赤霉素诱导葡萄无核形成机理研究进展

doi:10.16420/j.issn.0513-353x.2024-0030

摘要/Abstract

摘要：

赤霉素能够有效诱导葡萄胚珠败育，被广泛应用于葡萄的无核化生产。赤霉素诱导葡萄胚囊和花粉发育异常，造成内部生理代谢失调，该过程还受到一些植物激素合成代谢及信号通路相关基因、种子发育时期赤霉素响应基因以及特异microRNA的动态调控。本文中从胚胎学、生理生化和分子机制层面综述了赤霉素诱导葡萄无核的形成机理，以期为进一步揭示赤霉素诱导葡萄单性结实的潜在调控机制和今后利用赤霉素进行优质无核葡萄的生产提供依据。

关键词: 葡萄, 无核, 赤霉素, 单性结实, 调控机理

Abstract:

Gibberellins（GAs）can effectively induce ovule abortion and are widely utilized in the production of seedless grape. Gibberellins induces abnormal development of grape embryo sac and pollen，resulting in internal physiological metabolism disorders. This process is also dynamically regulated by some genes related to plant hormone anabolism and signaling pathways，gibberellins response genes and specific microRNA during seed development. This review summarizes the mechanism of GAs-induced seedlessness in grape from the aspects of embryology，physiology，biochemistry and molecular mechanisms，with the aim of further understanding the potential regulatory mechanism of GAs-induced parthenocarpy of grape and providing a basis for using GAs in the future production of high-quality seedless grape.

Key words: grape, seedless, gibberellins, parthenocarpy, regulatory mechanism

刘苗苗, 姚锦, 包敏, 楚言言, 王西平. 赤霉素诱导葡萄无核形成机理研究进展[J]. 园艺学报, 2024, 51(7): 1610-1622.

LIU Miaomiao, YAO Jin, BAO Min, CHU Yanyan, WANG Xiping. Research Progress on the Mechanism of GAs-Induced Seedlessness in Grape[J]. Acta Horticulturae Sinica, 2024, 51(7): 1610-1622.

