Hormonal and Transcriptome Analysis of Ripe Fruits of Non-Climacteric and Climacteric Melon

doi:10.16420/j.issn.0513-353x.2024-0371

Acta Horticulturae Sinica ›› 2025, Vol. 52 ›› Issue (4): 883-896.doi: 10.16420/j.issn.0513-353x.2024-0371

• Genetic & Breeding·Germplasm Resources·Molecular Biology • Previous Articles Next Articles

Hormonal and Transcriptome Analysis of Ripe Fruits of Non-Climacteric and Climacteric Melon

TANG Lingli¹^,², HE Yuhua¹, WANG Qingtao¹, AN Lulu¹, XU Yongyang¹, ZHANG Jian¹, KONG Weihu¹, HU Keyun¹, ZHAO Guangwei¹^,²^,^*()

¹ Zhengzhou Fruit Research Institute，Chinese Academy of Agricultural Sciences，Zhengzhou 450009，China
² Zhongyuan Research Center，Chinese Academy of Agricultural Sciences，Xinxiang，Henan 453500，China

Received:2024-12-18 Revised:2025-02-19 Online:2025-05-08 Published:2025-04-25
Contact: ZHAO Guangwei

Abstract

Abstract:

Melon can be divided into non-climacteric and climacteric according to mature habituation，but the most important hormones，sugar metabolites and genetic regulatory mechanisms remain elusive between non-climacteric and climacteric melon. Plant hormone metabolism detection revealed that abscisic acid（ABA），and the content of 1-aminocyclop ropanecarboxylic acid（ACC），trans-Zeatin Riboside-5′-monophosphate Disodium Salt（tZRMP）and auxin were higher in mature non-climacteric melon. The KEGG pathway enrichment analysis of these differentially expressed genes（DEGs）revealed“metabolic pathways”，“plant hormone signal transduction”and“biosynthesis of various alkaloids” were significantly different in the ripe fruits of non-climacteric and climacteric melons. The RNA-seq and qRT-PCR analyses were performed to identify important genes，enzymes and pathways that are involved in sugar and hormone metabolites. CmARF，CmARR2-like，CmGAI-like，CmPP2C，CmACO and CmEIL associated with plant hormone metabolism were identified in mature melon. ABA，Indole，ACC and GA pathways play key roles in non-climacteric melon. While besides ABA，IAA，GA，and ethylene（ETH）signaling pathways were also involved in regulating the maturation of climacteric melon.

Key words: melon, fruit ripening, phytohormone, transcriptome, gene expression

TANG Lingli, HE Yuhua, WANG Qingtao, AN Lulu, XU Yongyang, ZHANG Jian, KONG Weihu, HU Keyun, ZHAO Guangwei. Hormonal and Transcriptome Analysis of Ripe Fruits of Non-Climacteric and Climacteric Melon[J]. Acta Horticulturae Sinica, 2025, 52(4): 883-896.

Figures/Tables 9

Table 1 The qRT-PCR primers used in this study

引物名称 Primer name	引物序列（5′-3′） Primer sequence	引物名称 Primer name	引物序列（5′-3′） Primer sequence
RT-CmARF-F	GGCAGTCTTGTGTTTCGGC	RT-CmGH3.3-2-R	TCCCCATTGGCAATACGGT
RT-CmARF-R	GCTCCAGATTCGTCGATGC	RT-CmACO-F	GAGCGACCGACCATCTTAG
RT-CmARR2-like-F	GTCTACGGCCAGTTCCAGT	RT-CmACO-R	CGTTTAGTCCTTTGCTGGC
RT-CmARR2-like-R	GAGTGCATCCTCCGCTCG	RT-CmGAI-like-F	CGGAGGATCGTGAAAGGAG
RT-CmEIL-F	GCTCCACCAACGGATCCTG	RT-CmGAI-like-R	GCTCCAACTTCAAAGCGAC
RT-CmEIL-R	GCCTGGTCTTGAGACTGTCG	RT-CmACTIN-F	GCATCTCTAAGTACCTTCC
RT-CmGH3.3-2-F	TGTCACTACCAACGCCGAC	RT-CmACTIN-R	CCAACTAAAGGGAAATAAC

Fig. 1 Fruit phenotypes of the non-climacteric（P15，P45）and climacteric（X751，X757）melon FL：Fruit length/cm；FD：Fruit width/cm；SS：Soluble solids/°Bx

