中草药

灰毡毛忍冬花冠不展开表型与花冠细胞形态及XTH/EXP-PME表达特征的关联研究

PDF下载

敖志杰 ¹, , 付学森 ¹, , 周思敏 ¹, , 瞿美玲 ¹, , 朱佳源 ¹, , 袁林祥 ¹,,², , 周日宝 ³,⁴,⁵,¹,,², , 刘湘丹 ³,⁴,⁵,¹,,²,

中草药 | 药材与资源 2026,57(16): 6498-6509

收起

中草药 |药材与资源 2026 , 57 (16) : 6498 -6509

灰毡毛忍冬花冠不展开表型与花冠细胞形态及XTH/EXP-PME表达特征的关联研究

全屏

敖志杰¹,, 付学森¹,, 周思敏¹,, 瞿美玲¹,, 朱佳源¹,, 袁林祥¹,,²,, 周日宝³,⁴,⁵,¹,,²,, 刘湘丹³,⁴,⁵,¹,,²,

作者信息

^1.湖南中医药大学药学院, 湖南长沙 410208

^2.道地药材种质创制湖南省工程研究中心, 湖南长沙 410208

^3.湘产大宗道地药材种质资源及规范化种植重点研究室, 湖南长沙 410208

^4.湖南省普通高等学校中药现代化研究重点实验室, 湖南长沙 410208

^5.湖南省中药饮片标准化及功能工程技术研究中心, 湖南长沙 410208

^6.湖南中医药大学药学院, 湖南长沙 410208

^7.道地药材种质创制湖南省工程研究中心, 湖南长沙 410208

通讯作者:

周日宝

作者简介:

敖志杰: 敖志杰(2002—),男,硕士研究生,从事中药资源与质量研究。Tel:13873572934 E-mail:PolygalaeAozj@163.com

Correlation research of corolla non-opening phenotype with cell morphology and XTH/EXP-PME expression characteristics in Lonicera macranthoides

Affiliations

出版时间: 2026-08-28 doi: 10.7501/j.issn.0253-2670.2026.16.022

文章导航

摘要

收起

目的灰毡毛忍冬Lonicera macranthoides干燥的花蕾或带初开的花作为山银花入药，由其自然突变株培育的湘蕾品种具有花冠不展、花蕾期长的特殊表型，拟从细胞和分子水平探究湘蕾品种花冠不展开特殊表型的可能原因。方法以常规和湘蕾型灰毡毛忍冬花为研究对象，采用光学显微镜观察花冠近轴面表皮细胞以及花冠纵切面组织形态；qRT-PCR检测木葡聚糖内转糖苷酶/水解酶LmXTH6、LmXTH7、LmXTH33，扩展蛋白LmEXP7以及果胶甲酯酶LmPME28、LmPME45的表达模式；STRING数据库预测蛋白互作网络；钌红染色结合Image J定量评估花冠纵切面细胞壁果胶去甲酯化程度。结果常规型灰毡毛忍冬花冠近轴面表皮细胞呈现顺序扩张趋势，表现为基部优先扩张，随后中部扩张的趋势；且近轴面细胞的扩张速率大于远轴面；相比之下，湘蕾近轴面细胞扩张速率显著降低，细胞扩张程度不明显。分子层面，LmXTH6、LmXTH7、LmXTH33在常规品种开花前期及开花期高表达，在湘蕾品种中表达受到显著抑制；LmEXP7在常规花冠展开前表达量达到峰值，同一时期在湘蕾中表达较低。钌红染色结果表明湘蕾的整体去甲酯化程度高于常规，与LmPME28、LmPME45表达趋势一致。结论灰毡毛忍冬湘蕾品种花冠不展开与花冠细胞扩张受到抑制密切相关，XTHs及EXP的低表达共同导致了湘蕾花冠细胞的扩张能力下降，PMEs的上调增加了细胞壁果胶去甲酯化，可能增强了细胞壁刚性。XTH/EXP-PME协同可能是导致湘蕾花冠不展开的重要原因，其为湘蕾灰毡毛忍冬优良表型调控机制研究提供了实验依据，也为理解植物花器官形态建成的细胞壁调控机制提供了新见解。

