Transcriptome and metabolome integrated analysis of flavonoids in Xinyu grape peel under different ground mulch 5ypes

doi:10.6048/j.issn.1001-4330.2024.01.008

Abstract

Abstract:

【Objective】 To clarify the flavonoid related components and synthase genes information of table grapes under two different ground covering methods of horticultural cloth covering and grass cultivation in the hope of laying the theoretical foundation for proposing a new mode of high efficiency and high quality cultivation of Xinyu grapes in production. 【Methods】 In this study,the ground mulched by black geotextile and grass,clean tillage was taken as the control,the cDNA of Xinyu grape peel flavonoids were sequenced and were analyzed by using the bioinformatics methods subsequently,such as sequencing assess and gene function annotation,at the same time,the flavonoid metabolites in grape peel were determined by LC-MS/MS. 【Results】 (1) Compared the transcriptome of the black geotextile mulch and grass,587 differentially expressed gene(DEGs) in black geotextile mulch were found,including 317 up-regulated gene and 270 down-regulated genes,177 differentially expressed gene(DEGs) in grass were found,including 56 up-regulated gene and 121 down-regulated genes.(2) GO function annotation gene of black geotextile mulch was divided into 45 function categories,and the GO annotation gene of grass type was divided into 37 function categories,in which many function categories were mainly involved,such as cellular component,molecular function and biological processes. KOG database classified the 175 and 86 DEGs annotated to,respectively,with 22 and 16 functional categories respectively,and 193 DEGs for black geotextile mulch were annotated into KEGG pathway,biosynthesis of secondary metabolites,phenylpropanoid biosynthesis,flavonoid biosynthesis,phenylalanine biosynthesis,circadian rhythm-plant,stilbenoid,diarylheptanoid and gingerol biosynthesis were associated with flavonoid enrichment pathway,61 DEGs for grass were annotated into KEGG pathway,biosynthesis of secondary metabolites was associated with flavonoid enrichment pathway.(3) A total of 60 flavonoids metabolites were detected for Xinyu grape peel and divided into 12 categories,including 7 differential metabolites(2 down-regulated and 5 up-regulated) for black geotextile mulch and 11 differential metabolites(all down-regulated) for grass cultivation. Flavanols proportion content was the highest,with 52.41%-63.70%,flavanols content were higher for grass cultivation and black geotextile mulch than for the control,with 19.42% and 7.55%.(4) The flavonoid biosynthesis related enrichment pathways for black geotextile mulch were mainly concentrated in the flavonoid biosynthesis,while the flavonoid biosynthesis related enrichment pathways of grass mulch were mainly concentrated in the secondary metabolites and flavonoid biosynthesis,seven candidate genes were screened including C4H,CHS,F3H,LAR,HCT,FLS and C12RT1 for black geotextile mulch,five candidate genes were screened including CHS,F3H,F3'5'H,FLS and LAR for grass. 【Conclusion】 By transcriptome and metabolite analysis,candidate genes for flavonoid biosynthesis of Xinyu grape were screened and analyzed for black geotextile and grass mulch,grass cultivation was the preferable cultivation method for Xinyu grape in Xinjiang Turpan extreme drought area.

