum, LbWaxy family genes, amylase, expression analysis, atmospheric CO₂ concentration

张婷, 王博涛, 张磊, 宋丽华, 曹兵, 马亚平. 宁夏枸杞LbWaxy基因家族鉴定及高浓度CO₂处理下的基因表达特征分析[J]. 园艺学报, 2025, 52(9): 2395-2409.

ZHANG Ting, WANG Botao, ZHANG Lei, SONG Lihua, CAO Bing, MA Yaping. Identification of the LbWaxy Gene Family in Lycium barbarum and Analysis of Gene Expression Characteristics Under High Concentrations of CO₂[J]. Acta Horticulturae Sinica, 2025, 52(9): 2395-2409.

图/表 11

图1 ‘宁杞1号’果实5个发育阶段及根、茎、叶、花、果形态特征

Fig. 1 Five fruit development stages and morphologicalcharacteristics of the root，stem，leaf，flower，and fruit in goji berry

表1 荧光定量PCR引物

Table 1 Primer sequences of quantitative real time PCR

基因 Gene	上游引物（5′-3′） Forward primer	下游引物（5′-3′） Reverse primer
ACTIN	CCATCTACGAGGGTTACGCTTTG	AGTCAAGAGCCACATAGGCAAGC
LbWaxy1	GATGGTTCTGGCTCTGTTGTTGG	CAGCATCCTCACTCAATTCATCTCC
LbWaxy2	GCCATTGTTCCTCCTCACTCATTC	GCGTCTACTGAAGTTGCTCCAC
LbWaxy3	CACAGATGATTGGAATGCGTTATGG	TCCTTCAAAGACAAACCCGTTAGC
LbWaxy4	GCAAGGTGGTCAGCAAGGTTC	AAGGAGATTAAGAACGGCAGACTTG
LbWaxy5	GATGGTCGGGATTGGTTCAACTC	AGTAGTCAAGAGCAGGTCGGTTC
LbWaxy6	AGCAAGCATCCATTATTCTCGTAGG	ACACCTCTCCTTGAAGCCATCC
LbWaxy7	GGCAGGCATGACCTTGAACAG	CCAGCAGTTATACGATGAGCAGTC

表2 宁夏枸杞LbWaxy蛋白理化性质特征分析

Table 2 Feature analysis of LbWaxy in Lycium barbarum L.

基因名称 Gene name	基因ID Gene ID	外显子长度/bp CDS	理化性质 Physicochemical properties
基因名称 Gene name	基因ID Gene ID	外显子长度/bp CDS	分子量/KD Molecular weight	等电点 Theoretical Pi	不稳定指数 Instability index	脂肪族指数 Aliphatic index	亲水性 Grand average of hydrophilicity
LbWaxy1	Lbar03T009702.1	2 427	71.14	5.33	32.62	85.42	-0.165
LbWaxy2	Lbar04T011539.1	3 573	67.19	8.29	26.61	89.36	-0.076
LbWaxy3	Lbar11T036582.1	2 421	69.50	5.71	40.99	86.48	-0.207
LbWaxy4	Lbar12T040012.1	4 352	118.86	5.66	52.65	90.11	-0.463
LbWaxy5	Lbar12T040660.1	4 693	158.23	5.06	41.41	78.66	-0.508
LbWaxy6	Lbar12T041078.5	1 863	70.42	6.12	40.81	93.87	-0.244
LbWaxy7	Lbar12T041516.1	3 000	83.44	6.06	43.01	79.14	-0.410
基因名称 Gene name	基因ID Gene ID	蛋白长度/aa Protein	第二结构 Secondary structure			亚细胞定位 Subcellular
基因名称 Gene name	基因ID Gene ID	蛋白长度/aa Protein	α螺旋/% α helix	自由基/% Random coil	延伸链/% Extended strand	亚细胞定位 Subcellular
LbWaxy1	Lbar03T009702.1	646	33.59	50.77	15.63	细胞质 Cytoplasmic
LbWaxy2	Lbar04T011539.1	613	34.42	53.18	12.40	叶绿体 Chloroplast
LbWaxy3	Lbar11T036582.1	619	34.73	52.67	12.60	细胞质 Cytoplasmic
LbWaxy4	Lbar12T040012.1	1 043	57.62	34.32	8.05	细胞核 Nuclear
LbWaxy5	Lbar12T040660.1	1 408	36.79	48.79	14.42	细胞质 Cytoplasmic
LbWaxy6	Lbar12T041078.5	620	44.19	43.39	12.42	细胞质 Cytoplasmic
LbWaxy7	Lbar12T041516.1	754	39.52	49.20	11.27	叶绿体 Chloroplast