关键词

灰毡毛忍冬 / 花冠运动 / 细胞壁 / 木葡聚糖内转糖苷酶/水解酶 / 扩展蛋白 / 果胶甲酯酶

Abstract

收起

Objective Dried flower buds or newly opened flowers of Lonicera macranthoides are used as Shanyinhua (Lonicerae Flos) for medicinal purposes. The Xianglei cultivar, developed from a natural mutant, exhibits a distinctive phenotype characterized by non-opening corolla and prolonged bud stage. Aiming to elucidate the cellular and molecular mechanisms underlying this unique corolla phenotype. Methods The flower of conventional cultivar (WT) and the Xianglei cultivar (XL) of L. macranthoides were used as materials. Corolla adaxial epidermal cells and longitudinal sections were observed using light microscopy. The expression patterns of xyloglucan endotransglucosylase/hydrolase genes (LmXTH6, LmXTH7, LmXTH33), expansin gene (LmEXP7), and pectin methylesterase genes (LmPME28, LmPME45) were analyzed by qRT-PCR. Protein-protein interaction networks were predicted using the STRING database. Ruthenium red staining coupled with Image J quantitative analysis was performed to assess the degree of pectin de-methylesterification in corolla cell walls. Results In the WT, the adaxial cells of corolla displayed a spatially sequential expansion, with the basal cells expanding first, followed by the middle cells. Moreover, the expansion rate of the adaxial cells is higher than that of the abaxial cells. In contrast, the expansion rate of the adaxial cells is significantly decreased in Xianglei, and the extend of cell expansion is not evident. Molecular analyses revealed that LmXTH6, LmXTH7, and LmXTH33 were highly expressed during the pre-anthesis and anthesis stages in WT, but were significantly suppressed in Xianglei. Expression of LmEXP7 peaked just before corolla opening in WT but remained low in Xianglei at the same stage. Furthermore, ruthenium staining showed a higher overall degree of pectin de-methylesterification in Xianglei compared with WT, which is consistent with the expressin patterns of LmPME28 and LmPME45. Conclusion The non-opening of the corolla in the L. macranthoides cultivar Xianglei is closely associated with inhibited expansion of corolla cells. The low expression of XTHs and EXP collectively contributes to the reduced expansion capacity of corolla cells in Xianglei, while the up-regulation of PMEs increases pectin de-methylesterification in the cell wall, potentially enhancing cell wall stiffness. The synergistic effect of XTH/EXP and PME may be an important factor leading to corolla non-opening in Xianglei. These findings provide experimental evidence for understanding the regulatory mechanisms underlying the superior phenotype of the Xianglei cultivar and offer new insights into the cell wall regulatory mechanisms governing floral organ morphogenesis in plants.

Key words

Lonicera macranthoides Hand.-Mazz. / corolla movement / cell wall / xyloglucan endotransglucosylase/hydrolase / expansin / pectin methylesterase

引用本文

敖志杰, 付学森, 周思敏, 瞿美玲, 朱佳源, 袁林祥, 周日宝, 刘湘丹. 灰毡毛忍冬花冠不展开表型与花冠细胞形态及XTH/EXP-PME表达特征的关联研究. 中草药, 2026 , 57 (16) : 6498 -6509 . DOI: 10.7501/j.issn.0253-2670.2026.16.022

AO Zhijie, FU Xuesen, ZHOU Simin, QU Meiling, ZHU Jiayuan, YUAN Linxiang, ZHOU Ribao, LIU Xiangdan. Correlation research of corolla non-opening phenotype with cell morphology and XTH/EXP-PME expression characteristics in Lonicera macranthoides[J]. Chinese Traditional and Herbal Drugs, 2026 , 57 (16) : 6498 -6509 . DOI: 10.7501/j.issn.0253-2670.2026.16.022

基金

收起

国家自然科学基金资助项目
国家现代农业产业技术体系
湖南省中药材产业技术体系专项
2020年湖南省一流专业建设点：中药资源与开发
湖南中医药大学重点学科中药学科
2025湖南中医药大学研究生科研创新项目

参考文献

收起

文献

收起

参考文献引证文献

排序方式：

中国药典[S]. 一部. 2025: 56.

刘思思, 乔中全, 刘新民, 等. 花蕾型和普通型灰毡毛忍冬花冠开裂过程的转录组比较分析[J]. 分子植物育种, 2021, 19(6): 1822-1829.

王晓明. 灰毡毛忍冬新品种ISSR分子标记及组织培养的研究[D]. 长沙: 中南林业科技大学, 2012.

彭美晨, 刘湘丹, 徐玉琴, 等. 灰毡毛忍冬Lm-XL-AP1基因克隆、生物信息学和时空表达分析[J]. 中草药, 2018(7): 1652-1660.

俞亚欣, 龙丽君, 李昌珠, 等. 灰毡毛忍冬AP1同源基因的克隆及互作蛋白鉴定[J]. 药学学报, 2024, 59(10): 2880-2888.

王珊. 灰毡毛忍冬转录组测序及AP1和AGL19基因的克隆与表达分析[D]. 长沙: 湖南中医药大学, 2019.

蔡嘉洛, 朱贻霖, 谢舒平, 等. 灰毡毛忍冬内参基因筛选和Mads-box家族基因AGL15的时空表达分析[J]. 中草药, 2016, 47(15): 2727-2733.

易刚强, 蔡嘉洛, 朱贻霖, 等. 灰毡毛忍冬MADS-box基因家族AGL15基因的克隆、生物信息学和表达分析[J]. 中草药, 2016, 47(4): 640-647.

