纬度和海拔对主要苹果品种花芽分化期的影响
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摘要
花芽分化调控是苹果优质高产高效栽培的关键环节之一,准确把握花芽分化时期是精准调控的前提和基础,为探究纬度和海拔对苹果花芽分化期的影响,在陕西省杨凌示范区、甘肃省静宁县和四川省茂县3个苹果产区,用摘叶和摘果的方法研究了茂县(海拔1 425、1 680和2 050 m)、静宁(海拔1 601 m)‘长富2号’苹果,杨凌地区(海拔525 m)‘长富2号’、‘烟富6号’、‘嘎拉’和‘秦冠’苹果的花芽分化差异。结果表明:在杨凌地区花芽生理分化的时间为‘长富2号’56 d,‘烟富6号’49 d,‘嘎拉’56 d,‘秦冠’42 d。在茂县不同海拔试验点,‘长富2号’花芽生理分化期持续的时间长短为低海拔>高海拔(海拔1 425 m试验点为75 d,1 680 m为70 d,2 050 m为65 d)。‘长富2号’在不同地区,花芽分化持续时间的长短为低纬度>高纬度[茂县(31o33′N)为70 d,杨凌(34o18′N)为56 d,静宁(35o41′N)为49 d]。枝条停长时间与花芽分化密切相关,枝条停长越晚越不易形成花芽。在高纬度和高海拔地区枝条停长晚,但是花芽分化持续时间相对短。‘嘎拉’和‘长富2号’花芽分化从6月初开始至10月底分为6个时期,每个时期有明显的特征,各个时期相互交叉重叠;‘长富2号’各分化时期比‘嘎拉’开始的早,结束的晚,并且持续时间长,相对分散,认为这可能与富士苹果成花难有关。
Regulation of flower bud differentiation is one of the key ways to improve yield and quality of apple products. Flower bud differentiation was regulated by many external factors,such as latitude and altitude. In order to explore the influence of latitude and altitude on the period of apple flower bud differentiation,this research was performed in three apple producing areas,including Yangling,Shaanxi Province,Jingning,Gansu Province,and Maoxian,Sichuan Province.‘Nagafu 2'at different altitudes(1 425,1 680,and 2 050 m)from Maoxian and Jingning(1 601 m),as well as different cultivars(‘Nagafu 2',‘Yanfu 6',‘Gala',‘Qinguan')from Yangling were used to determine their flower buds physiological differentiation. De-leaves and de-fruits were used to investigate the start time of floral induction. Dissection was used to observe morphological difference during flower bud development. The results show different cultivars need different time to finish flower bud physiological differentiation:‘Nagafu 2'lasts 56 days,‘Yanfu 6'needs 49 days,‘Gala'maintains 56 days and‘Qinguan'requests 42 days. In same place,flower buds physiological differentiation lasts longer time at low altitude than that at high altitude. At the altitude of 1 425 m,it needs about 75 days,while at an altitude of 1 680 m and 2050 m,it only lasts 70 and 65 days. In different regions,flower buds differentiation of‘Nagafu 2'from the low latitude was longer than that from the high latitude,the results showed as below:Maoxian(70 d),Yangling(56 d),Jingning(49 d). Growing stop of current-year shoots indicates beginning of flower buds physiological differentiation. At high altitude and high latitude regions,the growth of branches stopped late and a duration of flower buds physiological differentiation became short. Flower bud morphological differentiation of‘Gala'and‘Nagafu 2'started from the beginning of June to the end of October,and they are classified into six periods with significantly marked features,and some periods are overlapped with each other.‘Nagafu 2'always shows an earlier start and a later ending when compared with‘Gala'.‘Nagafu 2'also shows a long differentiation time.‘Gala'shows a more concentrated differentiation time than that of‘Nagafu 2',which is believed to be related to the flower formation difficulty for Fuji apple.
Regulation of flower bud differentiation is one of the key ways to improve yield and quality of apple products. Flower bud differentiation was regulated by many external factors,such as latitude and altitude. In order to explore the influence of latitude and altitude on the period of apple flower bud differentiation,this research was performed in three apple producing areas,including Yangling,Shaanxi Province,Jingning,Gansu Province,and Maoxian,Sichuan Province.‘Nagafu 2'at different altitudes(1 425,1 680,and 2 050 m)from Maoxian and Jingning(1 601 m),as well as different cultivars(‘Nagafu 2',‘Yanfu 6',‘Gala',‘Qinguan')from Yangling were used to determine their flower buds physiological differentiation. De-leaves and de-fruits were used to investigate the start time of floral induction. Dissection was used to observe morphological difference during flower bud development. The results show different cultivars need different time to finish flower bud physiological differentiation:‘Nagafu 2'lasts 56 days,‘Yanfu 6'needs 49 days,‘Gala'maintains 56 days and‘Qinguan'requests 42 days. In same place,flower buds physiological differentiation lasts longer time at low altitude than that at high altitude. At the altitude of 1 425 m,it needs about 75 days,while at an altitude of 1 680 m and 2050 m,it only lasts 70 and 65 days. In different regions,flower buds differentiation of‘Nagafu 2'from the low latitude was longer than that from the high latitude,the results showed as below:Maoxian(70 d),Yangling(56 d),Jingning(49 d). Growing stop of current-year shoots indicates beginning of flower buds physiological differentiation. At high altitude and high latitude regions,the growth of branches stopped late and a duration of flower buds physiological differentiation became short. Flower bud morphological differentiation of‘Gala'and‘Nagafu 2'started from the beginning of June to the end of October,and they are classified into six periods with significantly marked features,and some periods are overlapped with each other.‘Nagafu 2'always shows an earlier start and a later ending when compared with‘Gala'.‘Nagafu 2'also shows a long differentiation time.‘Gala'shows a more concentrated differentiation time than that of‘Nagafu 2',which is believed to be related to the flower formation difficulty for Fuji apple.