Key words: Xinyu grape; ground mulch; flavonoids; transcriptome; metabolomics

摘要：

【目的】研究园艺地布覆盖和生草栽培两种不同地面覆盖方式下鲜食葡萄新郁果皮黄酮相关组分及其合成酶基因信息,为筛选新郁葡萄高效优质栽培新模式提供科学依据。【方法】以7年生新郁葡萄为试材,以清耕栽培为对照,分别进行行间铺设园艺地布和行间生草栽培两种地面覆盖方式处理,分析不同地面覆盖方式下葡萄果皮黄酮转录组和代谢物含量的差异。【结果】(1)园艺地布覆盖获得差异表达基因587个,其中上调基因317个、下调基因270个,生草栽培获得差异表达基因177个,其中上调基因56个、下调基因121个。(2)GO数据库将园艺地布覆盖注释的基因涉及45个功能组,将生草栽培覆盖注释的基因组涉及37个功能组,均主要集中在细胞组分、分子功能和生物学过程;KOG数据库分别将其注释到的175个和86个DEGs进行直系同源分类,各自获得22个和16个功能分类,园艺地布覆盖的193个差异基因被注释到KEGG通路中,与黄酮相关富集途径有次级代谢物生物合成、苯丙醇生物合成、类黄酮生物合成、苯丙氨酸生物合成、植物昼夜节律、二苯乙烯类二芳庚类和姜辣素类的生物合成;生草栽培的61个差异基因被注释到KEGG通路中,与黄酮相关富集途径有次级代谢物生物合成。(3)新郁葡萄果皮共检测到12类60个黄酮代谢物,其中地布覆盖差异代谢物7个(下调2个,上调5个),生草覆盖差异代谢物11个(全部下调)。黄烷醇含量占总黄酮含量的52.41%~63.70%,所占比例最高,生草栽培和园艺地布栽培的黄烷醇类含量均高于对照,分别比对照提高了19.42%和7.55%。(4)与黄酮合成相关富集途径主要集中在类黄酮生物合成、黄酮和黄酮醇生物合成,园艺地布覆盖筛选出7个候选基因:C4H、CHS、F3H、LAR、HCT、FLS和C12RT1,生草筛选出5个候选基因:CHS、F3H、F3'5'H、FLS和LAR。【结论】筛选出在地布和生草栽培模式下新郁葡萄黄酮生物合成候选基因。在新疆吐鲁番极端干旱地区鲜食葡萄新郁地面覆盖方式可选择生草栽培。

关键词: 新郁葡萄, 地面覆盖, 黄酮, 转录组, 代谢组

CLC Number:

S662.1

HU Jinge, BAI Shijian, CHEN Guang, CAI Junshe. Transcriptome and metabolome integrated analysis of flavonoids in Xinyu grape peel under different ground mulch 5ypes[J]. Xinjiang Agricultural Sciences, 2024, 61(1): 63-78.

户金鸽, 白世践, 陈光, 蔡军社. 不同地面覆盖方式下新郁葡萄果皮黄酮转录组和代谢组联合分析[J]. 新疆农业科学, 2024, 61(1): 63-78.