付学森, 刘紫璇, 王玲, 等. 灰毡毛忍冬MADS-box基因家族鉴定与CMB1基因克隆[J]. 湖南中医药大学学报, 2024, 44(3): 383-394.

龙丽君. 灰毡毛忍冬花器官发育相关MADS: box基因的克隆及功能分析[D]. 长沙: 中南林业科技大学, 2024.

孙梦姗, 周日宝, 彭晓丹, 等. 灰毡毛忍冬不同品种开花时期内源乙烯动态变化的研究[J]. 中成药, 2013, 35(9): 1969-1972.

刘畅宇. 灰毡毛忍冬乙烯生物合成途径关键酶ACS、ACO基因的全长克隆及表达分析[D]. 长沙: 湖南中医药大学, 2019.

Long Y Q, Zeng J, Yang M, et al. Comparative transcriptome analysis to reveal key ethylene genes involved in a Lonicera macranthoides mutant [J]. Genes Genom, 2023, 45(4): 437-450.

张曼, 张叶卓, 何其邹洪, 等. 植物细胞壁结构及成像技术研究进展[J]. 生物技术通报, 2023, 39(7): 113-122.

Zhang B C, Gao Y H, Zhang L J, et al. The plant cell wall: Biosynthesis, construction, and functions [J]. J Integr Plant Biol, 2021, 63(1): 251-272.

Samalova M, Melnikava A, Elsayad K, et al. Hormone-regulated expansins: Expression, localization, and cell wall biomechanics in Arabidopsis root growth [J]. Plant Physiol, 2023, 194(1): 209-228.

Miedes E, Zarra I, Hoson T, et al. Xyloglucan endotransglucosylase and cell wall extensibility [J]. J Plant Physiol, 2011, 168(3): 196-203.

Watanabe Y, Niki T, Norikoshi R, et al. Soluble carbohydrate concentration and expression of expansin and xyloglucan endotransglucosylase/hydrolase genes in epidermal and parenchyma cells during lily flower opening [J]. J Plant Physiol, 2022, 270: 153615.

Singh A P, Dubey S, Lakhwani D, et al. Differential expression of several xyloglucan endotransglucosylase/ hydrolase genes regulates flower opening and petal abscission in roses [J]. Aob PLANTS, 2013, 5: plt030.

Harada T, Torii Y, Morita S, et al. Cloning, characterization, and expression of xyloglucan endotransglucosylase/hydrolase and expansin genes associated with petal growth and development during carnation flower opening [J]. J Exp Bot, 2011, 62(2): 815-823.

Yang Y, Miao Y F, Zhong S W, et al. Genome-wide identification and expression analysis of XTH gene family during flower-opening stages in Osmanthus fragrans [J]. Plants, 2022, 11(8): 1015.

Del Corpo D, Fullone M R, Miele R, et al. AtPME17 is a functional Arabidopsis thaliana pectin methylesterase regulated by its PRO region that triggers PME activity in the resistance to Botrytis cinerea [J]. Mol Plant Pathol, 2020, 21(12): 1620-1633.

Wang S J, Li R M, Zhou Y J, et al. Integrated characterization of cassava (Manihot esculenta) pectin methylesterase (MePME) genes to filter candidate gene responses to multiple abiotic stresses [J]. Plants, 2023, 12(13): 2529.

Sun J H, Tian Z, Li X X, et al. Systematic analysis of the pectin methylesterase gene family in Nicotiana tabacum and reveal their multiple roles in plant development and abiotic stresses [J]. Front Plant Sci, 2022, 13: 998841.

Wang Y, Peng Y Z, Shangguan X C, et al. The pectin methylesterase OsPME14 modifies the cell wall to confer copper tolerance in Oryza sativa L. [J]. Plant J, 2025, 122(4): e70173.

Shi Y N, Li B J, Grierson D, et al. Insights into cell wall changes during fruit softening from transgenic and naturally occurring mutants [J]. Plant Physiol, 2023, 192(3): 1671-1683.

Liao J Q, Shang Y K, Han Z G, et al. Dynamic remodeling of pectin and hemicellulose for contrastive cadmium accumulations with root cell walls in distinct Salvia miltiorrhiza ecotypes [J]. Int J Biol Macromol, 2025, 329: 147782.

周思敏, 瞿美玲, 曾娟, 等. 灰毡毛忍冬AP2/ERF家族成员鉴定及表达分析[J]. 中国中药杂志, 2025, 50(15): 4248-4262.

刘湘丹, 徐玉琴, 王珊, 等. 适用于基因全长克隆的灰毡毛忍冬不同器官总RNA提取方法筛选[J]. 湖南中医药大学学报, 2015, 35(12): 60-63.

敖志杰, 张惊宇, 曾娟, 等. 灰毡毛忍冬CHS1基因克隆及功能分析[J]. 中草药, 2025, 56(17): 6334-6345.

