遮荫对‘花手鞠’绣球花色和花青素苷组成的影响

doi:10.16420/j.issn.0513-353x.2022-0528

摘要/Abstract

摘要：

以‘花手鞠’绣球（Hydrangea macrophylla‘Hanatemari’）为试材，研究全光照和50%遮荫对其不同开花时期花色和花青素苷组成的影响。采用英国皇家园艺学会比色卡和测色仪测定花色表型；利用高效液相色谱质谱联用技术（LC-MS）定性和定量分析花青素苷组分，并测定色素生物合成关键酶活性和金属离子含量等生理指标；运用多元线性回归法分析花色和生理指标间的关系。结果表明：与全光照相比，50%遮荫的萼片红度值（a^*）更低，黄度值（b^*）更高，在色彩上更偏粉色；从盛花期到末花期，全光照的花萼红度值（a^*）下降88.90%；而50%遮荫仅下降65.67%，有效减缓了花萼的褪色速率。花萼中共检测到矢车菊素、天竺葵素、飞燕草素、锦葵素和矮牵牛素等5种花青素苷元形成的11种花青素苷；主要组分为飞燕草-3-O-葡萄糖苷，占总花青素苷组分的92.12%，是‘花手鞠’花萼主要的呈色物质。50%遮荫有利于花青素苷的累积，其总花青素苷含量（1 286.96 μg · g^-1FW）较全光照（944.10 μg · g^-1FW）显著增加了36.32%；初开期苯丙氨酸解氨酶（PAL）活性和查尔酮异构酶（CHI）活性较高，现蕾期50%遮荫下PAL酶活性显著高于全光照；遮荫并未引起花萼中Al、Fe、Ca和Mg元素含量的显著变化。多元回归结果显示，飞燕草素、锦葵素和矮牵牛素的累积有利于绣球花萼呈现红色；总花青素苷含量对绣球花萼呈色起主导作用，并与总黄酮、总酚、PAL酶和CHI酶协同作用，共同影响花色的呈现。

关键词: 绣球, 花青素苷, 花色, 遮荫

Abstract:

The effects of full light and 50% shading on sepal color and pigment composition in different blooming periods of Hydrangea macrophylla‘Hanatemari’were studied. The color phenotype was determined by the Royal Horticultural Society colorimetric card and colorimeter. Anthocyanin composition was qualitatively and quantitatively analyzed by high-performance liquid chromatography and mass spectrometry（LC-MS）. Physiological indexes，such as the biosynthetic enzyme activities and metal ion contents，were determined. The correlation between sepal color and physiological indexes was analyzed through multiple linear regression analysis. Results showed that shading considerably affected the sepal color of H. macrophylla. The sepal color under 50% shading had lower redness value（a^*），higher yellowness value（b^*），and presented to be more pinkish（greyed-purple group）than that under full light. Fading speed of sepal color was retarded under 50% shading. From the full blooming to the withering period of flowers，the redness value a^* under full light decreased by 88.90%，whereas it only decreased by 65.67% under 50% shading. Five anthocyanidins（cyanidin，pelargonidin，delphinidin，malvidin，and petunidin）were detected in the sepal，and 11 anthocyanins were formed through glycosylation modification. Delphinium-3-O-glucoside，accounting for 92.12% of anthocyanin components，was the main reason for the coloration of hydrangea sepal. Later，50% shading was beneficial for anthocyanin accumulation，and the total anthocyanin content（1 286.96 µg · g^-1FW）was significantly increased by 36.32% compared with full light（944.10 µg · g^-1FW）. The activities of phenylalanine ammonia-lyase （PAL）and chalcone isomerase（CHI）were higher at the initiation stage，and the PAL activity was significantly higher under 50% shading at the budding stage than under full light. However，shading did not cause considerable changes in Al，Fe，Ca，and Mg contents. Multiple regression analysis showed that delphinidin，malvidin，and petunidin accumulations were beneficial to the redness of sepal color. Total anthocyanins played a dominant role in the coloration of H. macrophylla sepal under different treatments，and through synergistic effects of flavonoids，total phenols，PAL，and CHI enzyme activities，jointly affected the sepal color presentation.