图2 7个LbWaxy蛋白的三级结构

Fig. 2 Tertiary structure of seven LbWaxy proteins

图3 宁夏枸杞LbWaxy基因家族系统发育树（a），基序（b）和基因结构（c）

Fig. 3 Phylogenetic tree（a），conserved motif（b）and gene structure（c）in LbWaxy gene family

图4 宁夏枸杞、辣椒、茄子、番茄、马铃薯和拟南芥Waxy基因家族进化树分析

Fig. 4 The unrooted phylogenetic tree of Waxy gene family in Lycium barbarum，Capsicum annuum，Solanum melongena，S. lycopersicum，S. tuberosum and Arabidopsis thaliana

图5 LbWaxy基因家族的染色体分布（a）及顺式作用元件预测（b）

Fig. 5 Chromosomal localization（a）and predicted cis-acting elements（b）of LbWaxy gene family

图6 不同物种Waxy基因的共线性分析

Fig. 6 Covariance analysis of Waxy genes in different species

图7 LbWaxy蛋白相互作用网络

Fig. 7 Interaction network of LbWaxy proteins

图8 CO2浓度升高处理下宁夏枸杞各个器官的转录组表达分析 CK：自然CO2浓度处理；CO2：CO2浓度倍增处理；每个处理3次重复

Fig. 8 Transcriptome analysis of Lycium barbarum treated with elevated CO2 concentration CK：Normal CO2 concentration；CO2：Elevated CO2 concentration；each experiment was repeated three times

图 9 CO2升高处理下宁夏枸杞5个器官中Waxy基因表达分析 S1：幼果期；S2：青果期；S3：转色期；S4：初熟期；S5：成熟期

Fig. 9 Expression analysis of LbWaxy genes in five organs under elevated CO2 treatment S1：Young fruit stage；S2：Green fruit stage；S3：Color turning stage；S4：Initial ripening stage；S5：Mature stage

