阿克苏地区“天红2号”苹果花芽分化期不同器官植物内源激素的变化规律研究

Study on the Variation of Endogenous Hormones in Different Organs of“Tianhong No.2”Apple during Flower Bud Differentiation in Aksu Region

摘要

揭示阿克苏地区“天红2号”苹果花芽分化过程中内源激素的动态变化规律及其调控机制。在花芽分化的四个关键时期（2024年6月20日、7月30日、9月10日和10月20日），系统测定芽、叶片、果实、当年生枝和两年生枝中吲哚乙酸（IAA）、赤霉素（GA3）、细胞分裂素（CTK）和脱落酸（ABA）的含量，并结合比值分析、相关性分析和聚类分析进行综合研究。结果表明，IAA含量整体下降，GA3呈“先升后降”趋势，CTK在中期显著升高，ABA在整个过程中持续积累；IAA/ABA与（IAA+GA3+CTK）/ABA比值随分化推进显著下降，其中果实降幅最大；相关性分析表明IAA与ABA显著负相关，GA3与CTK显著正相关，提示IAA—ABA拮抗和GA3—CTK协同是花芽分化的重要机制；聚类分析则显示芽与果实在分化后期趋于一致，反映出分生组织活性和碳水分配的相似性。“天红2号”苹果花芽分化过程中呈现出“IAA/GA3抑制作用减弱—CTK/ABA促进作用增强”的激素调控格局，为实现区域苹果稳产高质及开展外源激素调控提供理论依据。

Abstract

To reveal the dynamic changes and regulatory mechanisms of endogenous hormones during flower bud differentiation of“Tianhong No.2”apple in Aksu region．Endogenous levels of indole-3-acetic acid（IAA），gibberellin（GA₃），cytokinin（CTK），and abscisic acid（ABA）were systematically measured in buds，leaves，fruits，current-year shoots，and two-year-old shoots at four key stages of flower bud differentiation（The dates are June 20，July 30，September 10，and October 20，2024）．Ratio analysis，correlation analysis，and cluster analysis were further conducted for comprehensive evaluation．The results showed that IAA content an overall decline，GA₃ exhibited a“rise-then-fall”trend，CTK significantly increased in the mid-stage，while ABA accumulated continuously throughout the process．Both IAA/ABA and （IAA+GA₃+CTK）/ABA ratios decreased significantly with the progression of differentiation，with fruits showing the largest decline．Correlation analysis indicated a significant negative correlation between IAA and ABA，and a significant positive correlation between GA₃ and CTK，suggesting that IAA-ABA antagonism and GA₃-CTK synergy are important mechanisms in flower bud differentiation．Cluster analysis revealed that buds and fruits tended to converge at the later stage of differentiation，reflecting similar characteristics of meristem activity and carbohydrate allocation．Flower bud differentiation of“Tianhong No.2”apple is characterized by A hormonal regulatory pattern of“weakened inhibitory effect of IAA/GA₃ and enhanced promotive effect of CTK/ABA”，providing a theoretical basis for stable and high-quality apple production and exogenous hormone regulation in the region．

关键词

阿克苏 / “天红2号”苹果 / 花芽分化 / 内源激素 / 器官特异性

Key words

Aksu / “Tianhong No.2”apple / flower bud differentiation / endogenous hormones / organ specificity

参考文献

[1] Plant Hormones: Physiology,Biochemistry, and Molecular Biology[M].Springer Science & Business Media, 2010.

[2] Hanke M V, Flachowsky H, Peil A. No flower no fruit:genetic potentials to trigger flowering in fruit trees[J]. Genes Genomes Genomics, 2007, 1(1):1-20.

[3] Hwang I, Sheen J, Müller B. Cytokinin signaling networks[J]. Annual Review of Plant Biology, 2012(63):353-380.

[4] 张晶楠. 三种苹果树形的冠层结构与果实品质的相关性研究[D]. 泰安: 山东农业大学, 2010.

[5] ZHANG Jingnan. Correlation between canopy structure and fruit quality of three apple tree forms[D]. Taian: Shandong Agricultural University, 2010.

[6] 蔡勇, 李晓东, 王华. 苹果花芽分化与内源激素关系的研究[J]. 果树学报, 2014, 31(3):21-25.

[7] CAI Yong, LI Xiaodong, WANG Hua. Relationship between flower bud differentiation and endogenous hormones in apple[J]. Journal of Fruit Science, 2014, 31(3):21-25.

[8] 曾骧. 植物激素和果树的生长发育[J]. 中国果树, 1980 (S1):6-19.

[9] ZENG Xiang. Plant hormones and growth and development of fruit trees[J]. China Fruits, 1980(S1):6-19.