Key words: Hydrangea macrophylla, anthocyanins, flower color, shade

杨君, 孔羽, 刘群录, 叶康, 秦俊. 遮荫对‘花手鞠’绣球花色和花青素苷组成的影响[J]. 园艺学报, 2023, 50(7): 1467-1481.

YANG Jun, KONG Yu, LIU Qunlu, YE Kang, QIN Jun. Effects of Shading on the Variation of Sepal Color and Pigment Composition in Hydrangea macrophylla‘Hanatemari’[J]. Acta Horticulturae Sinica, 2023, 50(7): 1467-1481.

图/表 11

参考文献 38

[1]	Cai Jian-guo, Wei Meng-qi, Zhang Yi, Wei Yun-long. 2017. Effects of shading on photosynthetic characteristics and chlorophyll fluorescence parameters in leaves of Hydrangea macrophylla. Chinese Journal of Plant Ecology, 41 (5):570-576. (in Chinese) doi: 10.17521/cjpe.2016.0245
	蔡建国, 韦孟琪, 章毅, 魏云龙. 2017. 遮荫对绣球光合特性和叶绿素荧光参数的影响. 植物生态学报, 49 (5):570-576.
[2]	Cao Yun-lin, Xing Meng-yun, Xu Chang-jie, Li Xian. 2018. Biosynthesis of flavonol and its regulation in plants. Acta Horticulturae Sinica, 45 (1):177-192. (in Chinese) doi: 10.16420/j.issn.0513-353x.2017-0306
	曹运琳, 邢梦云, 徐昌杰, 李鲜. 2018. 植物黄酮醇生物合成及其调控研究进展. 园艺学报, 45 (1):177-192. doi: 10.16420/j.issn.0513-353x.2017-0306
[3]	Cavalcanti R N, Santos D T, Meireles M A A. 2011. Non-thermal stabilization mechanisms of anthocyanins in model and food systems—an overview. Food Research International, 44 (2):499-509. doi: 10.1016/j.foodres.2010.12.007 URL
[4]	Chen G L, Chen S G, Xiao Y, Fu N L. 2018. Antioxidant capacities and total phenolic contents of 30 flowers. Industrial Crops and Products, 111:430-445. doi: 10.1016/j.indcrop.2017.10.051 URL
[5]	Chen X J, Wang P J, Gu M Y, Hou B H, Zhang C R, Zheng Y C, Sun Y, Jin S, Ye N X. 2022. Identification of PAL genes related to anthocyanin synthesis in tea plants and its correlation with anthocyanin content. Horticultural Plant Journal, 8 (3):381-394. doi: 10.1016/j.hpj.2021.12.005 URL
[6]	Dai Si-lan, Hong Yan. 2016. Molecular breeding for flower colors modification on ornamental plants based on the mechanism of anthocyanins biosynthesis and coloration studies. Scientia Agricultura Sinica, 49 (3):529-542. (in Chinese) doi: 10.3864/j.issn.0578-1752.2016.03.011
	戴思兰, 洪艳. 2016. 基于花青素苷合成和呈色机理的观赏植物花色改良分子育种. 中国农业科学, 49 (3):529-542. doi: 10.3864/j.issn.0578-1752.2016.03.011
[7]	Fukuoka N, Suzuki T, Minamide K. 2014. Effect of shading on anthocyanin and non-flavonoid polyphenol biosynthesis of Gynura bicolor leaves in midsummer. HortScience, 49 (9):1148-1153. doi: 10.21273/HORTSCI.49.9.1148 URL
[8]	Gong Zhong-xing, He Yong, Yang Jing, Song Ya, Ye Zhen-xiao, Zhu Zhu-jun. 2017. Mechanism of exogenous Al₂(SO₄)₃ on regulating the anthocyanin concentration in Hydrangea macrophylla petal. Journal of Plant Nutrition and Fertilizer, 23 (3):821-826. (in Chinese)
	龚仲幸, 何勇, 杨静, 宋亚, 叶真逍, 朱祝军. 2017. 外源硫酸铝调节八仙花花青苷组成和含量变化的分子生物学机制. 植物营养与肥料学报, 23 (3):821-826.
[9]	Guo L, Wang Y, Silva J, Fan Y, Yu X. 2019. Transcriptome and chemical analysis reveal putative genes involved in flower color change in Paeonia ‘Coral Sunset’. Plant Physiology and Biochemistry, 138:130-139. doi: 10.1016/j.plaphy.2019.02.025 URL
[10]	He F, Lin M, Yan G L, Liang N N, Pan Q H, Wang J, Reeves M J. 2010. Biosynthesis of anthocyanins and their regulation in colored grapes. Molecules, 15 (12):9057-9091. doi: 10.3390/molecules15129057 pmid: 21150825
[11]	Hong Yan, Wu Yuwei, Song Xiang, Li Mengling, Dai Silan. 2021. Molecular mechanism of light-induced anthocyanin biosynthesis in horticultural crops. Acta Horticulturae Sinica, 48 (10):1983-2000. (in Chinese) doi: 10.16420/j.issn.0513-353x.2021-0497
	洪艳, 武宇薇, 宋想, 李梦灵, 戴思兰. 2021. 光照调控园艺作物花青素苷生物合成的分子机制. 园艺学报, 48 (10):1983-2000. doi: 10.16420/j.issn.0513-353x.2021-0497
[12]	Hong Y, Yang L W, Li M L. 2016. Comparative analyses of light-induced anthocyanin accumulation and gene expression between the ray florets and leaves in chrysanthemum. Plant Physiology and Biochemistry, 103:120-132. doi: 10.1016/j.plaphy.2016.03.006 pmid: 26990403