Figures/Tables 12

References 35

[1]	骆强伟, 孙锋, 蔡军社, 等. 葡萄新品种新郁[J]. 园艺学报, 2007, 34(3):797.
	LUO Qiangwei, SUN Feng, CAI Junshe, et al. A new grape variety Xinyu[J]. Acta Horticulturae Sinica, 2007, 34(3):797.
[2]	Morimoto M, Tanimoto K, Nakano S, et al. Insect antifeedant activity of flavones and chromones against Spodoptcra litura[J]. Journal of Agriculture and Food Chemistry, 2003, 51(2):389-393. DOI URL
[3]	张波, 韩舜愈, 马腾臻, 等. 红葡萄酒中花色苷衍生物结构研究进展[J]. 食品科学, 2018, 39(5):284-295. DOI
	ZHANG Bo, HAN Shunyu, MA Tengzhen, et al. Progress in understanding structures of anthocyanins derivatives in red wines[J]. Food Science, 2018, 39(5):284-295. DOI
[4]	Cook N C, Samman S. Flavonoids-Chemistry,metabolism,cardioprotective effects,and dietary sources[J]. The Journal of Nutritional Biochemistry, 1996, 7(2):66-76. DOI URL
[5]	Springob K, Nakajima J I, Yamazaki M, et al. Recent advances in the biosynthesis and accumulation of anthocyanins[J]. Natural Product Reports, 2003, 20(3):288-303.. DOI PMID
[6]	赵德英. 梨园树盘覆盖的土壤生态效应及树体生理响应研究[D]. 北京: 中国农业科学院, 2013.
	ZHAO Deying. Study on the soil ecological effects and physiological response in different ground cover pear tree[D]. Beijing: Chinese Academy of Agricultural Sciences, 2013.
[7]	罗玲, 钟奇, 王进, 等. 不同覆盖材料对避雨栽培葡萄园土壤微生物特征及葡萄生长与品质的影响[J]. 核农学报, 2021, 35(2):471-480. DOI
	LUO Ling, ZHONG Qi, WANG Jin, et al. Influence of Different Mulching Materials on Soil Microbe and Grape Growth in Rai-Shelter Vineyard[J]. Journal of Nuclear Agricultural Sciences, 2021, 35(2):471-480. DOI
[8]	刘思, 王志磊, 张军翔. 葡萄行内覆盖对园区微域生态环境及果实品质的影响[J]. 西北农林科技大学学报(自然科学版), 2019, 47(6):73-79,88.
	LIU Si, WANG Zhilei, ZHANG Junxiang. Effects of within-row mulching on soil microsites in vineyard and fruit quality[J]. Journal of Northwest A&F University (Natural Science Edition), 2019, 47(6):73-79,88.
[9]	侯婷, 闫鹏科, 庞群虎, 等. 行内覆盖对果园土壤特性及酿酒葡萄产量和品质的影响[J]. 河南农业大学学报, 2019, 53(6):869-875.
	HOU Ting, YAN Pengke, PANG Qunhu, et al. Effects of intra-row coverage on orchard soil features and wine grape yield and quality[J]. Journal of Henan Agricultural University, 2019, 53(6):869 - 875.
[10]	王锐, 闫鹏科, 马婷慧, 等. 行内生草对土壤微环境和酿酒葡萄品质的影响[J]. 干旱地区农业研究, 2020, 38(3):195-203.
	WANG Rui, YAN Pengke, MA Tinghui, et al. Effects of intra-row planted grass on soil microenvironment and wine grape quality[J]. Agricultural Research in the Arid Areas, 2020, 38(3):195-203.
[11]	Rai A, Saito K, Yamazaki M. Integrated omics analysis of specialized metabolism in medicinal plants[J]. The Plant Journal, 2017, 90(4):764-787. DOI PMID
[12]	张少平, 邱珊莲, 张帅, 等. 紫背天葵叶片中花青素种类及合成调控基因转录组分析[J]. 西北植物学报, 2019, 39(5):808-816.
	ZHANG Shaoping, QIU Shanlian, ZHANG Shuai, et al. Transcriptome analysis of anthocyanidins and their synthesis regulatory genes in Gynura bicolor leaves[J]. Acta Botanica Boreali-Occidentalia Sinica, 2019, 39(5):808-816.
[13]	卢素文, 郑暄昂, 王佳洋, 等. 葡萄类黄酮代谢研究进展[J]. 园艺学报, 2021, 48(12):2506-2524. DOI
	LU Suwen, ZHENG Xuanang, WANG Jiayang, et al. Research progress on the metabolism of flavonoids in grape[J]. Acta Horticulturae Sinica, 2021, 48(12):2506-2524. DOI
[14]	刘伟, 王俊燚, 李萌, 等. 基于转录组测序的银杏类黄酮生物合成关键基因表达分析[J]. 中草药, 2018, 49(23):5633-5639.
	LIU Wei, WANG Junyi, LI Meng, et al. Transcriptome sequencing analysis of gene expression of flavonoid biosynthesis in Ginkgo biloba[J]. Chinese Traditional and Herbal Drugs, 2018, 49(23):5633-5639.
[15]	姚运法, 张少平, 练冬梅, 等. 黄秋葵花和果荚转录组测序及类黄酮代谢差异表达分析[J]. 西北植物学报, 2018, 38(11):2000-2009.
	YAO Yunfa, ZHANG Shaoping, LIAN Dongmei, et al. Transcriptome sequencing and differential expression analysis of flavonoid metabolism in flowers and fruits of Okra[J]. Acta Botanica Boreali-Occidentalia Sinica, 2018, 38(11):2000-2009.
[16]	陈为凯. 一年两收栽培模式下葡萄果实靶向代谢组和转录组研究[D]. 北京: 中国农业大学, 2018.
	CHEN Weikai. Study of targeted metabolome and transcriptome in grape berries grown under double cropping viticulture system[D]. Beijing: China Agricultural University, 2018.