[10] Wang J, Luo T, Zhang H, et al. Variation of endogenous hormones during flower and leaf buds development in‘Tianhong 2’apple[J]. Hortscience, 2020, 55(11):1794- 1 798.

[11] Milyaev A, Kofler J, Moya Y A T, et al. Profiling of phytohormones in apple fruit and buds regarding their role as potential regulators of flower bud formation[J]. Tree Physiology, 2022, 42(11):2319-2 335.

[12] 阿克苏市人民政府. 阿克苏市自然地理概况[EB/OL].2023-09- 15.

[13] Aksu Municipal People's Government. General situation of Aksu natural geography[EB/OL].2023-09-15.

[14] 中国气象局新疆气象台. 新疆主要农业气候区划[M]. 乌鲁木齐: 新疆科技出版社, 2019.

[15] Xinjiang Meteorological Bureau. Main agro-climatic regionalization of Xinjiang[M]. Urumqi: Xinjiang Science and Technology Press, 2019.

[16] 周晏起, 卜庆雁. 干旱胁迫下果树内源激素变化规律研究进展[J]. 北方果树, 2011(3):1-4.

[17] ZHOU Yanqi, BU Qingyan. Advances in research on endogenous hormone changes in fruit trees under drought stress[J]. Northern Fruits, 2011(3):1-4.

[18] 王华, 刘晓东. 新疆苹果种植现状与发展趋势[J]. 中国果树, 2018(2):12-17.

[19] WANG Hua, LIU Xiaodong. Current situation and development trend of apple cultivation in Xinjiang[J]. China Fruits, 2018(2):12-17.

[20] 黄细松. 白菜开花时间相关基因的分子标记及春化相关基因的克隆和表达分析[D]. 杭州: 浙江大学, 2006.

[21] HUANG Xisong. Molecular markers of flowering time-related genes and cloning and expression analysis of vernalization-related genes in Chinese cabbage[D]. Hangzhou: Zhejiang University, 2006.

[22] Cajlachjan C, Chailakhyan M K. New facts in support of the hormonal theory of plant development[J]. C. R. Acad. Sci. URSS, 1936, 13(1):79-83.

[23] Nakajima Y, Susanto S, Hasegawa K. Effect of winter-chilling treatment on flower bud induction in young Pomelo trees[J]. Journal of the Japanese Society for Horticultural Science, 1992, 61(2):287-293.

[24] 孟繁静. 植物花发育的分子生物学[M]. 北京: 中国农业出版社, 2000.

[25] MENG Fanjing. Molecular biology of plant flower development[M]. Beijing: China Agriculture Press, 2000.

[26] 沈元月, 贾克功. 温度对桃花器官发育的影响[J]. 园艺学报, 1999, 26(1):1-6.

[27] SHEN Yuanyue, JIA Kegong. Effects of temperature on peach flower organ development[J]. Journal of Horticultural Science, 1999, 26(1):1-6.

[28] 黄胜琴, 郭建军, 叶庆生, 等. 温度对蝴蝶兰成花诱导的研究初探[J]. 中山大学学报(自然科学版), 2003, 42 (4):132-134.

[29] HUANG Shengqin, GUO Jianjun, YE Qingsheng, et al. Preliminary study on temperature-induced flowering of phalaenopsis[J]. Journal of Sun Yat-sen University(Natural Science Edition), 2003, 42(4):132-134.

[30] 阳周华, 蔡华. 苗期低温处理对孔雀草开花的影响[J]. 安徽农学通报, 2008, 14(14):118-119.

[31] YANG Zhouhua, CAI Hua. Effect of low temperature treatment at seedling stage on flowering of Tagetes erecta[J]. Anhui Agricultural Bulletin, 2008, 14(14):118-119.

[32] 李丽, 张慧. 植物激素与环境胁迫响应[M]. 北京: 科学出版社, 2018.

[33] LI Li, ZHANG Hui. Plant hormones and environmental stress responses[M]. Beijing: Science Press, 2018.

[34] 廖明安, 张光伦, 刘镕山, 等. 两类生态型区苹果器官内源激素含量及生长发育的变化[J]. 园艺学报, 1999(4):57-59.

[35] LIAO Mingan, ZHANG Guanglun, LIU Rongshan, et al. Changes in endogenous hormone contents and growth development of apple organs in two ecological types[J]. Journal of Horticultural Science, 1999(4):57-59.

[36] 邵建柱, 徐继忠, 马宝焜, 等. 苹果短枝型新品种‘Tianhong No. 2’[J]. 园艺学报, 2008, 35(1):147.

[37] SHAO Jianzhu, XU Jizhong, MA Baokun, et al. Anew spur-type apple cultivar‘Tianhong No.2’[J]. Journal of Horticultural Science, 2008, 35(1):147.