[13]	Hu Ke, Han Ke-ting, Dai Si-lan. 2010. Regulation of plant anthocyanin synthesis and pigmentation by environmental factors. Chinese Bulletin of Botany, 45 (3):307-317. (in Chinese)
	胡可, 韩科厅, 戴思兰. 2010. 环境因子调控植物花青素苷合成及呈色的机理. 植物学报, 45 (3):307-317.
[14]	Hu Yan-ping, Zhao Di, Li Huan-ru, Feng Zhi-yi, Li Ke, Feng Su-xiang. 2021. UPLC-Q Exactive-Orbitrap high-resolution mass spectrometry quantitative analysis of the content of nine components in Ligustri Lucidi Fructus from different producing areas and before and after processing. Natural Product Research and Development, 33 (8):1308-1319. (in Chinese)
	胡雁萍, 赵迪, 李焕茹, 冯志毅, 李柯, 冯素香. 2021. UPLC-Q-Exactive Orbitrap高分辨质谱定量分析不同产地、炮制前后女贞子中9种成分的含量. 天然产物研究与开发, 33 (8):1308-1319.
[15]	Ito T, Aoki D, Fukushima K, Yoshida K. 2019. Direct mapping of hydrangea blue-complex in sepal tissues of Hydrangea macrophylla. Scientific Reports, 9:5450. doi: 10.1038/s41598-019-41968-7
[16]	Kodama M, Tanabe Y, Nakayama M. 2016. Analyses of coloration-related components in Hydrangea sepals causing color variability according to soil condition. The Horticulture Journal, 85 (4):372-379. doi: 10.2503/hortj.MI-131 URL
[17]	Li Min. 2020. Study on the molecular mechanism of petal fading of Brunfelsia acuminata and the technology of adjusting light and temperature[Ph. D. Dissertation]. Fuzhou: Fujian Agriculture and Forestry University. (in Chinese)
	李敏. 2020. 鸳鸯茉莉花瓣褪色分子机理及光温调控技术的研究[博士论文]. 福州: 福建农林大学.
[18]	Li Qing-yun, Tang Qian-wen, Chen Guan-qun, Shen Xiao-hui. 2022. Extraction,identification and physical-chemical stability of anthocyanins from two Hydrangea varieties. Guihaia,(2):1-14. (in Chinese)
	李清韵, 唐倩雯, 陈冠群, 申晓辉. 2022. 两个八仙花品种花色苷的提取、鉴定和理化稳定性. 广西植物,(2):1-14.
[19]	Lister C E, Lancaster J E. Walker J R. 1996. Developmental changes in enzymes of flavonoid biosynthesis in the skins of red and green apple cultivars. Journal of the Science of Food and Agriculture, 71 (3):313-320. doi: 10.1002/(ISSN)1097-0010 URL
[20]	Liu Jin, Wei Jing-li, Liu Mei-yan, Song Yang, Feng Shou-qian, Wang Chuan-zeng, Chen Xue-sen. 2012. The relationships between the enzyme activity of anthocyanin biosynthesis ethylene release and anthocyanin accumulation in fruits of precocious apple cultivars. Acta Horticulturae Sinica, 39 (7):1235-1242. (in Chinese)
	刘金, 魏景立, 刘美艳, 宋杨, 冯守千, 王传增, 陈学森. 2012. 早熟苹果花青苷积累与其相关酶活性及乙烯生成之间的关系. 园艺学报, 39 (7):1235-1242.
[21]	Lu Suwen, Zheng Xuanang, Wang Jiayang, Fang Jinggui. 2021. Research progress on the metabolism of flavonoids in grape. Acta Horticulturae Sinica, 48 (12):2506-2524. (in Chinese) doi: 10.16420/j.issn.0513-353x.2021-0558
	卢素文, 郑暄昂, 王佳洋, 房经贵. 2021. 葡萄类黄酮代谢研究进展. 园艺学报, 48 (12):2506-2524. doi: 10.16420/j.issn.0513-353x.2021-0558
[22]	Lubell J D, Brand M H. 2017. Flower color,color stability,and flower longevity in red-flowered elepidote rhododendrons. HortTechnology, 27 (5):607-610. doi: 10.21273/HORTTECH03792-17 URL
[23]	Mitra S, Irshad M, Debnath B, Lu X, Li M, Dash C K, Rizwan H M, Qiu Z. 2018. Effect of vineyard soil variability on chlorophyll fluorescence,yield and quality of table grape as influenced by soil moisture,grown under double cropping system in protected condition. PeerJ, 6:e5592.
[24]	Pan Yue, Zhang Xian-quan, Ye Kang, Liu Qun-lu, Qin Jun. 2021. Physiological responses of ten Hydrangea macrophylla cultivars to shading and evaluation of strong light tolerance. Journal of Fujian Agriculture and Forestry University(Natural Science Edition), 50 (1):36-48. (in Chinese)
	潘月, 张宪权, 叶康, 刘群录, 秦俊. 2021. 不同八仙花品种对遮荫和强光处理的生理响应与评价. 福建农林大学学报(自然科学版), 50 (1):36-48.
[25]	Peng J Q, Dong X J, Xue C, Liu Z M, Cao F X. 2021. Exploring the molecular mechanism of blue flower color formation in Hydrangea macrophylla cv.“forever summer”. Frontiers in Plant Science, 12:585-665.