Fig. 2 The comparative hormone content analysis in the non-climacteric（P15，P45）and climacteric（X751，X757）melon The significant differences in hormones contents listed in P15，P45（non-climacteric melon）and X751，X757（climacteric melon），when comparing the hormone content between the two different groups. Student’s t-test，** P < 0.01

Fig. 3 KEGG enrichment blot analysis of differential metabolites of the non-climacteric（P15，P45）and climacteric（X751，X757）melon

Fig. 4 Principal component（A）and DEGs analysis（B，C）of the non-climacteric（P15，P45）and climacteric melon（X751，X757）

Fig. 5 Genes related to sugar metabolite of the non-climacteric（P15，P45）and climacteric（X751，X757）melon Comparison of gene expression between non-climacteric and climacteric fruits. Student’s t-test，∗∗ P < 0.01

Fig. 6 Hormone and gene co-expression network analysis of the non-climacteric（A）and climacteric（B）melon The network of the most highly correlated genes which were in the blue and cyan module. In this network，the corresponding topological overlap is above a threshold of 0.08

Fig. 7 DEGs involved in plant hormone signal transduction of the non-climacteric（P15，P45）and climacteric（X751，X757）melon The plant hormone signal transduction pathways were modified from the Kanehisa et al.，2023

Fig. 8 The qRT-PCR detection of hormone signaling pathway gene expression in non-climacteric（P45）and climacteric（X751）melon fruits CmACTIN was used as an internal control，and three biological replicates were calculated. Error bars indicate the standard error of the mean. * P < 0.05，** P < 0.01