图/表 4

参考文献 60

[1]	Acheampong A K, Hu J, Rotman A, Zheng C, Halaly T, Takebayashi Y, Jikumaru Y, Kamiya Y, Lichter A, Sun T, Or E. 2015. Functional characterization and developmental expression profiling of gibberellin signalling components in Vitis vinifera. Journal of Experimental Botany, 66 (5):1463-1476. doi: 10.1093/jxb/eru504 pmid: 25588745
[2]	Bai Y, Wang Z, Luo L, Xuan X, Tang W, Qu Z, Dong T, Qi Z, Yu M, Wu W, Fang J, Wang C. 2023. Characterization of VvmiR166s-target modules and their interaction pathways in modulation of gibberellic-acid-induced grape seedless berries. International Journal of Molecular Sciences, 24 (22):16279.
[3]	Bai Y, Zhang X, Xuan X, Sadeghnezhad E, Liu F, Dong T, Pei D, Fang J, Wang C. 2022. miR3633a-GA3ox2 module conducts grape seed-embryo abortion in response to gibberellin. International Journal of Molecular Sciences, 23 (15):8767-8767.
[4]	Bai Yunhe, Wang Wenran, Dong Tianyu, Guan Le, Su Ziwen, Jia Haifeng, Fang Jinggui, Wang Chen. 2020. vvi-miR160s in mediating VvARF18response to gibberellin regulation of grape seed development. Scientia Agricultura Sinica, 53 (9):1890-1903. (in Chinese)
	白云赫, 王文然, 董天宇, 管乐, 宿子文, 贾海锋, 房经贵, 王晨. 2020. vvi-miR160s介导VvARF18应答赤霉素调控葡萄种子的发育. 中国农业科学, 53 (9):1890-1903. doi: 10.3864/j.issn.0578-1752.2020.09.015
[5]	Baydar N G, Harmankaya N. 2005. Changes in endogenous hormone levels during the ripening of grape cultivars having different berry set mechanisms. Turkish Journal of Agriculture and Forestry, 29 (3):205-210.
[6]	Bouquet A, Danglot Y. 1996. Inheritance of seedlessness in grapevine(Vitis viniferal L.). Vitis, 35 (1):35-42.
[7]	Cheng C, Jiao C, Singer S D, Gao M, Xu X, Zhou Y, Li Z, Fei Z, Wang Y, Wang X. 2015. Gibberellin-induced changes in the transcriptome of grapevine(Vitis labrusca × V. vinifera)cv. Kyoho flowers. BMC Genomics, 16 (1):128.
[8]	Cheng C, Xu X, Singer S D, Li J, Zhang H, Gao M, Zhang H, Gao M, Wang L, Song J, Wang X, Blazquez M A. 2013. Effect of GA₃ treatment on seed development and seed-related gene expression in grape. PLoS ONE, 8 (11):e80044.
[9]	Cui M, Wang C, Zhang W, Pervaiz T, Haider M S, Tang W, Fang J. 2018. Characterization of Vv-miR156:Vv-SPL pairs involved in the modulation of grape berry development and ripening. Molecular Genetics and Genomics, 293:1333-1354.
[10]	Cui M, Wang W, Guo F, Fan X, Liu C. 2020. Characterization and temporal-spatial expression analysis of LEC1 gene in the development of seedless berries in grape induced by gibberellin. Plant Growth Regulation, 90 (44):585-596.
[11]	Cui Mengjie, Guo Fengfei, Wang Chen, Jiu Songtao, Zhu Xudong, Fang Jinggui. 2019. Identification and roles of VvAGL11 and VvAGL15 gene in the development process of seedless grape berry induced by gibberellin. Journal of Nanjing Agricultural University, 42 (2):261-269. (in Chinese)
	崔梦杰, 郭凤菲, 王晨, 纠松涛, 朱旭东, 房经贵. 2019. 葡萄VvAGL11和VvAGL15基因的鉴定及其在赤霉素诱导葡萄无核果实发育过程中的作用. 南京农业大学学报, 42 (2):261-269.
[12]	Davière J M, Achard P. 2013. Gibberellin signaling in plants. Development, 140 (6):1147-1151.
[13]	Dong Shujuan, Guo Xiuwu. 2008. Change of protective enzyme activity,contents of MDA and inner nutriments during the embryonic development of several grape cultivars. Sino-Overseas Grapevine & Wine,(2):12-16. (in Chinese)
	董姝娟, 郭修武. 2008. 几个葡萄品种胚胎发育过程中保护酶活性、丙二醛及内含营养物质含量的变化. 中外葡萄与葡萄酒,(2):12-16.
[14]	Fos M, Proano K, Alabadi D, Nuez F, Carbonell J, Garcia-Martinez J L. 2003. Polyamine metabolism is altered in unpollinated parthenocarpic pat-2 tomato ovaries. Plant Physiology, 131:359-366.
[15]	Giacomelli L, Rota-Stabelli O, Masuero D, Acheampong A K, Moretto M, Caputi L, Vrhovsek U, Moser C. 2013. Gibberellin metabolism in Vitis vinifera L. during bloom and fruit-set:functional characterization and evolution of grapevine gibberellin oxidases. Journal of Experimental Botany, 64 (14):4403-4419. doi: 10.1093/jxb/ert251 pmid: 24006417
[16]	Gong Shufang, Yang Guohui, Wang Junhong. 2000. A study on the causes of gibberellin induced grape seedlessness. Northern Horticulture,(5):51. (in Chinese)
	龚束芳, 杨国慧, 王军虹. 2000. 赤霉素诱导葡萄无核原因研究. 北方园艺,(5):51.
[17]	Han J, Fang J, Wang C, Yin Y, Sun X, Leng X, Song C. 2014. Grapevine microRNAs responsive to exogenous gibberellin. BMC Genomics, 15 (1):111.
[18]	Hanania U, Velcheva M, Or E, Flaishman M, Sahar N, Perl A. 2007. Silencing of chaperonin 21,that was differentially expressed in inflorescence of seedless and seeded grapes,promoted seed abortion in tobacco and tomato fruits. Transgenic Research, 16:515-525. doi: 10.1007/s11248-006-9044-0 pmid: 17103240
[19]	Ishibashi Y, Tawaratsumida T, Kondo K, Kasa S, Sakamoto M, Aoki N, Zheng S, Yuasa T, lwaya-Inoue M. 2012. Reactive oxygen species are involved in gibberellin/abscisic acid signaling in barley aleurone cells. Plant Physiology, 158 (4):1705-1714. doi: 10.1104/pp.111.192740 pmid: 22291200
[20]	Ji Wei, Guo Rongrong, Wang Jingbo, Jiao Xiaobo, Yan Zhao, Chang Qinxiang, Dong Zhigang, Wang Yuejin. 2019. Grey correlation analysis of physiological and biochemical factors in embryo abortion of seedless grape. Acta Horticulturae Sinica, 46 (8):1473-1485. (in Chinese) doi: 10.16420/j.issn.0513-353x.2019-0011
	纪薇, 郭荣荣, 王静波, 焦晓博, 闫钊, 昌秦湘, 董志刚, 王跃进. 2019. 无核葡萄胚败育生理生化因子灰色关联分析. 园艺学报, 46 (8):1473-1485. doi: 10.16420/j.issn.0513-353x.2019-0011
[21]	Jin Yueqin, Chang Li, Wang Xiaofang, Han Bing, Ji Wei. 2020. Research progress of grape seedless formation and regulation mechanism. Journal of Shanxi Agricultural Sciences, 48 (9):1537-1542,1546. (in Chinese)
	靳月琴, 常丽, 王晓芳, 韩冰, 纪薇. 2020. 葡萄无核形成与调控机制研究进展. 山西农业科学, 48 (9):1537-1542,1546.
[22]	Jung C J, Hur Y Y, Yu H J, Noh J H, Park K S, Lee H J. 2014. Gibberellin application at pre-bloom in grapevines down-regulates the expressions of VvIAA9 and VvARF7,negative regulators of fruit set initiation,during parthenocarpic fruit development. PLoS ONE, 9 (4):e95634.
[23]	Jusoh M, Loh S H, Aziz A, San C T. 2019. Gibberellin promotes cell growth and induces changes in fatty acid biosynthesis and upregulates fatty acid biosynthetic genes in Chlorella vulgaris UMT-M1. Applied Biochemistry and Biotechnology, 188:450-459.
[24]	Lankinen A, Armbruster W S, Antonsen L. 2007. Delayed stigma receptivity in Collinsia heterophylla(Plantaginaceae):genetic variation and adaptive significance in relation to pollen competition,delayed self-pollination,and mating-system evolution. American Journal of Botany, 94 (7):1183-1192. doi: 10.3732/ajb.94.7.1183 pmid: 21636485
[25]	Li Shunyu. 2009. Morphology and physiological foundation of embryo abortion in stenospermocarpic grapes[M. D. Dissertation]. Guiyang: Guizhou University. (in Chinese)
	李顺雨. 2009. 种子败育型葡萄胚败育的形态学观察及生理基础[硕士论文]. 贵阳: 贵州大学.
[26]	Li Xiaopeng. 2019. Study on the physiological and molecular mechanism of gibberellin-induced seediess of‘Zuijinxiang’grape[Ph. D. Dissertation]. Nanjing: Nanjing Agricultural University. (in Chinese)
	李晓鹏. 2019. 赤霉素诱导‘醉金香’葡萄果实无核生理及分子机理的研究[博士论文]. 南京: 南京农业大学.
[27]	Liu C S, Xiao P S, Jiang F, Wang S Y, Liu Z, Song G Z, Li W, Lv T X, Li J, Wang D M, Li Y, Wu C B, Li T Z. 2022. Exogenous gibberellin treatment improves fruit quality in self-pollinated apple. Plant Physiology and Biochemistry, 174:11-21. doi: 10.1016/j.plaphy.2022.01.029 pmid: 35121481
[28]	Liu Jinbiao, Li Hongyan, Cao Xiongjun, Shi Xiaofang, Zhang Ying, Zhou Yongmei, Xie Taili. 2017. Effects of different treatments on seedlessness and fruit quality of‘Kyoho’grape. Southwest China Journal of Agricultural Sciences, 30 (12):2788-2792. (in Chinese)
	刘金标, 李洪艳, 曹雄军, 时晓芳, 张瑛, 周咏梅, 谢太理. 2017. 不同处理对‘巨峰’葡萄无核化及果实品质的影响. 西南农业学报, 30 (12):2788-2792.
[29]	Liu Jing, Dong Yang, Nong Huilan, Yu Xin, Huang Liyuan, Zheng Huan, Tao Jianmin. 2023. Changes in grape fruit shape and their physiological and molecular mechanisms under GA₃ treatment. Jiangsu Agricultural Sciences, 51 (2):153-159. (in Chinese)
	刘静, 董阳, 农慧兰, 余欣, 黄丽媛, 郑焕, 陶建敏. 2023. GA₃处理下葡萄果形变化及其生理和分子机制. 江苏农业科学, 51 (2):153-159.
[30]	Lu Long. 2016. The study on the molecular mechanism of gibberellin-induced parthenocarpy and improved fruit set in grapevine[Ph. D. Dissertation]. Beijing: China Agricultural University. (in Chinese)
	卢龙. 2016. 赤霉素诱导葡萄单性结实与促进坐果分子机制的研究[博士论文]. 北京: 中国农业大学.
[31]	Nishiyama S, Yoshimura D, Sato A, Yonemori K. 2022. Characterization of tissue-specific transcriptomic responses to seedlessness induction by gibberellin in table grape. The Horticulture Journal, 91 (2):157-168.
[32]	Okamoto G, Miura K. 2015. Effect of pre-bloom GA application on pollen tube growth in cv. Delaware grape pistils. Vitis Journal of Grapevine Research, 44 (4):157-160.