[17]	Petrussa E, Braidot E, Zancani M, et al. Plant flavonoids-biosynthesis,transport and involvement in stress responses[J]. International Journal of Molecular Sciences, 2013, 14(7):14950-14973. DOI PMID
[18]	Gouot J C, Smith J P, Holzapfel B P, et al. Grape berry flavonoids:A review of their biochemical responses to high and extreme high temperature[J]. Journal of Experimental Botany, 2019, 70(2):397-423. DOI URL
[19]	ØM Andersen, Markham K R, ØM Andersen, et al. Flavonoids. Chemistry,biochemistry and application[J]. Pesticide Science, 2006, 7(3):223-224. DOI URL
[20]	Williams Christine A, Grayer Renée J. Anthocyanins and other flavonoids[J]. Natural Product Reports, 2004, 21(4):539-573. DOI PMID
[21]	Iwashina T. The structure and distribution of the flavonoids in plant[J]. Journal of Plant Research, 2000, 113:287-299. DOI URL
[22]	赵婷, 吴佳颖, 陈黄曌, 等. 延迟采收对酿酒葡萄类黄酮物质的影响[J]. 食品科学, 2019, 40(14):229-235. DOI
	ZHAO Ting, WU Jiaying, CHEN Huangzhao, et al. Influence of Delayed Harvest on Flavonoids Compounds of Vitis vinifera Grape[J]. Food Science, 2019, 40(14):229-235. DOI
[23]	Nilgün, Göktürk, Baydar, et al. Total phenolic contents and antibacterial activities of grape(Vitis vinifera L.) extracts[J]. Food Control, 2004, 15(5):335-339. DOI URL
[24]	Hu J G, Bai S J, Zhao R H, et al. Effects of black geotextile much and grass mulch on the microclimate,fruit quality and anthocyanin components of ‘Xinyu’ table grape[J]. New Zealand Journal of Crop and Horticultural Science, 2022,1-18.
[25]	Wang Y, Gao X, Li H, et al. Microclimate changes caused by black inter-row mulch decreased flavonoids concentrations in grapes and wines under semi-arid climate[J]. Food Chemistry, 2021, 361:130064. DOI URL
[26]	Rienth M, Vigneron N, Darriet P, et al. Grape berry secondary metabolites and their modulation by abiotic factors in a climate change scenario-a review[J]. Frontiers in Plant Science, 2021,643258.
[27]	Forkmann G. Flavonoids as flower pigments:the formation of natural spectrum and its extension by genetic engineering[J]. Plant Breeding, 1991, 106(1):1-26. DOI URL
[28]	Pelletier M K, Murrell J R, Shirley B W. Characterization of flavonol synthase and leucoanthocyanidin dioxygenase genes in Arabidopsis(further evidence for differential regulation of early and late genes[J]. Plant Physiology, 1997, 113(4):1437-1445. PMID
[29]	Quattrocchio F, Wing J F, Leppen H T C, et al. Regulatory genes controlling anthocyanin pigmentation are functionally conserved among plant species and have distinct sets of garget genes[J]. The Plant Cell, 1993, 5(11):1497-1512. DOI PMID
[30]	Kubasek W L, Shirley B W, Mckillop A, et al. Regulation of flavonoid biosynthesis genes in germinating Arabidopsis seedlings[J]. Plant Cell, 1992, 4(10):1229-1236. DOI URL
[31]	Pelletier M K, Shirley B W. Analysis of flavanone 3-hydroxylase in Arabidopsis seedling(Coordinate regulation with chalcone synthase and chalcone isomerase)[J]. Plant Physiology, 1996, 111(1):339-345. DOI PMID
[32]	戴思兰, 洪艳. 基于花青素合成和成色机理的观赏植物花色改良分子育种[J]. 中国农业科学, 2016, 49(3):529-542. DOI
	DAI Silan, HONG Yan. Molecular breeding for flower colors modification on ornamental plants based on the mechanism of anthocyanins biosynthesis and coloration[J]. Scientia Agricultura Sinica, 2016, 49(3):529-542. DOI
[33]	Das K, Roychoudhury A. Reactive oxygen species(ROS) and response of antioxidants as ROS-scavengers during environmental stress in plants[J]. Frontiers in Environmental Science, 2014, 2:53.
[34]	Gaiotti F, Pastore C, Fillippetti I, et al. Low night temperature at veraison enhances the accumulation of anthocyanins in Corvina grapes(Vitis vinifera L.)[J]. Scientific Reports, 2018, 8(1):8719. DOI
[35]	Mori K, Goto-Yamamoto N, Kitayama M, et al. Effect of high temperature on anthocyanin composition and transcription of flavonoid hydroxylase genes in ‘Pinot noir’ grapes(Vitis vinifera). The Journal of Horticulture Science and Biotechnology, 2007, 82(2):199-206.