References 54

[1]	Adamowski M, Friml J. 2015. PIN-dependent auxin transport:action,regulation,and evolution. The Plant Cell,27:20-32.
[2]	Banasiak A, Biedron M, Dolzblasz A, Berezowski M. 2019. Ontogenetic changes in auxin biosynthesis and distribution determine the organogenic activity of the shoot apical meristem in pin1 mutants. Int J Mol Sci,20:1-20.
[3]	Beaudry A, Severson R, Black C. 1989. Banana ripening:implications of changes in glycolytie intermediate concentrations,glycolytic and gluconeogenic carbon flux,and fructose 2,6-bisphosphate concentration. Plant Physipl,91:1436-1444.
[4]	Burger Y, Schaffer A. 2007. The contribution of sucrose metabolism enzymes to sucrose accumulation in Cucumis melo. L. Journal of the American Society for Horticultural Science, 132 (5):704-712.
[5]	Busatto N, Farneti B, Tadiello A, Oberkofler V, Cellini A, Biasioli F, Delledonne M, Cestaro A, Noutsos C, Costa F. 2019. Wide transcriptional investigation unravel novel insights of the on-tree maturation and postharvest ripening of‘Abate Fetel’pear fruit. Horticulture Research,6:32.
[6]	Cao H, Chen J, Yue M, Xu C, Jian W, Liu Y, Song B, Gao Y, Cheng Y, Li Z. 2020. Tomato transcriptional repressor MYB 70 directly regulates ethylene‐dependent fruit ripening. The Plant Journal,104:1568-1581.
[7]	Chen C, Chen H, Zhang Y, Thomas H, Frank M, He Y, Xia R. 2020. TBtools:an integrative toolkit developed for interactive analyses of big biological data. Mol Plant,13:1194-1202.
[8]	Cheng H, Kong W, Tang T, Ren K, Zhang K, Wei H, Lin T. 2022. Identification of key gene networks controlling soluble sugar and organic acid metabolism during oriental melon fruit development by integrated analysis of metabolic and transcriptomic analyses. Frontiers in Plant Science,13:830517.
[9]	Dai C, Sahar U, Portnoy V, Tzuri G, Ophir R, Sherman A, Katzir N, Schaffer A, Hovav R. 2011. Metabolism of soluble sugars in developing melon fruit. Plant Mol Biol, 76:1-18.
[10]	Dai N, Petreikov M, Portnoy V, Katzir N, Pharr D, Schaffer A. 2006. Cloning and expression analysis of a UDP-Galactose/Glucose pyrophosphorylase from melon fruit provides evidence for the major metabolic pathway of galactose metabolism in raffinose oligosaccharide metabolizing plants. Plant Physiology,142:294-304.
[11]	Diao Q, Tian S, Cao Y, Yao D, Fan H, Zhang Y. 2023. Transcriptome analysis reveals association of carotenoid metabolism pathway with fruit color in melon. Scientific Reports,13:5004.
[12]	Diaz A, Fergany M, Formisano G. 2011. A consensus linkage map for molecular markers and Quantitative Trait Loci associated with economically important traits in melon(Cucumis melo L.). BMC Plant Biol,11:111.
[13]	Dos-Santos N, Bueso M, Díaz A, Moreno E, Garcia-Mas J, Monforte A, Fernández-Trujillo J. 2023. Thorough Characterization of ETHQB3.5,a QTL involved in melon fruit climacteric behavior and aroma volatile composition. Foods, 12 (2):376.
[14]	Fan D, Wang W, Hao Q, Jia W. 2022. Do non-climacteric fruits share a common ripening mechanism of hormonal regulation? Frontiers in Plant Science,13:923484.
[15]	Fenn M A, Giovannoni J J. 2021. Phytohormones in fruit development and maturation. Plant J, 105 (2):446-458.
[16]	Fortes A, Teixeira R, Agudelo-Romero P. 2015. Complex interplay of hormonal signals during grape berry ripening. Molecules,20:9326-9343.
[17]	Ge S, Son E, Yao R. 2018. iDEP:an integrated web application for differential expression and pathway analysis of RNA-Seq data. BMC Bioinformatics, 19 (1):534.
[18]	Green M, Sambrook J. 2018. Analysis and normalization of Real-Time Polymerase Chain Reaction(PCR)experimental data. Cold Spring Harb Protoc,(10): 10.1101/pdb.top095000.
[19]	Huggett J, Dheda K, Bustin S, Zumla A. 2005. Real-time RT-PCR normalisation;strategies and considerations. Genes Immun,6:279-284.