参考文献 24

[1]	Chen C J, Chen H, Zhang Y, Thomas H R, Frank M H, He Y H, Xia R. 2020. TBtools:an integrative toolkit developed for interactive analyses of big biological data. Molecular Plant, 13 (8):1194-1202.
[2]	Chen Haiyan, Li Dehong. 2010. Progress in effects of light,saccharide and hormones on the anthocyanin synthesis and accumulation in plants. Subtropical Plant Science, 39 (3):82-86. (in Chinese)
	程海燕, 李德红. 2010. 光、糖与激素影响植物花色素苷合成与积累的研究进展. 亚热带植物科学, 39 (3):82-86.
[3]	Feng Xuerui, Ma Yaping, Liu Jiaxin, Lu Hui, Li Yunmao, Cao Bing. 2024. Analysis of enzyme and gene expression related to sugar metabolismin Lycium barbarum under elevated CO₂ concentration treatment. Scientia Silvae Sinicae, 60 (3):10-21. (in Chinese)
	冯学瑞, 马亚平, 刘佳欣, 陆晖, 李运毛, 曹兵. 2024. CO₂浓度升高处理下宁夏枸杞糖代谢相关酶及基因表达分析. 林业科学, 60 (3):10-21.
[4]	Guo Changquan, Li Danqi, Hui Xinran, Zheng Jingya, Hou Menglu, Zhu Yongxing. 2024. Identification and expression pattern analysis of DUF 966 gene family members in ginger. Acta Horticulturae Sinica, 51 (9):2031-2047. (in Chinese)
	郭昌权, 李丹琪, 惠馨冉, 郑婧雅, 侯梦璐, 朱永兴. 2024. 姜DUF966基因家族成员鉴定与表达分析. 园艺学报, 51 (9):2031-2047.
[5]	Guo Fangyun, Cao Bing, Song Lihua, Ha Rong. 2020. Effects of elevated CO₂ concentration on Lycium barbarum fruit morphological parameters and sugar accumulation during development period in Ningxia. Journal of Nanjing Forestry University(Natural Sciences Edition), 44 (1):105-110. (in Chinese)
	郭芳芸, 曹兵, 宋丽华, 哈蓉. 2020. CO₂浓度升高对宁夏枸杞果实发育期形态指标及糖分积累影响. 南京林业大学学报(自然科学版), 44 (1):105-110. doi: 10.3969/j.issn.1000-2006.201901024
[6]	Guzmán C, Caballero L, Martín L M, Alvarez J B. 2012. Waxy genes from spelt wheat:new alleles for modern wheat breeding and new phylogenetic inferences about the origin of this species. Annals of Botany, 110 (6):1161-1171.
[7]	Liang Wangli, Yu Wenjing, Hu Jinhong, Song Fan, Wang Lingxia, Liang Wenyu. 2024. Differential expression of ABA metabolism-related genes in Lycium barbarum under NaCl stress. Acta Agriculturae Boreali-occidentalis Sinica, 33 (4):664-672. (in Chinese)
	梁旺利, 于雯静, 胡进红, 宋繁, 王玲霞, 梁文裕. 2024. NaCl胁迫下宁夏枸杞ABA代谢相关基因差异表达分析. 西北农业学报, 33 (4):664-672.
[8]	Ma Y P, Devi M J, R Reddy V, Song L H, Gao H D, Cao B. 2021. Cloning and characterization of three sugar metabolism genes(LBGAE, LBGALA,and LBMS)regulated in response to elevated CO₂ in Goji Berry(Lycium barbarum L.). Plants, 10 (2):321.
[9]	Ma Y P, Han Y R, Feng X R, Gao H D, Cao B, Song L H. 2022. Genome-wide identification of gene family in jujube and expression in response to abiotic stress. BMC Genomics, 23 (1):438.
[10]	Ma Yaping, Feng Xuerui, Gao Handong, Song Lihua, Cao Bing. 2024. Effects of simulated elevated CO₂ concentration and atmospheric temperature on quality formation of Lycium barbarum fruits. Scientia Silvae Sinicae, 60 (3):1-9. (in Chinese)
	马亚平, 冯学瑞, 高捍东, 宋丽华, 曹兵. 2024. 模拟CO₂浓度和温度升高对宁夏枸杞果实品质形成的影响. 林业科学, 60 (3):1-9.
[11]	Maeda H A, Fernie A R. 2021. Evolutionary history of plant metabolism. Annual Review of Plant Biology,72:185-216.
[12]	Momma M, Fujimoto Z. 2012. Interdomain disulfide bridge in the rice granule bound starch synthase I catalytic domain as elucidated by X-ray structure analysis. Biosci Biotechnol Biochem, 76 (8):1591-1605.
[13]	Niu Z G, He H L, Yu P T, Sitch S, Zhao Y, Wang Y H, Jain A K, Vuichard N, Si B C. 2023. Climate change and CO₂ fertilization have played important roles in the recent decadal vegetation greening trend on the chinese loess plateau. Remote Sensing, 15 (5):1233.
[14]	Nelson O E. 1975. The Waxy locus in maize III. effect of structural heterozygosity on intragenic recombination and flanking marker assortment. Genetics, 79 (1):31-44. doi: 10.1093/genetics/79.1.31 pmid: 17248676