样本ID Sample ID	原始读数 Raw reads	干净数据 Clean reads (bp)	碱基总数 Clean base (G)	错误率 Error rate (%)	测序准确度在99% Q20(%)	测序准确度在99% Q30 (%)	G和C 2种碱基占总碱基的百分比GC(%)
清耕Q-1 Clean tillage Q-1	57476100	55348990	8.30	0.03	97.00	91.94	46.03
清耕Q-2 Clean tillage Q-2	46791054	41704844	6.26	0.03	98.04	94.21	46.30
清耕Q-3 Clean tillage Q-3	47866130	41706034	6.26	0.03	98.01	94.16	46.10
园艺地布F-1 Horticultural ground cloth F-1	48420302	42003268	6.30	0.02	98.09	94.34	45.95
园艺地布F-2 Horticultural ground cloth F-2	48354116	44252400	6.64	0.03	97.89	93.91	46.46
园艺地布F-3 Horticultural ground cloth F-3	46448184	42513692	6.23	0.03	98.04	94.20	46.55
生草栽培S-1 Sod-culture-1	57847986	55154966	8.27	0.03	96.81	91.59	46.23
生草栽培S-2 Sod-culture-2	51485636	49924206	7.49	0.03	96.90	91.57	44.84
生草栽培S-3 Sod-culture-3	52146666	50057894	7.51	0.03	96.93	91.84	46.16