[26]	Schreiber H D, Jones A H, Lariviere C M, Mayhew K M, Cain J B. 2011. Role of aluminum in red-to-blue color changes in Hydrangea macrophylla sepals. Biometals, 24 (6):1005-1015. doi: 10.1007/s10534-011-9458-x pmid: 21584711
[27]	Toyama-Kato Y, Yoshida K, Fujimori E, Haraguchi H, Shimizu Y, Kondo T. 2003. Analysis of metal elements of Hydrangea sepals at various growing stages by ICP-AES. Biochemical Engineering Journal, 14:237-241. doi: 10.1016/S1369-703X(02)00220-6 URL
[28]	Xi Cong-fang, Zhen Li, Zi Shu-hui, Li Jian-bing, Chen Shu-ying, Wang You-guo. 2013. Effects of different shading on anthocyanins in the petals of Camellia reticulata. Acta Horticulturae Sinica, 40 (10):2006-2014. (in Chinese)
	希从芳, 郑丽, 字淑慧, 李建宾, 陈疏影, 王有国. 2013. 不同遮阴处理对滇山茶花瓣花青素苷构成的影响. 园艺学报, 40 (10):2006-2014.
[29]	Xu Hui, Liu Chao, Zhong Han-dong. 2014. Effects of different light intensity on the blossom of Hydrangea macrophylla. Northern Horticulture,(1):81-82. (in Chinese)
	徐慧, 刘超, 钟汉东. 2014. 不同光照强度对八仙花开花的影响. 北方园艺,(1):81-82.
[30]	Yamagishi M. 2022. High temperature enhances anthocyanin coloration in Asiatic hybrid lily flowers via upregulation of the MYB 12 positive regulator. Horticultural Plant Journal, 8 (6):769-776. doi: 10.1016/j.hpj.2022.05.003 URL
[31]	Yang Jun-feng, Shi Wen-jun, Yang Le, Gong Zhong-zhi, Li Bin-bin, Hou Zhi-xia. 2016. Effects of UV irradiation on accumulation of anthocyanins and the activity,expression of key enzyme in veraison‘Northland’blueberry. Acta Horticulturae Sinica, 43 (4):663-673. (in Chinese) doi: 10.16420/j.issn.0513-353x.2016-0036
	杨俊枫, 史文君, 杨乐, 宫中志, 李彬彬, 侯智霞. 2016. 紫外光对‘北陆’越橘转色果花青苷积累、关键酶活性及其基因表达的影响. 园艺学报, 43 (4):663-673. doi: 10.16420/j.issn.0513-353x.2016-0036
[32]	Yang Ling. 2007. The influence of environment factors to the growth and factors to the growth and the color expression of Cercis canadensis‘Forest Pansy’[M. D. Dissertation]. Harbin:Northeast Forestry University (in Chinese)
	杨羚. 2007. 环境因子对紫叶加拿大紫荆生长及叶色变化的影响[硕士论文]. 哈尔滨: 东北林业大学.
[33]	Yin Hantai, Yin Junmei, Liao Yi, Lu Shunjiao, Li Chonghui. 2021. Phenotype classification based on flower color,pigment distribution and epidermal cell shape of Dendrobium hybrids. Acta Horticulturae Sinica, 48 (10):1907-1920. (in Chinese) doi: 10.16420/j.issn.0513-353x.2021-0406
	殷涵泰, 尹俊梅, 廖易, 陆顺教, 李崇晖. 2021. 基于秋石斛花朵颜色,色素分布及表皮细胞形态的表型分类. 园艺学报, 48 (10):1907-1920. doi: 10.16420/j.issn.0513-353x.2021-0406
[34]	Yoshida K, Mori M, Kondo T. 2009. Blue flower color development by anthocyanins: from chemical structure to cell physiology. Natural Product Reports, 26 (7):884-891. doi: 10.1039/b800165k pmid: 19554240
[35]	Yue Gao-feng, Han Zhi-qiang, Ma Jun-qiang, Xu Xian-xin, Yang Wen-yue. 2020. Effects of different shading treatments on flowering period and color quality of peony. Hubei Agricultural Sciences, 59 (17):83-86. (in Chinese)
	岳高峰, 韩志强, 马俊强, 许显鑫, 杨文月. 2020. 不同遮阴处理对牡丹花期及花色品质的影响. 湖北农业科学, 59 (17):83-86.
[36]	Zhao Bing, Zhang Dong-lin, Li Hou-hua. 2016. Cultivation and management. Chinese Hydrangea. Beijing: China Forestry Publishing House. (in Chinese)
	赵冰, 张冬林, 李厚华. 2016. 栽培管理技术. 中国八仙花. 北京: 中国林业出版社.
[37]	Zhao D, Hao Z, Tao J. 2012. Effects of shade on plant growth and flower quality in the herbaceous peony(Paeonia lactiflora Pall.). Plant Physiology and Biochemistry, 61:187-196. doi: 10.1016/j.plaphy.2012.10.005 URL
[38]	Zhi Wei-te, Zhao Chang-ling, Chen Zhong-jian, Liao Kang-ru, Chen Wen-long, Mao Le-xin. 2012. Determination of the higher plant organ colors by anthocyanins and its influence factors. Journal of Tropical and Subtropical Botany, 20 (3):303-310. (in Chinese)
	支伟特, 赵昶灵, 陈中坚, 缪康汝, 陈文龙, 毛乐心. 2012. 花色苷对高等植物器官颜色的决定及其影响因素. 热带亚热带植物学报, 20 (3):303-310.