[15]	Park Y J, Nemoto K, Nishikawa T, Matsushima K, Minami M, Kawase M. 2011. Genetic diversity and expression analysis of granule bound starch synthase I gene in the new world grain amaranth(Amaranthus cruentus L.). Journal of Cereal Science, 53 (3):298-305.
[16]	Peng Bo, A Xinxiang, Zheng Mengyang, Guo Ying, Gu Yu, Zhang Kaixuan, Peng Juan, Song Xiaohua, Tian Xiayu, Liu Ziyue, Sun Yanfang, Pang Ruihua, Li Jintao, Wang Quanxiu, Zhou Wei. 2020. Detection and analysis of Waxy gene in Yunnan soft rice germplasm resources. Southwest China. Journal of Agricultural Sciences, 33 (12):2693-2701. (in Chinese)
	彭波, 阿新祥, 郑梦阳, 郭莹, 谷雨, 张凯璇, 彭娟, 宋晓华, 田夏雨, 刘子月, 孙艳芳, 庞瑞华, 李金涛, 汪全秀, 周伟. 2020. 云南软米水稻种质资源Waxy基因的检测与分析. 西南农业学报, 33 (12):2693-2701.
[17]	Qi X T, Wu H, Jiang H Y, Zhu J J, Huang C L, Zhang X, Liu C L, Cheng B J. 2020. Conversion of a normal maize hybrid into a waxy version using in vivo CRISPR/Cas 9 targeted mutation activity. The Crop Journal, 8 (3):440-448.
[18]	Qiang J, Ding R, Kang C, Xiao T T, Yan Y M. 2025. Impact of waxy protein deletions on the crystalline structure and physicochemical properties of wheat V-type resistant starch(RS5). Carbohydrate Polymers,347:122695.
[19]	Shure M, Wessler S, Fedoroff N. 1983. Molecular identification and isolation of the Waxy locus in maize. Cell,35:225-233.
[20]	Sun Tingzhen, Shu Qin, Ma Wei, Shi Yuzi, Zhang Meng, Xiang Chenggang, Bo Kailiang, Duan Ying, Wang Changlin. 2025. Identification and phylogenetic analysis of gibberellin oxidase gene family and its expression pattern in main vine growth in Cucurbita maxima. Acta Horticulturae Sinica, 52 (3):603-622. (in Chinese) doi: 10.16420/j.issn.0513-353x.2023-0974
	孙廷珍, 疏琴, 马玮, 史玉滋, 张蒙, 向成钢, 薄凯亮, 段颖, 王长林. 2025. 印度南瓜GAox家族基因鉴定及其在不同蔓长种质中的表达. 园艺学报, 52 (3):603-622. doi: 10.16420/j.issn.0513-353x.2023-0974
[21]	Sankoff D, Zheng C F. 2018. Whole genome duplication in plants:implications for evolutionary analysis. Methods in Molecular Biology,1704:291-315.
[22]	Victor T A, Yifeng W, Lijuan C, Huimei W, Wanning L, Xingyong L, Yicheng C, Xiaohong T, Jiezheng Y, Jian Z. 2021. Posttranslational modification of Waxy to genetically improve starch quality in rice grain. International Journal of Molecular Sciences, 22 (9):4845.
[23]	Wang Hongmei, Dong Kongjun, He Jihong, Bai Bin, Liu Tianpeng, Ren Ruiyu, Liu Xinxing. 2024. Analysis of amylose content and allelic variation of Waxy gene in broomcorn millet germplasm resources. Journal of Plant Genetic Resources, 25 (7):1140-1152. (in Chinese)
	王红梅, 董孔军, 何继红, 白斌, 刘天鹏, 任瑞玉, 刘新星. 2024. 糜子种质资源淀粉含量与Waxy基因等位变异分析. 植物遗传资源学报, 25 (7):1140-1152.
[25]	Wang Q P, Zhang L M, Xue C L, Zhang Y, Meng X R, Liu Z G, Liu M J, Zhao J. 2024. GST family genes in jujube actively respond to phytoplasma infection. Horticultural Plant Journal, 10 (1):77-90.
[26]	Wang Qian, Sun Wenjing, Bao Ying. 2017. Evolutionary pattern of the GBSS gene family in plants. Chinese Bulletin of Botany, 52 (2):179-187. (in Chinese)
	王倩, 孙文静, 包颖. 2017. 植物颗粒结合淀粉合酶GBSS基因家族的进化. 植物学报, 52 (2):179-187. doi: 10.11983/CBB16041
[27]	Wang Yanhong, Xi Keyong, Tian Ye, Liu Deqi, Zhou Kegui, Yin Junliang, Liu Yiqing, Zhu Yongxing. 2024. Identification and expression pattern analysis of GSTs in zingiber officinale. Acta Horticulturae Sinica, 51 (8):1803-1822. (in Chinese)
	王艳红, 席克勇, 田野, 刘德麒, 周克贵, 尹军良, 刘奕清, 朱永兴. 2024. 姜GST基因家族成员鉴定与表达分析. 园艺学报, 51 (8):1803-1822.
[28]	Yang Furong, Du Bing, Wang Chen, Zhang Fuhou, Men Chaomin. 2024. Identification of the CPP gene family and its response to exogenous selenium in foxtail millet. Acta Botanica Boreali-Occidentalia Sinica, 44 (8):1250-1260. (in Chinese)
	杨馥熔, 杜冰, 郭浩杰, 王成, 张富厚, 孟超敏. 2024. 谷子CPP基因家族鉴定及外源硒响应分析. 西北植物学报, 44 (8):1250-1260.
[24]	Wang L, Liu L L, Zhao J L, Li C L, Wu H L, Zhao H X, Wu Q. 2023. Granule-bound starch synthase in plants:towards an understanding of their evolution,regulatory mechanisms,applications and perspectives. Plant Science,336:111843.