瞿美玲, 周思敏, 张惊宇, 等. 灰毡毛忍冬bHLH转录基因家族的鉴定与表达分析[J]. 生物技术通报, 2025, 41(6): 256-268.

Chen C X, Hussain N, Ma Y X, et al. The ARF2–MYB6 module mediates auxin-regulated petal expansion in rose [J]. J Exp Bot, 2023, 74(15): 4489-4502.

Cheng C X, Yu Q, Wang Y R, et al. Ethylene-regulated asymmetric growth of the petal base promotes flower opening in rose (Rosa hybrida) [J]. Plant Cell, 2021, 33(4): 1229-1251.

Miao Y F, Yang Y, Zhu H J, et al. The basic/helix-loop-helix 79(bHLH79)-xyloglucan endotransglucosylase/ hydrolase 28(XTH28) regulatory module mediates flower opening in Osmanthus fragrans [J]. Int J Biol Macromol, 2025, 322: 146633.

Tsuchiya M, Satoh S, Iwai H. Distribution of XTH, expansin, and secondary-wall-related CesA in floral and fruit abscission zones during fruit development in tomato (Solanum lycopersicum) [J]. Front Plant Sci, 2015, 6: 323.

张瑞, 赵忠. 细胞壁动态调控及其对植物发育的影响[J]. 植物学报, 2026, 61(1): 15-25.

Pitaksaringkarn W, Matsuoka K, Asahina M, et al. XTH20 and XTH19 regulated by ANAC071 under auxin flow are involved in cell proliferation in incised Arabidopsis inflorescence stems [J]. Plant J, 2014, 80(4): 604-614.

Gallemí M, Montesinos J C, Zarevski N, et al. Dual role of pectin methyl esterase activity in the regulation of plant cell wall biophysical properties [J]. Front Plant Sci, 2025, 16: 1612366.

Chen YX. Research progress on plant cell wall structure and regulatory mechanism of pectin metabolism [J]. Int J Ecol, 2025, 14(3): 285-297.

Sénéchal F, Wattier C, Rustérucci C, et al. Homogalacturonan-modifying enzymes: Structure, expression, and roles in plants [J]. J Exp Bot, 2014, 65(18): 5125-5160.

Yao Y D, Yang Y, Ding Z Q, et al. Nitric oxide delays tomato fruit softening by inhibiting SlNAP2(NAC-like, activated by apetala3/pistillata2) transcription factor-activated transcription of soften-related genes [J]. Int J Biol Macromol, 2025, 309: 143148.

Wen B, Zhang F, Wu X Z, et al. Characterization of the tomato (Solanum lycopersicum) pectin methylesterases: Evolution, activity of isoforms and expression during fruit ripening [J]. Front Plant Sci, 2020, 11: 238.

Cai J F, Mo X L, Wen C J, et al. FvMYB79 positively regulates strawberry fruit softening via transcriptional activation of FvPME38[J]. Int J Mol Sci, 2022, 23(1): 101.

Xue C, Guan S C, Chen J Q, et al. Genome wide identification and functional characterization of strawberry pectin methylesterases related to fruit softening [J]. BMC Plant Biol, 2020, 20(1): 13.

Wang S Q, Liu C Y, Su X R, et al. Transcriptome analysis reveals key metabolic pathways and gene expression involving in cell wall polysaccharides-disassembling and postharvest fruit softening in custard apple (Annona squamosa L.) [J]. Int J Biol Macromol, 2023, 240: 124356.

Huang W N, Shi Y N, Yan H, et al. The calcium-mediated homogalacturonan pectin complexation in cell walls contributes the firmness increase in loquat fruit during postharvest storage [J]. J Adv Res, 2023, 49: 47-62.

Ochei C, Basiouny F M, Woods F M. Calcium-mediated postharvest changes in storageability and fruit quality of peaches [A] // Proceedings of the Florida State Horticultural Society [C]. New York: Florida State Horticultural Society, 1994,106:266-269.

Yang L, He J L, Qin S J, et al. MYB transcription factor MdMYB44 positively regulates fruit crispness by directly activating the expression of pectin methylesterase MdMPE3 in apple [J]. Plant Physiol Biochem, 2025, 224: 109936.

Ding W Y, Gou Y J, Li Y J, et al. A jasmonate-mediated regulatory network modulates diurnal floret opening time in rice [J]. New Phytol, 2024, 244(1): 176-191.

Wang M M, Zhu X P, Peng G Q, et al. Methylesterification of cell-wall pectin controls the diurnal flower-opening times in rice [J]. Mol Plant, 2022, 15(6): 956-972.

Pei Y Y, Wang Y K, Wei Z Z, et al. Pectin methylesterase inhibitors GhPMEI53 and AtPMEI19 improve seed germination by modulating cell wall plasticity in cotton and Arabidopsis [J]. J Integr Agric, 2024, 23(10): 3487-3505.