[33]	Pan Xuejun, Li Shunyu, Zhang Wene, Liu Chonghuai. 2011. Endogenous polyamines in stenospermocarpic grape ovules and their relationship with ovule development and abortion. Journal of Fruit Science, 28 (5):770-775. (in Chinese)
	潘学军, 李顺雨, 张文娥, 刘崇怀. 2011. 种子败育型葡萄胚珠中内源多胺含量与胚珠发育及败育的关系. 果树学报, 28 (5):770-775.
[34]	Pérez F J, Viani C, Retamales J. 2000. Bioactive gibberellins in seeded and seedless grapes:identification and changes in content during berry development. American Journal of Enology & Viticulture, 51 (4):315-318.
[35]	Sheng Z, Xuan X, Wang F, Sadeghnezhad E, Gong P, Xiao Y, Dong T, Zhang P, Wang X, Fang J, Wang C. 2022. Identification and characterization of auxin response factor gene family reveals their regulatory network to respond the multi-hormones crosstalk during GA-induced grape parthenocarpic berry. International Journal of Molecular Sciences, 23 (19):11108.
[36]	Suehiro Y, Mochida K, Tsuma M, Yasuda Y, Itamura H, Esumi T. 2019. Effects of gibberellic acid/cytokinin treatments on berry development and maturation in the yellow-green skinned‘shine muscat’grape. The Horticulture Journal, 88 (2):202-213.
[37]	Shiozaki S, Ogata T, Horiuchi S. 2000. Endogenous polyamines in the pericarp and seed of the grape berry during development and ripening. Scientia Horticulturae, 83 (1):33-41.
[38]	Su Z, Xuan X, Sheng Z, Wang F, Zhang X, Ye D, Wang X, Dong T, Pei D, Zhang P, Fang J, Wang C. 2022. Characterization and regulatory mechanism analysis of VvmiR156a-VvAGL80 pair during grapevine flowering and parthenocarpy process induced by gibberellin. The Plant Genome, 15:e20181.
[39]	Tao Jianmin, Han Chuanguang, Zhang Zhen, Xu Xilou. 2003. Application of GA₃ in grape production. Sino-Overseas Grapevine & Wine,(6):33-35. (in Chinese)
	陶建敏, 韩传光, 章镇, 徐喜楼. 2003. GA₃在葡萄生产上的应用. 中外葡萄与葡萄酒,(6):33-35.
[40]	Wang B, Wang J, Wang C, Shen W, Jia H, Zhu X, Li X. 2016. Study on modes of expression and cleavage role of miR156b/c/d and its target gene Vv-SPL9 during the whole growth stage of grapevine. Journal of Heredity, 107:626-634.
[41]	Wang C, Jogaiah S, Zhang W, Abdelrahman M, Fang J. 2018. Modulation of VvmiR159-mediated VvGAMYB involved in the floral development during grape parthenocarpy process induced by GA. Journal of Experimental Botany, 69:3639-3650.
[42]	Wang Fei, Wang Yuejin, Wan Yizhen, Ren Jie. 2004. Some physiological and biochemical changes of embryo abortion in hybrids of seedless grapes and Chinese wild grapes. Acta Horticulturae Sinica, 31 (5):651-653. (in Chinese)
	王飞, 王跃进, 万怡震, 任杰. 2004. 无核葡萄与中国野生葡萄杂种胚败育的某些生理生化变化. 园艺学报, 31 (5):651-653.
[43]	Wang H, Caruso L V, Downie A B, Perry S E. 2004. The embryo MADS domain protein AGAMOUS-Like 15 directly regulates expression of a gene encoding an enzyme involved in gibberellin metabolism. The Plant Cell, 16 (5):1206-1219.
[44]	Wang Huan. 2003. Study on technology of seedlessness and mechanism in fresh grapes[M. D. Dissertation]. Urumqi: Xinjiang Agricultural University. (in Chinese)
	王欢. 2003. 鲜食葡萄无核化技术及机理的研究[硕士论文]. 乌鲁木齐: 新疆农业大学.
[45]	Wang M, Sun X, Wang C, Cui L, Chen L, Zhang C, Shangguan L, Fang J. 2017. Characterization of miR061 and its target genes in grapevine responding to exogenous gibberellic acid. Functional & Integrative Genomics, 17:537-549.
[46]	Wang P, Xuan X, Su Z, Wang W, Abdelrahman M, Jiu S, Zhang X, Liu Z, Wang X, Wang C, Fang J. 2021. Identification of miRNAs-mediated seed and stone-hardening regulatory networks and their signal pathway of GA-induced seedless berries in grapevine(V. vinifera L.). BMC Plant Biology, 21 (1):442-442.
[47]	Wang Sha, Cheng Dawei, Gu Hong, Li Ming, He Shasha, Li Fangfei, Gu Shichao, Chen Jinyong. 2019. Effects of plant growth regulators on the seedless rate and fruit quality of‘Shine Muscat’grape. Journal of Fruit Science, 36 (12):1675-1682. (in Chinese)
	王莎, 程大伟, 顾红, 李明, 何莎莎, 李芳菲, 谷世超, 陈锦永. 2019. 植物生长调节剂对‘阳光玫瑰’葡萄果实无核及品质的影响. 果树学报, 36 (12):1675-1682.
[48]	Wang W, Bai Y, Koilkonda P, Guan L, Fang J. 2020. Genome-wide identification and characterization of gibberellin metabolic and signal transduction(GA MST)pathway mediating seed and berry development(SBD)in grape(Vitis vinifera L.). BMC Plant Biology, 20 (1):384-384.
[49]	Wang Wenran, Wang Chen, Xie Zhenqiang, Jia Haifeng, Tang Wei, Cui Mengjie, Fang Jinggui. 2018. Function analysis of VvmiR397a and its target genes VvLACs in grape berry development. Acta Horticulturae Sinica, 45 (8):1441-1455. (in Chinese)
	王文然, 王晨, 解振强, 贾海锋, 汤崴, 崔梦杰, 房经贵. 2018. VvmiR397a及其靶基因VvLACs在葡萄果实发育中的作用分析. 园艺学报, 45 (8):1441-1455. doi: 10.16420/j.issn.0513-353x.2018-0059
[50]	Wang Wenran, Xie Zhenqiang, Zhuge Yaxian, Bai Yunhe, Guan Le, Wu Weimin, Zhang Peian, Zheng Ting, Fang Jinggui, Wang Chen. 2021. Function analysis of GA₃ mediate miR171s and its target genes VvSCLs in grape seed development. Scientia Agricultura Sinica, 54 (2):357-369. (in Chinese)
	王文然, 解振强, 诸葛雅贤, 白云赫, 管乐, 吴伟民, 张培安, 郑婷, 房经贵, 王晨. 2021. GA₃介导miR171s及其靶基因VvSCLs调控葡萄种子发育的作用分析. 中国农业科学, 54 (2):357-369. doi: 10.3864/j.issn.0578-1752.2021.02.011
[51]	Wang Xicheng, Wang Chen, Fang Jinggui, Shangguan Lingfei, Liu Hong, Han Jian. 2012. Research advance of plant hormones on the control of parthenocarpic of fruit. Plant Physiology Journal, 48 (7):629-636. (in Chinese)
	王西成, 王晨, 房经贵, 上官凌飞, 刘洪, 韩键. 2012. 植物激素对果树单性结实调控的研究进展. 植物生理学报, 48 (7):629-636.
[52]	Wang Yue. 2023. Observation and mechanism analysis of seedless formation process of grape fruit in‘Shine Muscat’by GA₃and SM[M. D. Dissertation]. Yangzhou: Yangzhou University. (in Chinese)
	王悦. 2023. GA₃和SM对‘阳光玫瑰’葡萄果实无核形成过程观察及机理分析[硕士论文]. 扬州: 扬州大学.
[53]	Xuan Xuxian, Sheng Zilu, Xie Zhenqiang, Huang Yuqing, Gong Peijie, Zhang Chuan, Zheng Ting, Wang Chen, Fang Jinggui. 2021. Function analysis of vvi-miR172s and their target genes response to gibberellin regulation of grape berry development. Scientia Agricultura Sinica, 54 (6):1199-1217. (in Chinese) doi: 10.3864/j.issn.0578-1752.2021.06.011
	宣旭娴, 盛子璐, 解振强, 黄雨晴, 巩培杰, 张川, 郑婷, 王晨, 房经贵. 2021. vvi-miR172s及其靶基因响应赤霉素调控葡萄果实发育的作用分析. 中国农业科学, 54 (6):1199-1217. doi: 10.3864/j.issn.0578-1752.2021.06.011
[54]	Yang Yuyan, Wang Jianlou, Lu Jingyun, Qin Xudong, Li Tianzhong, Hu Jianfang. 2012. The relationship between grape fruit development and gibberellin(GA₃)treatment as well as VvSEP3 gene expression in‘Fenghou’grapes. Journal of China Agricultural University, 17 (1):72-79. (in Chinese)
	杨玉艳, 王建楼, 鲁井云, 秦旭冬, 李天忠, 胡建芳. 2012. ‘峰后’葡萄果实发育与赤霉素(GA₃)处理以及VvSEP3基因表达的关系. 中国农业大学学报, 17 (1):72-79.
[55]	Zhang S L, Wang L, Yao J, Wu N, Ahmad B, Nocker S V, Wu J U, Abudureheman R, Li Z, Wang X. 2023. Control of ovule development in Vitis vinifera by VvMADS28 and interacting genes. Horticulture Research, 10 (6):uhad070.
[56]	Zhang Wenying, Han Xu, Zhu Xudong, Xie Zhenqiang, Jiu Songtao, Huang Yuqing, Jia Haifeng, Fang Jinggui, Wang Chen. 2019a. Identification of the target genes of VvmiR159s and their regulation in response to GA in different tissues of grape berry. Scientia Agricultura Sinica, 52 (16):2858-2870. (in Chinese)
	张文颖, 韩旭, 朱旭东, 解振强, 纠松涛, 黄雨晴, 贾海锋, 房经贵, 王晨. 2019a. 葡萄miR159s靶基因的鉴定及其应答GA在果实不同组织的调控作用. 中国农业科学, 52 (16):2858-2870.
[57]	Zhang Wenying, Wang Chen, Zhu Xudong, Ma Chao, Wang Wenran, Leng Xiangpeng, Zheng Ting, Fang Jinggui. 2018. Genome-wide identification and expression of DELLA protein gene family during the development of grape berry induced by exogenous GA. Scientia Agricultura Sinica, 51 (16):3130-3146. (in Chinese) doi: 10.3864/j.issn.0578-1752.2018.16.009
	张文颖, 王晨, 朱旭东, 马超, 王文然, 冷翔鹏, 郑婷, 房经贵. 2018. 葡萄全基因组DELLA蛋白基因家族鉴定及其应答外源赤霉素调控葡萄果实发育的特征. 中国农业科学, 51 (16):3130-3146. doi: 10.3864/j.issn.0578-1752.2018.16.009
[58]	Zhang Wenying, Zhu Xudong, Cui Tengfei, Jia Haifeng, Fang Jinggui, Wang Chen. 2019b. VvRGL genes regulate seedless fruit development by responding to GA in grapevine. Acta Botanica Boreali-Occidentalia Sinica, 39 (3):381-390. (in Chinese)
	张文颖, 朱旭东, 崔腾飞, 贾海锋, 房经贵, 王晨. 2019b. 葡萄VvRGL基因应答GA调控无核果实发育的研究. 西北植物学报, 39 (3):381-390.
[59]	Zhang Ying, Bai Xianjin, Cao Xiongjun, Lin Ling, Li Hongyan, Che Jianglü, Wen Rende. 2013. Effects of four plant growth regulators on inducing seedless fruit and quality of Muscat Hamburg grape. Journal of Southern Agriculture, 44 (5):819-822. (in Chinese)
	张瑛, 白先进, 曹雄军, 林玲, 李洪艳, 车江旅, 文仁德. 2013. 四种植物生长调节剂对玫瑰香葡萄无核化及果实品质的影响. 南方农业学报, 44 (5):819-822.
[60]	Zhao Maoxiang, Zheng Weiwei, Javed H U, Wang Shiping. 2020. Efficiency and mechanism of seedless berrying induced by streptomycin and gibberellins in grape (Vitis vinifera). Journal of Fruit Science, 37 (6):830-837. (in Chinese)
	赵茂香, 郑伟尉, Hafiz Umer Javed, 王世平. 2020. 链霉素和赤霉素诱导欧亚种葡萄‘玫瑰香’无核结实的效果及机制研究. 果树学报, 37 (6):830-837.