处理 Treatment	时期 Stage	RHSCC	CIE Lab表色系统 CIE Lab coordinate
处理 Treatment	时期 Stage	RHSCC	L^*	a^*	b^*	C	h
全光照 Full sunlight	现蕾期 Budding stage 初开期 Initiation stage 初开末期 Final initiation stage 盛花期 Full bloom stage 盛花末期 Final full bloom stage 末花期 Final flowering stage	150C N74D 73A N74C 73B 73D	49.59 ± 7.13 d 70.30 ± 5.22 b 58.19 ± 4.11 c 59.77 ± 0.94 c 61.71 ± 5.35 c 76.23 ± 3.68 a	-3.02 ± 2.59 d -4.38 ± 4.63 d 26.48 ± 3.23 a 26.40 ± 1.57 a 20.96 ± 2.42 b 2.93 ± 1.33 c	22.95 ± 10.08 a 13.20 ± 5.68 b 5.68 ± 2.88 c -5.72 ± 1.72 d -5.87 ± 1.51 d 16.22 ± 3.54 b	23.20 ± 10.27 ab 14.26 ± 6.45 c 27.16 ± 3.68 a 27.07 ± 1.78 a 21.78 ± 2.64 ab 16.56 ± 3.39 bc	-0.96 ± 1.18 c -0.25 ± 0.29 b 0.20 ± 0.08 b -0.21 ± 0.06 b -0.27 ± 0.05 b 1.38 ± 0.11 a
50%遮荫 50% shading	现蕾期 Budding stage 初开期 Initiation stage 初开末期 Final initiation stage 盛花期 Full bloom stage 盛花末期 Final full bloom stage 末花期 Final flowering stage	150C 55C 55C 186C 186D 56D	62.11 ± 3.95 c 76.00 ± 4.14 a 69.47 ± 7.49 b 65.06 ± 3.14 bc 65.75 ± 2.33 bc 75.43 ± 2.69 a	-5.97 ± 1.65 e -5.57 ± 1.93 e 13.14 ± 4.46 c 22.37 ± 2.43 a 17.36 ± 4.09 b 7.68 ± 3.09 d	41.89 ± 5.19 a 34.90 ± 4.12 b 6.88 ± 3.00 c -3.63 ± 2.45 d -3.98 ± 2.88 d 4.91 ± 2.62 c	42.33 ± 5.26 a 35.36 ± 4.30 b 15.31 ± 3.43 d 22.75 ± 2.63 c 17.92 ± 4.52 d 9.72 ± 1.68 e	-1.43 ± 0.03 c -1.42 ± 0.04 c 0.52 ± 0.29 a -0.15 ± 0.10 b -0.21 ± 0.12 b 0.61 ± 0.42 a