样本ID Sample ID	原始读数 Raw reads	干净数据 Clean reads (bp)	碱基总数 Clean base (G)	错误率 Error rate (%)	测序准确度在99% Q20(%)	测序准确度在99% Q30 (%)	G和C 2种碱基占总碱基的百分比GC(%)
清耕Q-1 Clean tillage Q-1	57476100	55348990	8.30	0.03	97.00	91.94	46.03
清耕Q-2 Clean tillage Q-2	46791054	41704844	6.26	0.03	98.04	94.21	46.30
清耕Q-3 Clean tillage Q-3	47866130	41706034	6.26	0.03	98.01	94.16	46.10
园艺地布F-1 Horticultural ground cloth F-1	48420302	42003268	6.30	0.02	98.09	94.34	45.95
园艺地布F-2 Horticultural ground cloth F-2	48354116	44252400	6.64	0.03	97.89	93.91	46.46
园艺地布F-3 Horticultural ground cloth F-3	46448184	42513692	6.23	0.03	98.04	94.20	46.55
生草栽培S-1 Sod-culture-1	57847986	55154966	8.27	0.03	96.81	91.59	46.23
生草栽培S-2 Sod-culture-2	51485636	49924206	7.49	0.03	96.90	91.57	44.84
生草栽培S-3 Sod-culture-3	52146666	50057894	7.51	0.03	96.93	91.84	46.16

差异表达基因集 Differentially expressed gene set	DEG数(个)Number of DEG			有GO功能注释的DEG数 Number of DEGs with GO function annotations	DEG总数 Total of DEG
差异表达基因集 Differentially expressed gene set	生物学过程 Biological process	细胞组分 Cellular component	分子功能 Molecular function	有GO功能注释的DEG数 Number of DEGs with GO function annotations	DEG总数 Total of DEG
清耕vs园艺地布Q_vs_F Clean tillage vs Horticultural ground cloth Q_vs_F	425	23	308	155	364
清耕vs生草栽培Q_vs_S Clean tillage vs Sod-culture Q_vs_S	119	15	132	57	108

差异表达基因集 Differentially expressed gene set	DEG数(个)Number of DEG			有GO功能注释的DEG数 Number of DEGs with GO function annotations	DEG总数 Total of DEG
差异表达基因集 Differentially expressed gene set	生物学过程 Biological process	细胞组分 Cellular component	分子功能 Molecular function	有GO功能注释的DEG数 Number of DEGs with GO function annotations	DEG总数 Total of DEG
清耕vs园艺地布Q_vs_F Clean tillage vs Horticultural ground cloth Q_vs_F	425	23	308	155	364
清耕vs生草栽培Q_vs_S Clean tillage vs Sod-culture Q_vs_S	119	15	132	57	108