[20]	Jiang Bo,Lü Yuanda,Liu Shumei,Yan Huaxue. 2024. Research advances in the regulation of plant hormones in citrus fruit maturation. Acta Horticulturae Sinica, 51 (12):2928-2944. (in Chinese)
	姜波, 吕远达, 刘淑梅, 闫化学. 2024. 植物激素调控柑橘果实成熟的研究进展. 园艺学报, 51 (12):2928-2944.
[21]	Kanehisa M, Furumichi M, Sato Y, Kawashima M, Ishiguro-Watanabe M. 2023. KEGG for taxonomy-based analysis of pathways and genomes. Nucleic Acids Research,51:587-592.
[22]	Karlova R, Chapman N, David K, Angenent G, Seymour G, de Maagd R. 2014. Transcriptional control of fleshy fruit development and ripening. Journal of Experimental Botany,65:4527-4541.
[23]	Kou X, Zhou J, Wu C, Yang S, Liu Y, Chai L, Xue Z. 2021. The interplay between ABA/ethylene and NAC TFs in tomato fruit ripening:a review. Plant Molecular Biology,106:223-238.
[24]	Langfelder P, Horvath S. 2008. WGCNA:an R package for weighted correlation network analysis. BMC Bioinformatics,9:559.
[25]	Leida C, Moser C, Esteras C, Sulpice R, Lunn J, de Langen F, Monforte A, Picó B. 2015. Variability of candidate genes,genetic structure and association with sugar accumulation and climacteric behavior in a broad germplasm collection of melon(Cucumis melo L.). BMC Genetics,16:28.
[26]	Li T, Dai Z, Zeng B, Li J, Ouyang J, Kang L, Wang W, Jia W. 2022. Autocatalytic biosynthesis of abscisic acid and its synergistic action with auxin to regulate strawberry fruit ripening. Horticulture Research, 9 (9):uhab076.
[27]	Li Y, He L, Song Y, Zhang P, Chen D, Guan L, Liu S. 2023. Comprehensive study of volatile compounds and transcriptome data providing genes for grape aroma. BMC Plant Biology, 23 (1):171.
[28]	Liu B, Santo Domingo M, Mayobre C, Martín-Hernández A, Pujol M, Garcia-Mas J. 2022. Knock-out of CmNAC-NOR affects melon climacteric fruit ripening. Frontiers in Plant Science,13:878037.
[29]	Liu M, Pirrello J, Chervin C, Roustan J, Bouzayen M. 2015. Ethylene control of fruit ripening:revisiting the complex network of transcriptional regulation. Plant Physiology, 169 (4):2380-2390.
[30]	Livak K, Schmittgen T. 2001. Analysis of relative gene expression data using real-time quantitative PCR and the 2^-ΔΔCT method. Methods,25:402-408.
[31]	Lü P, Yu S, Zhu N, Chen Y, Zhou B, Pan Y, Tzeng D, Fabi J, Argyris J, Garcia-Mas J. 2018. Genome encode analyses reveal the basis of convergent evolution of fleshy fruit ripening. Nature Plants,4:784-791.
[32]	Ma X C, Chang Y Y, Li F F, Yang J F, Ye L, Zhou T, Jin Y, Sheng L, Lu X P. 2024. CsABF3-activated CsSUT1pathway is implicated in pre-harvest water deficit inducing sucrose accumulation in citrus fruit. Horticultural Plant Journal, 10 (1):103-114.
[33]	Monforte J. 2013. Interaction between QTLs induces an advance in ethylene biosynthesis during melon fruit ripening. Theor Appl Genet,126:1531-1544.
[34]	Nishiyama K, Guis M, Rose J, Kubo Y, Bennett K, Wang L, Kato K, Ushijima K, Nakano R, Inaba A. 2007. Ethylene regulation of fruit softening and cell wall disassembly in Charentais melon. Journal of Experimental Botany,58:1281-1290.
[35]	Nonaka S, Ito M, Ezura H. 2023. Targeted modification of CmACO1 by CRISPR/Cas9 extends the shelf-life of Cucumis melo var. reticulatus melon. Frontiers in Genome Editing,5:1176125.
[36]	Pan L, Zeng W, Niu L,Lu,Z, Liu H, Cui G, Zhu Y, Chu J, Li W, Fang W. 2015. PpYUC11,a strong candidate gene for the stony hard phenotype in peach(Prunus persica L. Batsch),participates in IAA biosynthesis during fruit ripening. Journal of Experimental Botany,66:7031-7044.
[37]	Paul V, Pandey R, Srivastava G. 2011. The fading distinctions between classical patterns of ripening in climacteric and non-climacteric fruit and the ubiquity of ethylene-an overview. Journal of Food Science and Technology,49:1-21.