途径 Pathway	基因名称 Gene name	表达模式 Expression pattern	主要调控作用 Major regulatory effect	参考文献 Reference
赤霉素合成代谢 GAs metabolism	VvGA2oxs VvGA3oxs VvGA20oxs	下调Down 下调Down 上调Up	抑制内源赤霉素合成，降低内源赤霉素活性水平 Inhibit endogenous GAs synthesis and decrease the level of endogenous active GAs	Bai et al.，2022
赤霉素信号转导 GAs signal transduction	VvGAI1 VvRGA VvSLR1	下调Down	赤霉素信号转导通路的保守抑制因子，通过应答赤霉素信号调控果皮、果肉和种子的发育 Conservative inhibitors of GAs signaling pathway，regulate the development of berry pericarp, berry flesh and seed by responding to GAs	张文颖等，2018
赤霉素信号转导 GAs signal transduction	VvRGL	下调Down	GRAS转录因子家族成员，参与赤霉素信号传导和种子胚乳的发育 GRAS transcription factor family members，involved in GAs signaling and seed endosperm development	张文颖等，2019b
	VvGID1	下调Down	感知和接收赤霉素信号，参与赤霉素介导的多激素信号网络 Perceive and receive GAs signal，participate in GAs-mediated multi hormone signaling network	Suehiro et al.，2019
生长素途径 Auxin pathway	VvIAA9 VvARF7	下调Down	葡萄坐果起始的负调控因子，联系生长素信号与赤霉素信号 The negative regulatory factors of fruit setting initiation in grape，link the auxin signaling and GAs signaling	Jung et al.，2014
	VvARF4-2 VvARF6-1	上调Up	参与赤霉素介导的多激素信号网络，调节葡萄单性结实 Participate in GAs-mediated multi-hormone signaling network，regulate grape parthenocarpy	Sheng et al.，2022
	VvYUC VvAUX	上调Up	影响内源生长素的合成，参与赤霉素介导的生长素信号通路 Affect the synthesis of endogenous auxin，articipate in GAs-mediated auxin signaling network	刘静等，2023
细胞分裂素途径 Cytokinin pathway	VvAHK-1	下调Down	细胞分裂素受体蛋白，参与赤霉素介导的CTK信号通路，调节单性结实 Cytokinin receptor protein，participate in GAs-mediated CTK signaling pathway，regulate parthenocarpy	Sheng et al.，2022
细胞分裂素途径 Cytokinin pathway	VvCHK	下调Down	调控细胞分裂素生物合成，参与赤霉素介导的CK信号转导 Regulate cytokinin biosynthesis，participate in GAs-mediated CTK signaling pathway	Suehiro et al.，2019
脱落酸途径 Abscisic acid pathway	VvPYR1	下调Down	ABA信号通路受体蛋白，参与赤霉素介导的ABA信号通路，调节单性结实 Receptor proteins of ABA signaling pathway，participate in GAs-mediated ABA signaling pathway，regulate parthenocarpy	Sheng et al.，2022
乙烯途径 Ethylene pathway	VvCTR1	下调Down	乙烯信号途径关键基因，参与赤霉素介导的ETH信号通路，调节单性结实 The key gene of ethylene signaling pathway，participate in GAs-mediated ETH signaling pathway，regulate parthenogenesis	Sheng et al.，2022
乙烯途径 Ethylene pathway	VvACO	上调Down	乙烯生物合成相关，参与赤霉素介导的ETH信号通路 Ethylene biosynthesis related，participate in GAs-mediated ETH signaling pathway	Suehiro et al.，2019
茉莉酸途径 Jasmonic acid pathway	VvTGA2.2	下调Down	参与SA信号通路，响应赤霉素调节葡萄单性结实 Participate in SA signaling pathway and regulate grape parthenocarpy in response to GAs	Sheng et al.，2022