[38] 王久照, 王欢欢, 陈奇凌, 等. 不同栽培模式对‘Tianhong No. 2’苹果果实品质的影响[J]. 绿洲农业科学与工程, 2023, 9(2):64-69.

[39] WANG Jiuzhao, WANG Huanhuan, CHEN Qiling, et al. Effects of different cultivation models on fruit quality of‘Tianhong No. 2’apple[J].Oasis Agricultural Science and Engineering, 2023, 9(2):64-69.

[40] Li W F, Mao J, Su J, et al. Exogenous ABA and its inhibitor regulate flower bud induction of apple cv.‘Nagafu No.2’ grafted on different roots tocks[J]. Trees, 2021, 35:609-620.

[41] Tan M, Li G F, Chen X L, et al An N., Role of cytokinin, strigolactone,and auxin export on outgrowth of axillary buds in apple[J]. Frontiers in Plant Science, 2019, 10:616.

[42] Fan S, Zhang D, Gao C, et al. Mediation of flower induction by gibberellin and its inhibitor paclobutrazol:mRNA and miRNA integration comprises complex regulatory cross-talk in apple[J]. Plant and Cell Physiology, 2018, 59(11):2288- 2 307.

[43] Zhang S W, Zhang D, Sheng Fan S, et al. Effect of exogenous GA3 and its inhibitor paclobutrazol on floral formation,endogenous hormones,and flowering-associated genes in ‹Fuji› apple(Malus domestica Borkh.)[J].Plant Physiology and Biochemistry,2016(107):178-186.

[44] Jiang Y Y, Liu Y C, Gao Y C, et al. Gibberellin induced transcriptome profiles reveal gene regulation of loquat flowering[J]. Frontiers in Genetics, 2021(12):703+688.

[45] Li G F, Tan M, Cheng F, et al. Molecular role of cytokinin in bud activation and outgrowth in apple branching based on transcriptomic analysis[J]. Plant Molecular Biology, 2018 (98):261-274.

[46] 周学明, 马焕普, 王凤珍, 等. 不同时期苹果花芽和叶芽中内源赤霉素、脱落酸和细胞分裂素活性的变化[J]. 中国农业科学, 1988(3):41-45.

[47] ZHOU Xueming, MA Huanpu, WANG Fengzhen, et al. Changes in endogenous gibberellin,abscisic acid and cytokinin activities in apple flower and leaf buds at different stages[J]. Scientia Agricultura Sinica, 1988(3):41-45.

[48] Bharath P, Gahir S, Raghavendra A S. Abscisic acid-induced stomatal closure:an important component of plant defense against abiotic and biotic stress[J]. Frontiers in Plant Science, 2021(12):615+114.

[49] Xing L B, Zhang D, Li Y M, et al. Transcription profiles reveal sugar and hormone signaling pathways mediating flower induction in apple(Malus domestica Borkh.)[J]. Plant and Cell Physiology, 2015, 56(10):2052-2 068.

[50] Koshita Y, Takahara T, Ogata T, et al. Involvement of endogenous plant hormones(IAA,ABA,GAs)in leaves and flower bud formation of satsuma mandarin(Citrus unshiu Marc.)[J]. Scientia Horticulturae, 1999, 79(3-4):185-194.

[51] 蔡坤秀, 牛先前, 王龙平, 等. 叶底红花芽分化期叶片内源激素的动态变化[J]. 福建农业学报, 2017, 32(3):278-281.

[52] CAI KunXiu, NIU Xianqian, WANG Longping, et al. The dynamic changes of endogenous hormones in the leaves during the bud differentiation period of red flowers at the base of the leaves[J]. Fujian Journal of Agricultural Sciences, 2017, 32 (3):278-281.

[53] 章希娟, 许玲, 余东, 等. 莲雾催花过程中叶片内源激素的变化[J]. 中国农学通报, 2016, 32(34):68-71.

[54] ZHANG Xijuan, XU Ling, YU Dong, et al. Changes of endogenous hormones in leaves during flowering induction of wax apple[J]. Chinese Agricultural Bulletin, 2016, 32 (34):68-71.

[55] Huang X Z, Liu L, Qiang X J, et al. Integrative metabolomic and transcriptomic analysis elucidates that the mechanism of phytohormones regulates floral bud development in alfalfa[J]. Plants, 2024, 13(8):1 078.

[56] Zhu L C, Zhang C X, Yang N X, et al. Apple vacuolar sugar transporters regulated by MdDREB2A enhance drought resistance by promoting accumulation of soluble sugars and activating ABA signaling[J]. Horticulture Research, 2024, 11(12):251.

基金资助

新疆农垦科学院创新工程项目(NCG202403)

新疆农垦科学院创新工程项目(2025YJQN11)

新疆维吾尔自治区“2+5”人才项目(XJLGCYJSTX04-2024-18)