[38]	Perpiñá G, Cebolla-Cornejo J, Esteras C, Monforte A, Picó B. 2017. ‘MAK-10’:a long shelf-life charentais breeding nine developed by introgression of a genomic region from Makuwa Melon. HortScience,52:1633-1638.
[39]	Pujol M, Garcia-Mas J, Costa F. 2023. Regulation of climacteric fruit ripening in melon:recent advances and future challenges. Journal of Experimental Botany,74:6224-6236.
[40]	Shao Fengqing, Luo Xiurong, Wang Qi, Zhang Xianzhi, Wang Wencai. 2023. Advances in research of DNA methylation regulation during fruit ripening. Acta Horticulturae Sinica, 50 (1):197-208. (in Chinese)
	邵凤清, 罗秀荣, 王奇, 张宪智, 王文彩. 2023. 果实成熟过程中的DNA 甲基化调控研究进展. 园艺学报, 50(1):197-208.
[41]	Shen Q, Wu X, Tao Y, Yan G, Wang X, Cao S, Wang C, He W. 2022. Mining candidate genes related to heavy metals in mature melon(Cucumis melo L.) peel and pulp using WGCNA. Genes(Basel), 13 (10):1767.
[42]	Stepanova A, Yun J, Likhacheva A, Alonso J. 2007. Multilevel interactions between ethylene and auxin in Arabidopsis roots. The Plant Cell,19:2169-2185.
[43]	Wang J, Tian S, Yu Y, Ren Y, Guo S, Zhang J, Li M, Zhang H, Gong G, Wang M. 2022. Natural variation in the NAC transcription factor NONRIPENING contributes to melon fruit ripening. Journal of Integrative Plant Biology,64:1448-1461.
[44]	Wang X, Meng J, Deng L, Wang Y, Liu H, Yao J, Nieuwenhuizen N, Wang Z, Zeng W. 2021a. Diverse functions of IAA-leucine resistant PpILR1 provide a genic basis for auxin-ethylene crosstalk during peach fruit ripening. Frontiers in Plant Science,12:655758.
[45]	Wang X, Zeng W, Ding Y, Wang Y, Niu L, Yao J, Pan L, Lu Z, Cui G, Li G. 2019. PpERF 3 positively regulates ABA biosynthesis by activating PpNCED2/3 transcription during fruit ripening in peach. Horticulture Research, 6:19.
[46]	Wang Y, Deng L, Meng J, Niu L, Pan L, Lu Z, Cui G, Wang Z, Zeng W. 2021b. Transcriptomic and metabolic analyses reveal the mechanism of ethylene production in stony hard peach fruit during cold storage. International Journal of Molecular Sciences, 22 (21):11308.
[47]	Xu M, Zhou W, Geng W, Zhao S, Pan Y, Fan G, Zhang S, Wang Y, Liao K. 2021. Transcriptome analysis insight into ethylene metabolism and pectinase activity of apricot(Prunus armeniaca L.) development and ripening. Scientific Reports, 11 (1):13569.
[48]	Yang T, Deng L, Wang Q, Sun C, Ali M, Wu F, Zhai H, Xu Q, Xin P, Cheng S. 2024. Tomato CYP94C 1 inactivates bioactive JA-Ile to attenuate jasmonate-mediated defense during fruit ripening. Molecular Plant,17:509-512.
[49]	Yin H N, Wang Z X, Wang L, Cao J H, Wang J K, Xi Z M. 2024. Effects of mesoclimate and microclimate variations mediated by high altitude and row orientation on sucrose metabolism and anthocyanin synthesis in grape berries. Horticultural Plant Journal, 10 (3):713-731.
[50]	Young M D, Wakefield M J, Smyth G K, Oshlack A. 2010. Gene ontology analysis for RNA-seq accounting for selection bias. Genome Biology, 11 (2):R14.
[51]	Zhang Y, Su Z, Luo L, Wang P, Zhu X, Liu J, Wang C. 2023. Exogenous auxin regulates the growth and development of peach fruit at the expansion stage by mediating multiple-hormone signaling. BMC Plant Biology, 23 (1):499.
[52]	Zhang Yin,Hu Luyan,Wang Shuming,Jing Danlong,Guo Qigao,Liang Guolu. 2023. Research advances in ABA-mediated fruit ripening. Acta Horticulturae Sinica, 50 (9):1889-1898. (in Chinese)
	张印, 胡路艳, 王淑明, 景丹龙, 郭启高, 梁国鲁. 2023. ABA调控果实成熟研究进展. 园艺学报, 50 (9):1889-1898.
[53]	Zhao G, Lian Q, Zhang Z, Fu Q, He Y, Ma S, Ruggieri V, Monforte A, Wang P, Julca I. 2019. A comprehensive genome variation map of melon identifies multiple domestication events and loci influencing agronomic traits. Nat Genet,51:1607-1615.
[54]	Zhong S, Fei Z, Chen Y, Zheng Y, Huang M, Vrebalov J, McQuinn R, Gapper N, Liu B, Xiang J. 2013. Single-base resolution methylomes of tomato fruit development reveal epigenome modifications associated with ripening. Nature Biotechnology,31:154-159.