处理 Treatment	时期 Stage	RHSCC	CIE Lab表色系统 CIE Lab coordinate
处理 Treatment	时期 Stage	RHSCC	L^*	a^*	b^*	C	h
全光照 Full sunlight	现蕾期 Budding stage 初开期 Initiation stage 初开末期 Final initiation stage 盛花期 Full bloom stage 盛花末期 Final full bloom stage 末花期 Final flowering stage	150C N74D 73A N74C 73B 73D	49.59 ± 7.13 d 70.30 ± 5.22 b 58.19 ± 4.11 c 59.77 ± 0.94 c 61.71 ± 5.35 c 76.23 ± 3.68 a	-3.02 ± 2.59 d -4.38 ± 4.63 d 26.48 ± 3.23 a 26.40 ± 1.57 a 20.96 ± 2.42 b 2.93 ± 1.33 c	22.95 ± 10.08 a 13.20 ± 5.68 b 5.68 ± 2.88 c -5.72 ± 1.72 d -5.87 ± 1.51 d 16.22 ± 3.54 b	23.20 ± 10.27 ab 14.26 ± 6.45 c 27.16 ± 3.68 a 27.07 ± 1.78 a 21.78 ± 2.64 ab 16.56 ± 3.39 bc	-0.96 ± 1.18 c -0.25 ± 0.29 b 0.20 ± 0.08 b -0.21 ± 0.06 b -0.27 ± 0.05 b 1.38 ± 0.11 a
50%遮荫 50% shading	现蕾期 Budding stage 初开期 Initiation stage 初开末期 Final initiation stage 盛花期 Full bloom stage 盛花末期 Final full bloom stage 末花期 Final flowering stage	150C 55C 55C 186C 186D 56D	62.11 ± 3.95 c 76.00 ± 4.14 a 69.47 ± 7.49 b 65.06 ± 3.14 bc 65.75 ± 2.33 bc 75.43 ± 2.69 a	-5.97 ± 1.65 e -5.57 ± 1.93 e 13.14 ± 4.46 c 22.37 ± 2.43 a 17.36 ± 4.09 b 7.68 ± 3.09 d	41.89 ± 5.19 a 34.90 ± 4.12 b 6.88 ± 3.00 c -3.63 ± 2.45 d -3.98 ± 2.88 d 4.91 ± 2.62 c	42.33 ± 5.26 a 35.36 ± 4.30 b 15.31 ± 3.43 d 22.75 ± 2.63 c 17.92 ± 4.52 d 9.72 ± 1.68 e	-1.43 ± 0.03 c -1.42 ± 0.04 c 0.52 ± 0.29 a -0.15 ± 0.10 b -0.21 ± 0.12 b 0.61 ± 0.42 a