途径 Pathway	基因名称 Gene name	表达模式 Expression pattern	主要调控作用 Major regulatory effect	参考文献 Reference
赤霉素合成代谢 GAs metabolism	VvGA2oxs VvGA3oxs VvGA20oxs	下调Down 下调Down 上调Up	抑制内源赤霉素合成，降低内源赤霉素活性水平 Inhibit endogenous GAs synthesis and decrease the level of endogenous active GAs	Bai et al.，2022
赤霉素信号转导 GAs signal transduction	VvGAI1 VvRGA VvSLR1	下调Down	赤霉素信号转导通路的保守抑制因子，通过应答赤霉素信号调控果皮、果肉和种子的发育 Conservative inhibitors of GAs signaling pathway，regulate the development of berry pericarp, berry flesh and seed by responding to GAs	张文颖等，2018
赤霉素信号转导 GAs signal transduction	VvRGL	下调Down	GRAS转录因子家族成员，参与赤霉素信号传导和种子胚乳的发育 GRAS transcription factor family members，involved in GAs signaling and seed endosperm development	张文颖等，2019b
	VvGID1	下调Down	感知和接收赤霉素信号，参与赤霉素介导的多激素信号网络 Perceive and receive GAs signal，participate in GAs-mediated multi hormone signaling network	Suehiro et al.，2019
生长素途径 Auxin pathway	VvIAA9 VvARF7	下调Down	葡萄坐果起始的负调控因子，联系生长素信号与赤霉素信号 The negative regulatory factors of fruit setting initiation in grape，link the auxin signaling and GAs signaling	Jung et al.，2014
	VvARF4-2 VvARF6-1	上调Up	参与赤霉素介导的多激素信号网络，调节葡萄单性结实 Participate in GAs-mediated multi-hormone signaling network，regulate grape parthenocarpy	Sheng et al.，2022
	VvYUC VvAUX	上调Up	影响内源生长素的合成，参与赤霉素介导的生长素信号通路 Affect the synthesis of endogenous auxin，articipate in GAs-mediated auxin signaling network	刘静等，2023
细胞分裂素途径 Cytokinin pathway	VvAHK-1	下调Down	细胞分裂素受体蛋白，参与赤霉素介导的CTK信号通路，调节单性结实 Cytokinin receptor protein，participate in GAs-mediated CTK signaling pathway，regulate parthenocarpy	Sheng et al.，2022
细胞分裂素途径 Cytokinin pathway	VvCHK	下调Down	调控细胞分裂素生物合成，参与赤霉素介导的CK信号转导 Regulate cytokinin biosynthesis，participate in GAs-mediated CTK signaling pathway	Suehiro et al.，2019
脱落酸途径 Abscisic acid pathway	VvPYR1	下调Down	ABA信号通路受体蛋白，参与赤霉素介导的ABA信号通路，调节单性结实 Receptor proteins of ABA signaling pathway，participate in GAs-mediated ABA signaling pathway，regulate parthenocarpy	Sheng et al.，2022
乙烯途径 Ethylene pathway	VvCTR1	下调Down	乙烯信号途径关键基因，参与赤霉素介导的ETH信号通路，调节单性结实 The key gene of ethylene signaling pathway，participate in GAs-mediated ETH signaling pathway，regulate parthenogenesis	Sheng et al.，2022
乙烯途径 Ethylene pathway	VvACO	上调Down	乙烯生物合成相关，参与赤霉素介导的ETH信号通路 Ethylene biosynthesis related，participate in GAs-mediated ETH signaling pathway	Suehiro et al.，2019
茉莉酸途径 Jasmonic acid pathway	VvTGA2.2	下调Down	参与SA信号通路，响应赤霉素调节葡萄单性结实 Participate in SA signaling pathway and regulate grape parthenocarpy in response to GAs	Sheng et al.，2022