引文
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Hoover E, Silva N D, Mcartney S,Hirst P. 2004. Bud development and floral morphogenesis in four apple cultivars. Journal of Pomology&Horticultural Science, 79(6):981-984.
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LiXue-wei, Li Bing-zhi, Liu Fu-Ting, Zhang Lin-sen, Han Ming-yu, Xu Sheng-rong, Zhang Yong-wang, Liu Hang-kong. 2016. Effect of bud cutting, twig twisting and tip removal on branches water conductivity, hormone content and flower bud formation of Fuji apple. Chinese Horticulture Abstracts,46(1):3643-3650.(in Chinese)李雪薇,李丙智,刘富庭,张林森,韩明玉,胥生荣,张永旺,刘航空.2016.刻芽、扭枝和去顶梢对富士苹果枝条导水率、激素含量和花芽形成的影响.中国园艺文摘,46(1):3643-3650.
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Zhang S, Zhang D,Fan S,Du L,Shen Y,Xing L,Li Y,Ma J,Han M. 2016. Effect of exogenous GA3 and its inhibitor paclobutrazol on floral formation, endogenous hormones,and flowering-associated genes in'Fuji'apple(Malus domestica Borkh.). Plant Physiology and Biochemistry, 107(18):178-186.
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Cao Shang-yin,Tang Yi-zu,Zhang Jun-chang. 2001. Effects of GA3 and PP333 on the apple flower bud differentiation course and contents of endogenous hormone. Journal of Fruit Science, 18(6):313-316.(in Chinese)曹尚银,汤一卒,张俊昌.2001. GA3和PP333对苹果花芽形态建成及其内源激素比例变化的影响.果树学报,18(6):313-316.
Cao Shang-yin,Zhang Qiu-ming,Wu Shun. 2003a. Advances in reaserch on the mechanism of flower-bud differentiation of fruit trees. Journal of Fruit Science, 20(5):345-350.(in Chinese)曹尚银,张秋明,吴顺.2003a.果树花芽分化机理研究进展.果树学报,20(5):345-350.
Cao Shang-yin, Zhang Qiu-ming, Zhang Jun-chang, Jiang Ai-hua. 2003b. Effects of alpeneprazole on hormone levels and flower bud differentiation of apple crown buds. Journal of Plant Physiology and Molecular Biology, 29(5):375-379.(in Chinese)曹尚银,张秋明,张俊畅,江爱华.2003b.喷施烯效唑对苹果顶芽激素水平和花芽分化的影响.植物生理与分子生物学学报,29(5):375-379.
Costes E. 2003. Winter bud content according to position in 3-year-old branching systems of'Granny Smith'apple. Ann Bot, 92(4):581-588.
Du Lisha,Qi Siyan,Ma Juanjuan,Xing Libo, Fan Sheng, Li Youmei,Zhang Dong, Zhao Caiping,Han Mingyu. 2017. Effects of de-fruit treatment on soluble sugar accumulation and expression of flowering-related genes in terminal spur buds and leaves of'Fuji'apple. Acta Horticulturae Sinica,44(4):622-632.(in Chinese)杜利莎,齐思言,马娟娟,邢利博,樊胜,李有梅,张东,赵彩平,韩明玉.2017.'长富2号'苹果短枝幼果摘除对芽叶可溶性糖积累和成花相关基因表达的影响.园艺学报,44(4):622-632.
Fan S,Zhang D,Gao C,Wan S,Lei C,Wang J,Zuo X,Dong F,Li Y,Shah K,Han M. 2018. Mediation of flower induction by gibberellin and its inhibitor paclobutrazol:mRNA and miRNA integration comprises complex regulatory cross-talk in apple. Plant and Cell Physiology,DOI:10.1093/pcp/pcy 154.
Fan S, Zhang D, Zhang L, Gao C, Xin M, Mobeen M. Li Y, Ma J, Han M. 2017. Comprehensive analysis of GASA family members in the Malus domestica genome:identification, characterization,and their expressions in response to apple flower induction. BMC Genomics, 18(1):827-846.