峰序号 Peak number	保留时期/min Retention time	母离子/ m/z [M⁺H]⁺	碎片离子/ m/z Fragment	推定结构 Tentative identification	含量/（μg · g^-1 FW）Content
峰序号 Peak number	保留时期/min Retention time	母离子/ m/z [M⁺H]⁺	碎片离子/ m/z Fragment	推定结构 Tentative identification	全光照 Full sunlight	50%遮荫 50% shading
1	6.58	287.0552	287.0552	矢车菊素 Cyanidin	10.97	10.14
2	6.12	419.0966	287.0552	矢车菊-3-O-阿拉伯糖苷 Cyanidin-3-O-ara	2.41^**	0.54^**
3	6.09	449.1072	287.0552	矢车菊-3-O-葡萄糖苷 Cyanidin-3-O-glucoside	8.10^**	4.21^**
4	6.24	595.1647	287.0550	矢车菊-3-O-芸香糖苷 Cyanidin-3-O-rutinoside	2.05	2.62
5	5.41	611.1591	287.0549，449.1082	矢车菊-3,5-二葡萄糖苷 Cyanidin-3,5-O-diglucoside	6.68^*	5.34^*
6	6.37	433.1121	271.0601	天竺葵-3-O-葡萄糖苷 Pelargonidin-3-O-glucoside	1.08^**	0.54^**
7	4.64	465.1021	303.0500	飞燕草-3-O-葡萄糖苷 Delphinidin-3-O-glucoside	838.53^*	1 227.95^*
8	6.98	493.1333	331.0812	锦葵-3-O-葡萄糖苷 Malvidin-3-O-glucoside	41.40^**	20.65^**
9	11.87	331.0805	287.0552，315.0500	锦葵素 Malvidin	23.13^**	8.17^**
10	7.71	479.1175	317.0657	矮牵牛素-3-O-葡萄糖苷 Petunidin-3-O-glucoside	9.40^*	6.47^*
11	9.29	317.0650	302.0422	矮牵牛素 Petunidin	0.34	0.32
合计Total					944.10^*	1 286.96^*