miRNA		靶基因 Target		调节作用 Regulation	参考文献 Reference
名称 Name	赤霉素应答 GAs response	名称 Name	赤霉素应答 GAs response	调节作用 Regulation	参考文献 Reference
VvmiR061	上调Up	VvREV VvHOX32	下调Down	调节开花和果实发育 Regulate flowering and fruit development	Wang et al.，2017
VvmiR156a	下调Down	VvAGL80	上调Up	调节开花、单性结实和种皮硬化 Regulate Flowering，parthenocarpy and seed-coat hardening	Su et al.，2022
VvmiR156b/c/d	上调Up	VvSPL	下调Down	调节果实发育和成熟以及胚的早期分化 Regulate the fruit development and ripening，as well as early differentiation of embryos	Wang et al.，2016；Cui et al.，2018
VvmiR159a/b/c	上调或下调 Up or down	VvGAMYB VvMYB48 VvAIX15 VvGRAS-1	上调或下调 Up or down	GA-DELLA(SLR1)-VvmiR159c-VvGAMYB是外源赤霉素诱导单性结实关键信号传导调控模型；调控果皮与果肉的发育 GA-DELLA(SLR1)-VvmiR159c-VvGAMYB is the key signaling regulatory model of GAs-induced grapevine parthenocarpy；Regulate the development of berry pericarp and berry flesh	Wang et al.，2018；张文颖等，2019a
Vvi-miR160a/b	上调Up	VvARF18	下调Down	生长素响应因子，调控赤霉素诱导葡萄种子败育的过程 Auxin response factor，regulate the GAs-induced grape seedless berry development.	白云赫等，2020
VvmiR166s	下调Down	VvHB15	上调Up	抑制果实中的木质素合成 Inhibit lignin synthesis in fruits	Bai et al.，2023
Vvi-miR172c	上调Up	VvAP2 VvRAP2-2 VvVvRAP2-3	下调Down	调控果皮与果肉组织的发育 Regulate the development of berry pericarp and berry flesh	宣旭娴等，2021
Vvi-miR172d	下调Down	VvRAP2-1	上调Up		宣旭娴等，2021
VvmiR31-3p	上调Up	VvCCoAOMT	下调Down	木质素合成酶基因，抑制种子硬化和胚胎发育 Lignin synthase gene，inhibits seed-coat hardening and embryonic development	Wang et al.，2021
VvmiR8-5p	上调Up	VvDCAF1	下调Down
VvmiR3633a	上调Up	VvGA3ox2	下调Down	抑制内源赤霉素的合成，降低活性赤霉素水平 Inhibit the synthesis of endogenous GAs and decrease the level of active GAs	Bai et al.，2022
VvmiR397a	下调Down	VvLAC	上调Up	调节木质素生物合成，影响葡萄果核的发育 Regulate the biosynthesis of lignin，affect the development of grape fruit pits	王文然等，2018