Wang X, Wilson L, Cosgrove D J. Pectin methylesterase selectively softens the onion epidermal wall yet reduces acid-induced creep [J]. J Exp Bot, 2020, 71(9): 2629-2640.

Zhang Q, Du J, Hao S, et al. Pectin metabolism influences phloem architecture and flowering time in Arabidopsis Thaliana [J]. Adv Sci, 2025, 12(39): e02980.

Thanomchit K, Imsabai W, Burns P, et al. Differential expression of ethylene biosynthetic and receptor genes in pollination-induced senescence of Dendrobium flowers [J]. Plant Physiol Biochem, 2022, 188: 38-46.

Beauzamy L, Nakayama N, Boudaoud A. Flowers under pressure: Ins and outs of turgor regulation in development [J]. Ann Bot, 2014, 114(7): 1517-1533.

Iakimova E T, Woltering E J. Revealing programmed cell death events during flower (petal) senescence [J]. Ann Bot, 2025: mcaf105.

Tieman D M, Handa A K. Reduction in pectin methylesterase activity modifies tissue integrity and cation levels in ripening tomato (Lycopersicon esculentum Mill.) fruits [J]. Plant Physiol, 1994, 106(2): 429-436.

Gong F F, Jing W K, Jin W C, et al. RhMYC₂ controls petal size through synergistic regulation of jasmonic acid and cytokinin signaling in rose [J]. Plant J, 2024, 120(2): 459-472.

Gong F F, Wang X Y, Zhao Q C, et al. The GAs–RhMYB70 feedback loop fine-tunes cell expansion and petal size by modulating cellulose content in rose [J]. Hortic Res, 2025, 12(8):uhaf134.

Jing W K, Gong F F, Liu G Q, et al. Petal size is controlled by the MYB73/TPL/HDA19-miR159-CKX6 module regulating cytokinin catabolism in Rosa hybrida [J]. Nat Commun, 2023, 14: 7106.

Guan Y X, Ding L, Jiang J F, et al. The TIFY family protein CmJAZ1-like negatively regulates petal size via interaction with the bHLH transcription factor CmBPE2 in Chrysanthemum morifolium [J]. Plant J, 2022, 112(6): 1489-1506.

Varaud E, Brioudes F, Szécsi J, et al. AUXIN RESPONSE FACTOR8 regulates Arabidopsis petal growth by interacting with the bHLH transcription factor BIGPETALp [J]. Plant Cell, 2011, 23(3): 973-983.

Pei H X, Ma N, Tian J, et al. An NAC transcription factor controls ethylene-regulated cell expansion in flower petals [J]. Plant Physiol, 2013, 163(2): 775-791.

Dai F W, Zhang C Q, Jiang X Q, et al. RhNAC2 and RhEXPA4 are involved in the regulation of dehydration tolerance during the expansion of rose petals [J]. Plant Physiol, 2012, 160(4):2064-2082.