Fan Sheng,Lei Chao,Xin Mingzhi,Shao Hongxia, Muhammad M T,Zhang Dong,Han Mingyu. 2017. Cloning and expression analysis of the gibberellin signal transduction factor MdGAMYB in Malus. Acta Horticulturae Sinica, 44(5):817-827.(in Chinese)樊胜,雷超,辛明志,邵红霞,Muhammad M T,张东,韩明玉.2017.苹果赤霉素信号转导因子MdGAMYB的克隆和表达分析.园艺学报,44(5):817-827.
Fornara F, Montaigu A,Coupland G. 2010. Snapshot:control of flowering in Arabidopsis. Cell, 141(3):550-565.
Gao Ai-ling,Li Jian-an, Liu Ru, He Zhi-xiang, Sun Ying. 2010. Advances in research on flower bud differentiation mechanism in higher plants.Nonwood Forest Research,28(2):131-136.(in Chinese)郜爱玲,李建安,刘儒,何志祥,孙颖.2010.高等植物花芽分化机理研究进展.经济林研究,28(2):131-136.
Han Ming-yu,Li Bingzhi,Fan Chonghui, Zhang Lixin, Zhang Linsen, Zhang Dong. 2015. Heory and practice of apple development regulation in the loess plateau. Beijing:China Agriculture Press:33-35.(in Chinese)韩明玉,李丙智,范崇辉,张立新、张林森、张东.2015.黄土高原苹果发育调控理论与实践.北京:中国农业出版社:33-35.
He Dong, Peng Jin-hui, Qiu Bo, Peng Liang, Hu Ling-xue, Hu Yao, Zhao Ying-ying. 2013. Research progress on the effect of temperature on flower bud differentiation of ornamental plants. Chinese Horticulture Abstracts, 29(3):40-42.(in Chinese)何东,彭尽晖,邱波,彭亮,胡凌雪,胡瑶,赵盈盈.2013.温度对观赏植物花芽分化影响的研究进展.中国园艺文摘,29(3):40-42.
Hoover E, Silva N D, Mcartney S,Hirst P. 2004. Bud development and floral morphogenesis in four apple cultivars. Journal of Pomology&Horticultural Science, 79(6):981-984.
Huang Hai. 1987. Study on flower bud differentiation of fruit trees(continued). Journal of Fruit Science, 4(3):41-47.(in Chinese)黃海.1987.关于果树花芽分化的研究(续).果树学报,4(3):41-47.
Huang Hai,Qiao Xian-sheng,Cao Shang-yin. 1986b. Studies on physiological differentiation of apple flower buds. Acta Horticulturae Sinica, 13(3):181-186.(in Chinese)黄海,乔宪生,曹尚银.1986.关于苹果花芽生理分化时期的研究.园艺学报,13(3):181-186.
Huet J. 1973. Floral initiation in pear trees. Nuclear Instruments&Methods in Physics Research, 267(21-22):3500-3504.
Lauri P, Maguylo K, Trottier C. 2006. Architecture and size relations:an essay on the apple(Malus×domestica, Rosaceae)tree. American Journal of Botany, 93(3):357-368.
Li Xian-li, Yuan Zhi-you,Gao Dong-sheng. 2002. The research achievements on mechanism of floral formation in plants. Acta Botanica Boreali-occidentalia Sinica, 22(1):173-183.(in Chinese)李宪利,袁志友,高东升.2002.高等植物成花分子机理研究现状及展望.西北植物学报,22(1):173-183.
LiXue-wei, Li Bing-zhi, Liu Fu-Ting, Zhang Lin-sen, Han Ming-yu, Xu Sheng-rong, Zhang Yong-wang, Liu Hang-kong. 2016. Effect of bud cutting, twig twisting and tip removal on branches water conductivity, hormone content and flower bud formation of Fuji apple. Chinese Horticulture Abstracts,46(1):3643-3650.(in Chinese)李雪薇,李丙智,刘富庭,张林森,韩明玉,胥生荣,张永旺,刘航空.2016.刻芽、扭枝和去顶梢对富士苹果枝条导水率、激素含量和花芽形成的影响.中国园艺文摘,46(1):3643-3650.
Li Y, Zhang D, Zhang L, Zuo X, Fan S, Zhang X, Shalmani A, Han M. 2017. Identification and expression analysis of cytokinin response-regulator genes during floral induction in apple(Malus domestica Borkh). Plant Growth Regulation, 85(12):117-135.Luckwill L C, Silva J M. 1979. The effects of daminozide and gibberellic acid on flower initiation, growth and fruiting of apple cv golden delicious.Journal of Pomology&Horticultural Science,54(3):217-223.
Michaels S D,Amasino R M. 2000b. Memories of winter:vernalization and the competence to flower. Plant Cell&Environment, 23(11):1145-1153.
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