miRNA		靶基因 Target		调节作用 Regulation	参考文献 Reference
名称 Name	赤霉素应答 GAs response	名称 Name	赤霉素应答 GAs response	调节作用 Regulation	参考文献 Reference
VvmiR061	上调Up	VvREV VvHOX32	下调Down	调节开花和果实发育 Regulate flowering and fruit development	Wang et al.，2017
VvmiR156a	下调Down	VvAGL80	上调Up	调节开花、单性结实和种皮硬化 Regulate Flowering，parthenocarpy and seed-coat hardening	Su et al.，2022
VvmiR156b/c/d	上调Up	VvSPL	下调Down	调节果实发育和成熟以及胚的早期分化 Regulate the fruit development and ripening，as well as early differentiation of embryos	Wang et al.，2016；Cui et al.，2018
VvmiR159a/b/c	上调或下调 Up or down	VvGAMYB VvMYB48 VvAIX15 VvGRAS-1	上调或下调 Up or down	GA-DELLA(SLR1)-VvmiR159c-VvGAMYB是外源赤霉素诱导单性结实关键信号传导调控模型；调控果皮与果肉的发育 GA-DELLA(SLR1)-VvmiR159c-VvGAMYB is the key signaling regulatory model of GAs-induced grapevine parthenocarpy；Regulate the development of berry pericarp and berry flesh	Wang et al.，2018；张文颖等，2019a
Vvi-miR160a/b	上调Up	VvARF18	下调Down	生长素响应因子，调控赤霉素诱导葡萄种子败育的过程 Auxin response factor，regulate the GAs-induced grape seedless berry development.	白云赫等，2020
VvmiR166s	下调Down	VvHB15	上调Up	抑制果实中的木质素合成 Inhibit lignin synthesis in fruits	Bai et al.，2023
Vvi-miR172c	上调Up	VvAP2 VvRAP2-2 VvVvRAP2-3	下调Down	调控果皮与果肉组织的发育 Regulate the development of berry pericarp and berry flesh	宣旭娴等，2021
Vvi-miR172d	下调Down	VvRAP2-1	上调Up		宣旭娴等，2021
VvmiR31-3p	上调Up	VvCCoAOMT	下调Down	木质素合成酶基因，抑制种子硬化和胚胎发育 Lignin synthase gene，inhibits seed-coat hardening and embryonic development	Wang et al.，2021
VvmiR8-5p	上调Up	VvDCAF1	下调Down
VvmiR3633a	上调Up	VvGA3ox2	下调Down	抑制内源赤霉素的合成，降低活性赤霉素水平 Inhibit the synthesis of endogenous GAs and decrease the level of active GAs	Bai et al.，2022
VvmiR397a	下调Down	VvLAC	上调Up	调节木质素生物合成，影响葡萄果核的发育 Regulate the biosynthesis of lignin，affect the development of grape fruit pits	王文然等，2018