代谢物ID Metabolite ID	物质 Compound	二级代谢产物 Class Ⅱ metabolite	清耕Q (对照,CK) Clean tillage		园艺地布 Horticultural ground cloth		生草栽培 Sod-culture
Flavonoid_145	原花青素B2	花青素	113.161±5.062^a (3.489%)		121.677±14.285^a (3.387%)		76.123±7.106^b (2.302%)
Flavonoid_65		查耳酮	17.546±1.872^a		18.213±2.625^a		15.875±0.745^a
Flavonoid_191	柚皮素查尔酮		0.198±0.058^b		0.345±0.048^a		0.186±0.049^b
Flavonoid_202			0.703±0.117^a		0.626±0.097^a		0.624±0.016^a
Flavonoid_22	新橙皮甙二氢查尔酮		0.023±0.017^ab		0.039±0.015^a		0.004±0.002^b
Flavonoid_31	柚皮苷二氢查尔酮		0.010±0.004^a		0.011±0.002^a		0.006±0.002^a
Flavonoid_177	总含量		0.008±0.001^b 18.848±1.946^a (0.57%)		0.018±0.004^a 19.253±2.706^a (0.54%)		0.005±0.001^b 16.700±0.683^a (0.51%)
Flavonoid_195	没食子儿茶素(-)	黄烷醇类	718.783±77.192^b		792.625±48.743^ab		903.325±68.297^a
Flavonoid_147	(-)-表儿茶素		397.187±41.006^a		455.475±48.453^a		383.830±17.308^a
Flavonoid_55	(-)-儿茶素		385.300±6.333^a		328.490±16.378^b		404.035±10.757^a
Flavonoid_84	(-)-Epigallocatechin		248.105±37.000^c		303.825±4.475^b		393.096±23.182^a
Flavonoid_150	儿茶素没食子酸酯(-)		9.069±1.240^b		10.132±0.997^b		12.403±0.370^a
Flavonoid_80	没食子儿茶素没食子酸酯(-) 总含量		6.282±1.771^b 1 764.726±137.344^b (54.41%)		7.469±0.628^b 1 898.014±115.200^ab (52.83%)		10.787±1.140^a 2 107.475±103.044^a (63.73%)
Flavonoid_176	柚皮素7-O-葡萄糖苷	二氢黄酮	2.280±0.397^a		2.186±0.387^a		1.484±0.028^b
Flavonoid_126	Eriodictyol		0.184±0.026^b		0.263±0.046^a		0.140±0.020^b
Flavonoid_78	圣草次甙		0.043±0.005^ab		0.047±0.014^a		0.026±0.006^b
Flavonoid_108	Pinocembrin		0.026±0.022^a 2.533±0.715^a (0.08%)		0.008±0.001^a 2.503±0.631^a (0.07%)		0.016±0.009^b 1.665±0.024^b (0.05%)
Flavonoid_01	落新妇苷	二氢黄酮醇	232.382±48.012^ab		262.966±16.701^a		176.495±16.215^b
Flavonoid_117	黄杞苷		24.027±6.243^b		33.018±2.330^a		16.988±1.595^b
Flavonoid_56	花旗松素		2.236±0.037^a		2.327±0.048^a		1.735±0.226^b
Flavonoid_40	二氢杨梅素		1.369±0.254^a		1.607±0.221^a		0.668±0.070^b
Flavonoid_139	二氢山奈酚		0.240±0.034^a		0.251±0.055^a		0.208±0.019^a
Flavonoid_04	异水飞蓟宾总含量		0.008±0.001^b 260.262±54.785^ab (8.03%)		0.011±0.002^a 300.181±18.845^a (8.36%)		0.000±0.000^c 196.094±17.889^b (5.93%)
Flavonoid_161	地奥司明	黄酮	82.793±9.443^b		135.293±20.719^a		43.813±5.059^c
Flavonoid_42	Nicotiflorin		13.405±2.472^b		20.647±2.846^a		6.015±0.631^c
Flavonoid_118	水仙苷		2.515±0.464^bA		4.043±0.852^a		1.741±0.480^b
Flavonoid_90	木犀草苷		2.121±0.035^a		1.932±0.411^a		1.790±0.241^a
Flavonoid_203	柠檬黄素		0.928±0.105^b		1.658±0.296^a		0.576±0.179^b
Flavonoid_165	高车前苷		0.217±0.017^a		0.201±0.026^a		0.157±0.014^b
Flavonoid_160	芹菜素7-葡萄糖苷		0.048±0.006^a		0.039±0.016^ab		0.014±0.014^b
Flavonoid_187	Tricetin		0.025±0.007^a		0.032±0.006^a		0.037±0.012^a
Flavonoid_50	木犀草素		0.007±0.002^a		0.008±0.001^a		0.009±0.004^a
Flavonoid_61	Chrysin		0.007±0.013^a		0.000±0.000^a		0.001±0.001^a
Flavonoid_06	香叶木素总含量		0.002±0.001^a 102.069±12.266^b (3.15%)		0.003±0.001^a 163.857±24.220^a (4.56%)		0.002±0.001^a 54.156±5.802^c (1.64%)
Flavonoid_02	槲皮素-3-O-葡萄糖醛酸苷	黄酮醇	705.296±59.981^ab		726.203±15.629^a		645.872±30.336^b
Flavonoid_98	金丝桃苷		84.123±16.962^ab		100.957±22.073^a		52.459±10.554^b
Flavonoid_57	芦丁		72.699±16.037^ab		86.735±11.412^a		51.383±3.789^b
Flavonoid_115	槲皮苷		42.880±9.875^a		39.482±4.102^ab		29.499±1.936^b
Flavonoid_197	异鼠李素-3-O-葡萄糖苷		35.671±4.306^b		65.029±19.704^a		19.015±4.202^b
Flavonoid_52	堪非醇3-新橙皮糖苷		5.296±1.359^a		5.693±0.679^a		2.181±0.177^b
Flavonoid_119	紫云英苷		4.158±0.833^ab		6.389±2.894^a		1.958±1.081^b
Flavonoid_194	槲皮素-7-O-β-D-葡萄糖苷		1.165±0.196^ab		1.420±0.329^a		0.878±0.168^b
Flavonoid_138	Baimaside		0.840±0.196^b		1.627±0.336^a		0.317±0.068^c
Flavonoid_54	异鼠李素-3-O-新橙皮苷		0.486±0.020^a		0.618±0.124^a		0.205±0.040^b
Flavonoid_23	槲皮素		0.326±0.043^b		0.537±0.090^a		0.285±0.018^b
Flavonoid_178	杨梅素		0.193±0.025^b		0.516±0.169^a		0.124±0.064^b
Flavonoid_92	杨梅素			0.167±0.044^b		0.518±0.199^a		0.063±0.018^b
Flavonoid_175	Afzelin		0.117±0.050^a		0.134±0.035^a		0.037±0.023^b
Flavonoid_58	异鼠李素		0.092±0.010^b		0.207±0.041^a		0.066±0.003^b
Flavonoid_188	西伯利亚落叶松黄酮		0.000±0.000^b		0.058±0.007^a		0.000±0.000^b
Flavonoid_123	2'-O-没食子酰基金丝桃苷总含量		0.000±0.000^a 953.342±39.099 ^ab (29.40%)		0.14±0.024^a 1036.137±62.291^a (28.84%)		0.000±0.000^a 804.342±39.099^b (24.32%)
Flavonoid_153	Glycitin	异黄酮	0.614±0.143^a		0.691±0.070^a		0.726±0.065^a
Flavonoid_73	Genistin		0.220±0.039^a		0.246±0.095^a		0.147±0.034^a
Flavonoid_96	毛蕊异黄酮苷		0.005±0.005^a 0.839±0.185^a (0.03%)		0.000±0.000^a 0.937±0.028^a (0.03%)		0.000±0.000^a 0.873±0.035^a (0.03%)
Flavonoid_122	7-Hydroxy-4H-chromen-4-one	酚酸类	0.396±0.116^a		0.302±0.085^a		0.395±0.050^a
Flavonoid_79	女贞苷		0.052±0.012^a		0.045±0.010^a		0.031±0.013^a
Flavonoid_35	丁香醛总含量		0.000±0.000^a 0.448±0.124^a (0.01%)		0.000±0.000^a 0.347±0.077^a (0.01%)		0.153±0.266^a 0.580±0.236^a (0.02%)
Flavonoid_157	异芒果苷	口山酮	0.024±0.004^a (0.00%)		0.021±0.001^a (0.00%)		0.021±0.001^a (0.00%)
Flavonoid_102	茶黄素	-	27.176±3.262^b (0.84%)		49.820±1.176^a (1.39%)		48.822±5.892^a (1.48%)
Flavonoid_154	黄酮碳糖苷	-	0.007±0.002^a (0.00%)		0.003±0.001^b (0.00%)		0.009±0.001^a (0.00%)