中国药典[S]. 一部. 2025: 56.
刘思思, 乔中全, 刘新民, 等. 花蕾型和普通型灰毡毛忍冬花冠开裂过程的转录组比较分析[J]. 分子植物育种, 2021, 19(6): 1822-1829.
王晓明. 灰毡毛忍冬新品种ISSR分子标记及组织培养的研究[D]. 长沙: 中南林业科技大学, 2012.
彭美晨, 刘湘丹, 徐玉琴, 等. 灰毡毛忍冬Lm-XL-AP1基因克隆、生物信息学和时空表达分析[J]. 中草药, 2018(7): 1652-1660.
俞亚欣, 龙丽君, 李昌珠, 等. 灰毡毛忍冬AP1同源基因的克隆及互作蛋白鉴定[J]. 药学学报, 2024, 59(10): 2880-2888.
王珊. 灰毡毛忍冬转录组测序及AP1和AGL19基因的克隆与表达分析[D]. 长沙: 湖南中医药大学, 2019.
蔡嘉洛, 朱贻霖, 谢舒平, 等. 灰毡毛忍冬内参基因筛选和Mads-box家族基因AGL15的时空表达分析[J]. 中草药, 2016, 47(15): 2727-2733.
易刚强, 蔡嘉洛, 朱贻霖, 等. 灰毡毛忍冬MADS-box基因家族AGL15基因的克隆、生物信息学和表达分析[J]. 中草药, 2016, 47(4): 640-647.
付学森, 刘紫璇, 王玲, 等. 灰毡毛忍冬MADS-box基因家族鉴定与CMB1基因克隆[J]. 湖南中医药大学学报, 2024, 44(3): 383-394.
龙丽君. 灰毡毛忍冬花器官发育相关MADS: box基因的克隆及功能分析[D]. 长沙: 中南林业科技大学, 2024.
孙梦姗, 周日宝, 彭晓丹, 等. 灰毡毛忍冬不同品种开花时期内源乙烯动态变化的研究[J]. 中成药, 2013, 35(9): 1969-1972.
刘畅宇. 灰毡毛忍冬乙烯生物合成途径关键酶ACS、ACO基因的全长克隆及表达分析[D]. 长沙: 湖南中医药大学, 2019.
Long Y Q, Zeng J, Yang M, et al. Comparative transcriptome analysis to reveal key ethylene genes involved in a Lonicera macranthoides mutant [J]. Genes Genom, 2023, 45(4): 437-450.
张曼, 张叶卓, 何其邹洪, 等. 植物细胞壁结构及成像技术研究进展[J]. 生物技术通报, 2023, 39(7): 113-122.
Zhang B C, Gao Y H, Zhang L J, et al. The plant cell wall: Biosynthesis, construction, and functions [J]. J Integr Plant Biol, 2021, 63(1): 251-272.
Samalova M, Melnikava A, Elsayad K, et al. Hormone-regulated expansins: Expression, localization, and cell wall biomechanics in Arabidopsis root growth [J]. Plant Physiol, 2023, 194(1): 209-228.
Miedes E, Zarra I, Hoson T, et al. Xyloglucan endotransglucosylase and cell wall extensibility [J]. J Plant Physiol, 2011, 168(3): 196-203.
Watanabe Y, Niki T, Norikoshi R, et al. Soluble carbohydrate concentration and expression of expansin and xyloglucan endotransglucosylase/hydrolase genes in epidermal and parenchyma cells during lily flower opening [J]. J Plant Physiol, 2022, 270: 153615.
Singh A P, Dubey S, Lakhwani D, et al. Differential expression of several xyloglucan endotransglucosylase/ hydrolase genes regulates flower opening and petal abscission in roses [J]. Aob PLANTS, 2013, 5: plt030.
Harada T, Torii Y, Morita S, et al. Cloning, characterization, and expression of xyloglucan endotransglucosylase/hydrolase and expansin genes associated with petal growth and development during carnation flower opening [J]. J Exp Bot, 2011, 62(2): 815-823.
Yang Y, Miao Y F, Zhong S W, et al. Genome-wide identification and expression analysis of XTH gene family during flower-opening stages in Osmanthus fragrans [J]. Plants, 2022, 11(8): 1015.
Del Corpo D, Fullone M R, Miele R, et al. AtPME17 is a functional Arabidopsis thaliana pectin methylesterase regulated by its PRO region that triggers PME activity in the resistance to Botrytis cinerea [J]. Mol Plant Pathol, 2020, 21(12): 1620-1633.
Wang S J, Li R M, Zhou Y J, et al. Integrated characterization of cassava (Manihot esculenta) pectin methylesterase (MePME) genes to filter candidate gene responses to multiple abiotic stresses [J]. Plants, 2023, 12(13): 2529.
Sun J H, Tian Z, Li X X, et al. Systematic analysis of the pectin methylesterase gene family in Nicotiana tabacum and reveal their multiple roles in plant development and abiotic stresses [J]. Front Plant Sci, 2022, 13: 998841.
Wang Y, Peng Y Z, Shangguan X C, et al. The pectin methylesterase OsPME14 modifies the cell wall to confer copper tolerance in Oryza sativa L. [J]. Plant J, 2025, 122(4): e70173.
Shi Y N, Li B J, Grierson D, et al. Insights into cell wall changes during fruit softening from transgenic and naturally occurring mutants [J]. Plant Physiol, 2023, 192(3): 1671-1683.
Liao J Q, Shang Y K, Han Z G, et al. Dynamic remodeling of pectin and hemicellulose for contrastive cadmium accumulations with root cell walls in distinct Salvia miltiorrhiza ecotypes [J]. Int J Biol Macromol, 2025, 329: 147782.
周思敏, 瞿美玲, 曾娟, 等. 灰毡毛忍冬AP2/ERF家族成员鉴定及表达分析[J]. 中国中药杂志, 2025, 50(15): 4248-4262.
刘湘丹, 徐玉琴, 王珊, 等. 适用于基因全长克隆的灰毡毛忍冬不同器官总RNA提取方法筛选[J]. 湖南中医药大学学报, 2015, 35(12): 60-63.
敖志杰, 张惊宇, 曾娟, 等. 灰毡毛忍冬CHS1基因克隆及功能分析[J]. 中草药, 2025, 56(17): 6334-6345.
瞿美玲, 周思敏, 张惊宇, 等. 灰毡毛忍冬bHLH转录基因家族的鉴定与表达分析[J]. 生物技术通报, 2025, 41(6): 256-268.
Chen C X, Hussain N, Ma Y X, et al. The ARF2–MYB6 module mediates auxin-regulated petal expansion in rose [J]. J Exp Bot, 2023, 74(15): 4489-4502.
Cheng C X, Yu Q, Wang Y R, et al. Ethylene-regulated asymmetric growth of the petal base promotes flower opening in rose (Rosa hybrida) [J]. Plant Cell, 2021, 33(4): 1229-1251.
Miao Y F, Yang Y, Zhu H J, et al. The basic/helix-loop-helix 79(bHLH79)-xyloglucan endotransglucosylase/ hydrolase 28(XTH28) regulatory module mediates flower opening in Osmanthus fragrans [J]. Int J Biol Macromol, 2025, 322: 146633.
Tsuchiya M, Satoh S, Iwai H. Distribution of XTH, expansin, and secondary-wall-related CesA in floral and fruit abscission zones during fruit development in tomato (Solanum lycopersicum) [J]. Front Plant Sci, 2015, 6: 323.
张瑞, 赵忠. 细胞壁动态调控及其对植物发育的影响[J]. 植物学报, 2026, 61(1): 15-25.
Pitaksaringkarn W, Matsuoka K, Asahina M, et al. XTH20 and XTH19 regulated by ANAC071 under auxin flow are involved in cell proliferation in incised Arabidopsis inflorescence stems [J]. Plant J, 2014, 80(4): 604-614.