Hormonal and Transcriptome Analysis of Ripe Fruits of Non-Climacteric and Climacteric Melon

RichHTML

PDF

Knowledge

Abstract

Cite this article

share this article

Figures/Tables 9

References 54

Related Articles 15

Recommended Articles

Metrics

Comments

[1]	LIU Weisheng, LIU Shuai, SUN Julong, BAI Zhigang, ZHANG Yangdong, ZHANG Tao, CUI Aihua, HU Qixing. A New Mid-Early Maturing Watermelon Cultivar‘Cuijia 76’ [J]. Acta Horticulturae Sinica, 2025, 52(S1): 147-148.
[2]	YU Rong, TIAN Mei, YANG Wanbang, GUO Song, LIU Shengfeng, DU Huiying. A New Watermelon Cultivar‘Ningnongke 10’ [J]. Acta Horticulturae Sinica, 2025, 52(S1): 149-150.
[3]	CUI Aihua, ZHANG Dezheng, SUN Julong, BAI Zhigang, HU Qixing, LIU Weisheng, ZHANG Yangdong, LIU Shuai, XIAO Yuanlong, XIAHOU Wei, WU Yisheng. A New Watermelon Cultivar‘Cuimi’ [J]. Acta Horticulturae Sinica, 2025, 52(S1): 151-152.
[4]	TIAN Hongmei, WANG Yuanlong, WANG Fei, PAN Hong, TAO Zhen, ZHANG Jian, WANG Pengcheng. A New Muskmelon Cultivar‘Jinzhongyuan Jintian’ [J]. Acta Horticulturae Sinica, 2025, 52(S1): 153-154.
[5]	HE Yuhua, XU Yongyang, TANG Lingli, HU Keyun, KONG Weihu, ZHANG Jian, ZHAO Guangwei. A New Melon Cultivar‘Zhongtian Hongxiaoxian’ [J]. Acta Horticulturae Sinica, 2025, 52(S1): 155-156.
[6]	WANG Yi, LIANG Zhenchang. Analysis of Genetic Mechanism of Grape Distant Hybridization Based on Population Transcriptome [J]. Acta Horticulturae Sinica, 2025, 52(4): 835-845.
[7]	YUAN Juanwei, JIA Li, WANG Han, YAN Congsheng, ZHANG Qi’an, YU Feifei, GAN Defang, JIANG Haikun. Screening of Resistant Germplasm and Identifying Candidate Gene for Phytophthora capsici Resistance in Pepper Germplasm Resources [J]. Acta Horticulturae Sinica, 2025, 52(4): 857-871.
[8]	SHAO Yifan, ZHU Baoqing, WANG Tongxin, LIAO Jianhe, WU Fanhua, YANG Siyi, FENG Jianhang, YU Xudong. Research of Genetic Regulation of Bombax ceiba Prickle Based on Hormone，Transcriptome and Metabolome [J]. Acta Horticulturae Sinica, 2025, 52(4): 933-946.
[9]	TIAN Yufeng, MA Songya, YANG An, HAN Xiaolei, ZHANG Caixia. Establishment and Optimization of Regeneration System for Apple Rootstock B9 [J]. Acta Horticulturae Sinica, 2025, 52(4): 947-958.
[10]	DUAN Qingqing, HAN Meimei, WANG Youping, CHANG Peipei, TAN Yueqiang, ZHANG Zikun. Effects of Exogenous Melatonin on Seed Germination of Watermelon Under Saline-Alkali Stress and Comprehensive Evaluation [J]. Acta Horticulturae Sinica, 2025, 52(4): 1020-1036.
[11]	LIN Ruxue, WANG Fei, ZHANG Yanjie, MA Li, LIU Xiaofeng, LI Shuran, LIU Yalong, WANG Chen, JIANG Shuling, OU Chunqing. Transcriptome Screening and Functional Analysis of Dwarf Related Genes in Pear [J]. Acta Horticulturae Sinica, 2025, 52(3): 545-560.
[12]	WANG Rui, WU Hongfei, ZHANG Changyuan, CAO Haishun, TAN Delong, GUO Jinju, WANG Yunlong, WANG Rufang, YUAN Yu, WU Tingquan. CsPUB54 Negatively Regulates Cucumber Resistance to Phytophthora Melonis by Interacting with PmRXLR1 Effector [J]. Acta Horticulturae Sinica, 2025, 52(3): 591-602.
[13]	SUN Tingzhen, SHU Qin, MA Wei, SHI Yuzi, ZHANG Meng, XIANG Chenggang, BO Kailiang, DUAN Ying, WANG Changlin. Identification and Phylogenetic Analysis of Gibberellin Oxidase Gene Family and Its Expression Pattern in Main Vine Growth in Cucurbita maxima [J]. Acta Horticulturae Sinica, 2025, 52(3): 603-622.
[14]	HE Dandan, HE Hongtai, WANG Wenting, ZHOU Wenmei, LIU Yanmin, LIU Sushuang. Identification of Melon GolS Genes Family and the Expression Analysis in Response to Low Temperature Stress [J]. Acta Horticulturae Sinica, 2025, 52(1): 136-148.
[15]	SHI Xingxiu, FENG Beibei, YAN Peng, GENG Wenjuan, Jumazira Sharshanmukhan. Physiological Changes Associated with Early Watercore in‘Orin’Apples [J]. Acta Horticulturae Sinica, 2025, 52(1): 171-184.