宁夏枸杞LbWaxy基因家族鉴定及高浓度CO₂处理下的基因表达特征分析

Identification of the LbWaxy Gene Family in Lycium barbarum and Analysis of Gene Expression Characteristics Under High Concentrations of CO₂

RichHTML

PDF

可视化

摘要/Abstract

引用本文

使用本文

图/表 11

参考文献 24

相关文章 15

编辑推荐

Metrics

本文评价

访问总数:　当日访问总数:　当前在线人数:

[1]	王宝琛, 罗陈, 闫晋强, 程晓欣, 莫仁连, 刘文睿, 张余洋, 江彪. 冬瓜BhYAB4b基因的克隆、表达及功能研究[J]. 园艺学报, 2025, 52(9): 2353-2362.
[2]	张拓, 聂梦涵, 刘媛媛, 段昕莲, 韩星, 谢宝贵, 严俊杰. 金针菇过氧化氢酶基因FfCAT1和FfCAT2的表达分析[J]. 园艺学报, 2025, 52(7): 1769-1779.
[3]	孙廷珍, 疏琴, 马玮, 史玉滋, 张蒙, 向成钢, 薄凯亮, 段颖, 王长林. 印度南瓜GAox家族基因鉴定及其在不同蔓长种质中的表达[J]. 园艺学报, 2025, 52(3): 603-622.
[4]	邓淑琴, 高莹瑞, 李雨桐, 王瑛, 龚春梅, 白娟. 茶树泛素连接酶基因CsPUB21对非生物胁迫的响应[J]. 园艺学报, 2025, 52(3): 655-670.
[5]	仝宗军, 韩星, 段昕莲, 刘媛媛, 林俊彬, 甘颖, 陈杰, 谢宝贵, 甘炳成, 严俊杰. 金针菇PRX家族基因鉴定及其在子实体发育过程中的表达分析[J]. 园艺学报, 2025, 52(2): 337-348.
[6]	贺丹丹, 何宏泰, 王文庭, 周文美, 刘燕敏, 刘骕骦. 甜瓜GolS家族基因鉴定及其响应低温胁迫的表达分析[J]. 园艺学报, 2025, 52(1): 136-148.
[7]	郭昌权, 李丹琪, 惠馨冉, 郑婧雅, 侯梦璐, 朱永兴. 姜DUF966基因家族成员鉴定与表达分析[J]. 园艺学报, 2024, 51(9): 2031-2047.
[8]	赵佳莹, 曾周婷, 岑欣颖, 施姣淇, 李效贤, 沈晓霞, 俞振明. 铁皮石斛CCO基因家族鉴定及其在花发育中的表达分析[J]. 园艺学报, 2024, 51(9): 2075-2088.
[9]	刘艳艳, 丁颖, 刘兴华, 郑佳秋, 刘志钦. 辣椒CaSYT1的鉴定及其在疫霉侵染过程中的功能初探[J]. 园艺学报, 2024, 51(3): 533-544.
[10]	杨雪, 胡进红, 李晶晶, 章英才, 王玲霞, 梁文裕. 宁夏枸杞TGA2转化拟南芥增强其耐盐性[J]. 园艺学报, 2024, 51(12): 2829-2842.
[11]	李宇腾, 陈瑶, 任恒泽, 李聪聪, 王浩乾, 曹红利, 岳川, 郝心愿, 王新超. 茶树CsIDM的鉴定、表达分析及互作验证[J]. 园艺学报, 2023, 50(8): 1679-1696.
[12]	薛珍珍, 关鸿发, 李娜, 李凌飞, 钟春梅. 铁十字秋海棠GASA家族全基因组鉴定及叶斑形成的初步探索[J]. 园艺学报, 2023, 50(7): 1482-1494.
[13]	王同欢, 吴雨馨, 武艺圆, 李鑫鑫, 刘梦阳, 杨莲莲, 李佳鹏, 张忠山, 曹访, 仲雪婷, 王占旗. 菊花脑GRAS家族鉴定及其低温胁迫响应表达分析[J]. 园艺学报, 2023, 50(4): 815-830.
[14]	刘慧, 黄婷, 陶建平, 张佳琪, 张榕蓉, 宋刘霞, 赵统敏, 游雄, 熊爱生. 番茄生物钟基因SlLNK1、SlEID1和SlELF3光周期响应分析[J]. 园艺学报, 2023, 50(10): 2079-2090.
[15]	王亦栖, 颜爽爽, 余炳伟, 甘雨薇, 邱正坤, 朱张生, 陈长明, 曹必好. 茄子青枯病抗性相关的E3泛素连接酶基因的筛选及鉴定[J]. 园艺学报, 2023, 50(10): 2271-2287.