峰序号 Peak number	保留时期/min Retention time	母离子/ m/z [M⁺H]⁺	碎片离子/ m/z Fragment	推定结构 Tentative identification	含量/（μg · g^-1 FW）Content
峰序号 Peak number	保留时期/min Retention time	母离子/ m/z [M⁺H]⁺	碎片离子/ m/z Fragment	推定结构 Tentative identification	全光照 Full sunlight	50%遮荫 50% shading
1	6.58	287.0552	287.0552	矢车菊素 Cyanidin	10.97	10.14
2	6.12	419.0966	287.0552	矢车菊-3-O-阿拉伯糖苷 Cyanidin-3-O-ara	2.41^**	0.54^**
3	6.09	449.1072	287.0552	矢车菊-3-O-葡萄糖苷 Cyanidin-3-O-glucoside	8.10^**	4.21^**
4	6.24	595.1647	287.0550	矢车菊-3-O-芸香糖苷 Cyanidin-3-O-rutinoside	2.05	2.62
5	5.41	611.1591	287.0549，449.1082	矢车菊-3,5-二葡萄糖苷 Cyanidin-3,5-O-diglucoside	6.68^*	5.34^*
6	6.37	433.1121	271.0601	天竺葵-3-O-葡萄糖苷 Pelargonidin-3-O-glucoside	1.08^**	0.54^**
7	4.64	465.1021	303.0500	飞燕草-3-O-葡萄糖苷 Delphinidin-3-O-glucoside	838.53^*	1 227.95^*
8	6.98	493.1333	331.0812	锦葵-3-O-葡萄糖苷 Malvidin-3-O-glucoside	41.40^**	20.65^**
9	11.87	331.0805	287.0552，315.0500	锦葵素 Malvidin	23.13^**	8.17^**
10	7.71	479.1175	317.0657	矮牵牛素-3-O-葡萄糖苷 Petunidin-3-O-glucoside	9.40^*	6.47^*
11	9.29	317.0650	302.0422	矮牵牛素 Petunidin	0.34	0.32
合计Total					944.10^*	1 286.96^*

处理 Treatment	化合物 Compound	不同开花阶段Different blooming stage
处理 Treatment	化合物 Compound	现蕾期 Budding stage	初开期 Initiation stage	初开末期 Final initiation stage	盛花期 Bloom stage	盛花末期 Final bloom stage	末花期 Final flowering stage
全光照	飞燕草素Delphinidin	819.94 ± 24.26	744.45 ± 37.98	773.89 ± 121.08	1 047.50 ± 159.79	832.83 ± 18.56	812.56 ± 248.45
Full	锦葵素Malvidin	87.67 ± 6.51	73.51 ± 3.28	98.14 ± 8.40	40.83 ± 4.09	17.54 ± 0.32	69.47 ± 1.16
sunlight	矢车菊素Cyanidin	37.36 ± 1.84	30.96 ± 1.80	28.55 ± 2.15	27.74 ± 5.18	22.47 ± 0.35	34.24 ± 7.89
	矮牵牛素Petunidin	4.89 ± 0.26	8.48 ± 0.46	9.74 ± 0.68	11.90 ± 1.72	10.10 ± 2.02	13.34 ± 3.09
	天竺葵素Pelargonidin	1.67 ± 0.13	1.35 ± 0.16	1.11 ± 0.06	0.93 ± 0.46	0.57 ± 0.11	0.82 ± 0.13
	总花青素苷 Total anthocyanin content	951.53 ± 32.48	858.76 ± 39.17	911.42 ± 127.86	1 128.90 ±167.76	883.52 ± 20.67	930.43 ± 241.40
50%遮荫	飞燕草素Delphinidin	908.59 ± 26.26	635.24 ± 26.42	1 026.26 ± 67.22	2 786.80 ± 364.88	934.18 ± 32.20	1 076.65 ± 77.92
50%	锦葵素Malvidin	16.17 ± 1.34	49.30 ± 4.56	20.12 ± 1.02	52.73 ± 1.06	20.21 ± 1.22	14.42 ± 1.86
shading	矢车菊素Cyanidin	20.98 ± 0.05	31.32 ± 8.09	16.47 ± 2.73	20.97 ± 6.22	23.57 ± 1.87	23.81 ± 7.50
	矮牵牛素Petunidin	8.54 ± 0.16	10.88 ± 8.17	4.74 ± 0.19	2.59 ± 0.32	9.44 ± 0.40	4.54 ± 0.21
	天竺葵素Pelargonidin	0.45 ± 0.03	0.95 ± 0.31	0.51 ± 0.11	0.52 ± 0.08	0.44 ± 0.06	0.35 ± 0.09
	总花青素苷 Total anthocyanin content	954.72 ± 24.67	727.69 ± 43.74	1 068.10 ± 68.97	2 863.61 ± 363.67	978.84 ± 32.17	1 119.77 ± 75.14