Gallemí M, Montesinos J C, Zarevski N, et al. Dual role of pectin methyl esterase activity in the regulation of plant cell wall biophysical properties [J]. Front Plant Sci, 2025, 16: 1612366.
Chen YX. Research progress on plant cell wall structure and regulatory mechanism of pectin metabolism [J]. Int J Ecol, 2025, 14(3): 285-297.
Sénéchal F, Wattier C, Rustérucci C, et al. Homogalacturonan-modifying enzymes: Structure, expression, and roles in plants [J]. J Exp Bot, 2014, 65(18): 5125-5160.
Yao Y D, Yang Y, Ding Z Q, et al. Nitric oxide delays tomato fruit softening by inhibiting SlNAP2(NAC-like, activated by apetala3/pistillata2) transcription factor-activated transcription of soften-related genes [J]. Int J Biol Macromol, 2025, 309: 143148.
Wen B, Zhang F, Wu X Z, et al. Characterization of the tomato (Solanum lycopersicum) pectin methylesterases: Evolution, activity of isoforms and expression during fruit ripening [J]. Front Plant Sci, 2020, 11: 238.
Cai J F, Mo X L, Wen C J, et al. FvMYB79 positively regulates strawberry fruit softening via transcriptional activation of FvPME38[J]. Int J Mol Sci, 2022, 23(1): 101.
Xue C, Guan S C, Chen J Q, et al. Genome wide identification and functional characterization of strawberry pectin methylesterases related to fruit softening [J]. BMC Plant Biol, 2020, 20(1): 13.
Wang S Q, Liu C Y, Su X R, et al. Transcriptome analysis reveals key metabolic pathways and gene expression involving in cell wall polysaccharides-disassembling and postharvest fruit softening in custard apple (Annona squamosa L.) [J]. Int J Biol Macromol, 2023, 240: 124356.
Huang W N, Shi Y N, Yan H, et al. The calcium-mediated homogalacturonan pectin complexation in cell walls contributes the firmness increase in loquat fruit during postharvest storage [J]. J Adv Res, 2023, 49: 47-62.
Ochei C, Basiouny F M, Woods F M. Calcium-mediated postharvest changes in storageability and fruit quality of peaches [A] // Proceedings of the Florida State Horticultural Society [C]. New York: Florida State Horticultural Society, 1994,106:266-269.
Yang L, He J L, Qin S J, et al. MYB transcription factor MdMYB44 positively regulates fruit crispness by directly activating the expression of pectin methylesterase MdMPE3 in apple [J]. Plant Physiol Biochem, 2025, 224: 109936.
Ding W Y, Gou Y J, Li Y J, et al. A jasmonate-mediated regulatory network modulates diurnal floret opening time in rice [J]. New Phytol, 2024, 244(1): 176-191.
Wang M M, Zhu X P, Peng G Q, et al. Methylesterification of cell-wall pectin controls the diurnal flower-opening times in rice [J]. Mol Plant, 2022, 15(6): 956-972.
Pei Y Y, Wang Y K, Wei Z Z, et al. Pectin methylesterase inhibitors GhPMEI53 and AtPMEI19 improve seed germination by modulating cell wall plasticity in cotton and Arabidopsis [J]. J Integr Agric, 2024, 23(10): 3487-3505.
Wang X, Wilson L, Cosgrove D J. Pectin methylesterase selectively softens the onion epidermal wall yet reduces acid-induced creep [J]. J Exp Bot, 2020, 71(9): 2629-2640.
Zhang Q, Du J, Hao S, et al. Pectin metabolism influences phloem architecture and flowering time in Arabidopsis Thaliana [J]. Adv Sci, 2025, 12(39): e02980.
Thanomchit K, Imsabai W, Burns P, et al. Differential expression of ethylene biosynthetic and receptor genes in pollination-induced senescence of Dendrobium flowers [J]. Plant Physiol Biochem, 2022, 188: 38-46.
Beauzamy L, Nakayama N, Boudaoud A. Flowers under pressure: Ins and outs of turgor regulation in development [J]. Ann Bot, 2014, 114(7): 1517-1533.
Iakimova E T, Woltering E J. Revealing programmed cell death events during flower (petal) senescence [J]. Ann Bot, 2025: mcaf105.
Tieman D M, Handa A K. Reduction in pectin methylesterase activity modifies tissue integrity and cation levels in ripening tomato (Lycopersicon esculentum Mill.) fruits [J]. Plant Physiol, 1994, 106(2): 429-436.
Gong F F, Jing W K, Jin W C, et al. RhMYC₂ controls petal size through synergistic regulation of jasmonic acid and cytokinin signaling in rose [J]. Plant J, 2024, 120(2): 459-472.
Gong F F, Wang X Y, Zhao Q C, et al. The GAs–RhMYB70 feedback loop fine-tunes cell expansion and petal size by modulating cellulose content in rose [J]. Hortic Res, 2025, 12(8):uhaf134.
Jing W K, Gong F F, Liu G Q, et al. Petal size is controlled by the MYB73/TPL/HDA19-miR159-CKX6 module regulating cytokinin catabolism in Rosa hybrida [J]. Nat Commun, 2023, 14: 7106.
Guan Y X, Ding L, Jiang J F, et al. The TIFY family protein CmJAZ1-like negatively regulates petal size via interaction with the bHLH transcription factor CmBPE2 in Chrysanthemum morifolium [J]. Plant J, 2022, 112(6): 1489-1506.
Varaud E, Brioudes F, Szécsi J, et al. AUXIN RESPONSE FACTOR8 regulates Arabidopsis petal growth by interacting with the bHLH transcription factor BIGPETALp [J]. Plant Cell, 2011, 23(3): 973-983.
Pei H X, Ma N, Tian J, et al. An NAC transcription factor controls ethylene-regulated cell expansion in flower petals [J]. Plant Physiol, 2013, 163(2): 775-791.
Dai F W, Zhang C Q, Jiang X Q, et al. RhNAC2 and RhEXPA4 are involved in the regulation of dehydration tolerance during the expansion of rose petals [J]. Plant Physiol, 2012, 160(4):2064-2082.