基因名称 Gene name	作用 Effect	参考文献 Reference
VvAGL11	响应赤霉素诱导影响种胚的正常发育 Affect the normal development of seed embryo in response to GAs induction	崔梦杰等，2019
VvAGL15	直接调节赤霉素代谢相关酶基因的表达，参与体细胞胚胎发生和种子萌发 Directly regulate the expression of GAs metabolism related enzyme genes and participated in somatic embryogenesis and seed germination	Wang et al.，2004
VvSEP3	参与胚珠组织种皮发育 Participate in seed formation and development	杨玉艳等，2012
VvLEC1	响应赤霉素诱导无核果实发育 Induce seedless fruit development in response to GAs	Cui et al.，2020
RBOH	调控细胞活性氧水平，抑制胚的发育 Regulate cellular reactive oxygen species levels and inhibiting embryonic development	Cheng et al.，2013
VvFeSOD3/VvMnSOD/ VvNADPH-E/VvFeRO2	抗氧化酶基因，影响抗氧化酶VvCAT和VvSOD的活性 Antioxidant enzyme genes，affect the activity of antioxidant enzymes VvCAT and VvSOD	Bai et al.，2022
VvGIN1	液泡转化酶基因，促进葡萄坐果 Vacuole converting enzyme gene，promote grape fruit setting	卢龙，2016