2026年第57卷第16期

PDF下载

115

引用本文

BibTeX

文章信息

doi: 10.7501/j.issn.0253-2670.2026.16.022

接收时间：2026-01-02
首发时间：2026-10-22
出版时间：2026-08-28

补充材料

相关文章

文章信息

作者

出版历史

收稿日期：2026-01-02

基金

国家自然科学基金资助项目

国家现代农业产业技术体系

湖南省中药材产业技术体系专项

2020年湖南省一流专业建设点：中药资源与开发

湖南中医药大学重点学科中药学科

2025湖南中医药大学研究生科研创新项目

作者信息

^1.湖南中医药大学药学院, 湖南长沙 410208

^2.道地药材种质创制湖南省工程研究中心, 湖南长沙 410208

^3.湘产大宗道地药材种质资源及规范化种植重点研究室, 湖南长沙 410208

^4.湖南省普通高等学校中药现代化研究重点实验室, 湖南长沙 410208

^5.湖南省中药饮片标准化及功能工程技术研究中心, 湖南长沙 410208

^6.湖南中医药大学药学院, 湖南长沙 410208

^7.道地药材种质创制湖南省工程研究中心, 湖南长沙 410208

通讯作者:

参考文献

分享链接

https://castjournals.cast.org.cn/joweb/zcy/CN/10.7501/j.issn.0253-2670.2026.16.022

分享至

全文二维码

扫描看全文

引用本文

BibTeX

本文的引用情况

2种不同金属材料的力学参数

科 Family	属数 Number of genus	种数 Number of species	占总种数比例 Percentage of total species (%)	属 Genus	种数 Number of species	占总种数比例 Percentage of total species (%)
鹅膏菌科Amanitaceae	2	11	5.26	鹅膏菌属 Amanita	10	4.78
小菇科 Mycenaceae	2	12	5.74	丝盖伞属 Inocybe	5	2.39
多孔菌科 Polyporaceae	8	14	6.70	蜡蘑属 Laccaria	5	2.39
红菇科 Russulaceae	3	23	11.00	小皮伞属 Marasmius	6	2.87
				小菇属 Mycena	11	5.26
				光柄菇属 Pluteus	5	2.39
				红菇属 Russula	17	8.13
				栓菌属 Trametes	5	2.39

关闭全屏

BibTeX
EndNote
RefWorks
TxT