乙烯代谢和能量状态对‘巴茨拉’牡丹切花瓶插品质的作用研究

doi:10.16420/j.issn.0513-353x.2020-0094

摘要/Abstract

摘要：

以发育至绽口期的‘巴茨拉’牡丹切花为试材,分别经蒸馏水和20 mg · L^-1 纳米银预处理2 h,再进行蒸馏水或保鲜液瓶插,测定瓶插过程中花瓣乙烯释放速率,1-氨基环丙烷-1-羧酸（ACC）、腺苷酸（ANP）和糖含量,乙烯和糖代谢相关酶活性。结果表明,纳米银预处理乙烯释放峰值降幅达40.9%,降低了ACC含量和ACC合成酶（ACS）活性,提高了琥珀酸脱氢酶（SDH）和H⁺-ATP酶活性,增加了糖和ATP含量,保持较高的能荷（EC）水平,切花瓶插寿命延长了40.7 h。保鲜液瓶插乙烯释放峰值降幅达35.1%,提高了SDH活性与糖及ATP含量,保持较高的EC值,最大花径增加了3.5 cm。纳米银预处理后保鲜液瓶插（NS + PS）乙烯释放峰值降幅达31.2%,降低了ACC含量和ACS活性,提高了SDH、CCO和H⁺-ATP酶活性,糖及ANP含量,维持了较高的EC水平,提高了可溶性蛋白质含量,瓶插寿命延长了46.4 h,最大花径增大3.6 cm。结果显示通过抑制乙烯代谢和增强能量状态可延缓‘巴茨拉’牡丹切花衰老提高瓶插品质。

关键词: 牡丹, 切花, 乙烯代谢, 能量状态, 瓶插品质

Abstract:

The regulatory effects of ethylene metabolism and energy status on the vase quality of cut Itoh peony‘Bartzella’flowers were studied by using nano silver and preservative solution treatment. Cut Itoh peony flowers with a developmental level of stageⅡwere employed as test materials and were pretreated with distilled water（the control）or 20 mg · L^-1 nano silver（NS）for 2 h. The samples were then vased in preservative solution（PS）directly after distilled water treatment or after nano silver treatment （NS + PS）. The release rate of ethylene,the contents of 1-aminocyclopropane-1-carboxylic acid（ACC）,adenine nucleotides（ATP + ADP + AMP）,sugar,and the activities of enzymes related to ethylene and sugar metabolism during the vase insertion process were measured. The results demonstrated that nano-silver pretreatment（NS）inhibited the peak of ethylene release by 40.9%,and reduced ACC content and ACC synthetase（ACS）activity,while increased the activities of succinic dehydrogenase（SDH）and H⁺-ATPase,and contents of sugar and ATP,which maintained a high energy charge（EC）level,therefore,prolonged vase life by 40.7 h. The PS also inhibited the peak of ethylene release by 35.1%,and increased SDH activity and sugar and ATP contents,which maintained a high EC value,and increased the maximum flower diameter by 3.5 cm. Moreover,NS + PS group showed inhibition of ethylene release peak by 31.2%,and reduction of ACC content and ACS activity,while SDH,CCO,and H⁺-ATPase activities,and the contents of sugar and ATP were all increased,which also maintained high EC levels and increased the soluble protein content,and therefore,increased flower vase life by 46.4 h,and the maximum flower diameter by 3.6 cm. The results showed that inhibition of ethylene metabolism and enhancement of energy status could delay the senescence and improve the vase quality of Itoh peony‘Bartzella’cut flowers.

Key words: tree peony, cut flower, ethylene metabolism, energy status, vase quality

中图分类号:

S685.11

王依, 王凯轩, 胡思源, 周爽, 史国安. 乙烯代谢和能量状态对‘巴茨拉’牡丹切花瓶插品质的作用研究[J]. 园艺学报, 2021, 48(6): 1135-1149.

WANG Yi, WANG Kaixuan, HU Siyuan, ZHOU Shuang, SHI Guoan. Effects of Ethylene Metabolism and Energy Status on Vase Quality of Cut Itoh Peony‘Bartzella’Flowers[J]. Acta Horticulturae Sinica, 2021, 48(6): 1135-1149.

图/表 8

图1 ‘巴茨拉’牡丹切花瓶插过程中的形态变化 H2O：蒸馏水对照;PS：保鲜液;NS：纳米银预处理;NS + PS：纳米银 + 保鲜液。下同。

Fig. 1 The morphological changes of cut Itoh peony‘Bartzella’flowers during vase periods H2O：Distilled water control;PS：Preservative solution;NS：Nano silver;NS + PS：Nano silver + Preservative solution. The same below.

表1 纳米银（NS）和保鲜液（PS）对‘巴茨拉’牡丹切花瓶插寿命、最佳观赏期和最大花径的影响

Table 1 Effects of nano silver and preservative solution on vase life,best viewing period and maximum flower diameter of cut Itoh peony ‘Bartzella’flowers in vase

处理 Treatment	瓶插寿命/h Vase life	最佳观赏期/h Best viewing period	最大花径/cm Max flower diameter
H₂O（对照Control） PS NS + H₂O NS + PS	93.3 ± 3.1 b 93.7 ± 3.1 b 134.0 ± 2.9 a 139.7 ± 4.3 a	41.5 ± 4.3 c 65.5 ± 5.3 b 62.7 ± 4.7 b 96.0 ± 6.1 a	18.1 ± 0.84 b 21.6 ± 1.52 a 18.5 ± 1.09 b 21.7 ± 1.60 a

图2 ‘巴茨拉’牡丹切花花枝鲜样质量和花径变化

Fig. 2 Changes of fresh mass and flower diameter of cut Itoh peony‘Bartzella’flowers during vase periods

图3 ‘巴茨拉’牡丹切花花枝水分平衡值和花瓣可溶性蛋白质含量的变化

Fig. 3 Changes of water balance value and soluble protein content of cut Itoh peony‘Bartzella’flowers during vase periods

图4 ‘巴茨拉’牡丹花瓣乙烯释放速率、ACC含量、ACS和ACO活性的变化

Fig. 4 Changes of ethylene production,ACC content,ACS and ACO activities of cut Itoh peony‘Bartzella’flowers during vase periods

图5 ‘巴茨拉’牡丹花瓣中ATP、ADP和AMP含量及能荷的变化

Fig. 5 Changes of ATP,ADP and AMP contents and energy charge of cut Itoh peony‘Bartzella’flowers during vase periods

图6 ‘巴茨拉’牡丹花瓣中可溶性糖含量的变化

Fig. 6 Changes of soluble sugar content of cut Itoh peony‘Bartzella’flowers during vase periods

图7 ‘巴茨拉’牡丹花瓣中SDH、CCO和ATP酶活性的变化

Fig. 7 Changes of SDH,CCO and ATPase activity of cut Itoh peony flowers during vase periods

参考文献 73

[1]	Aghdam M S, Jannatizadeh A, Luo Z S, Paliyath G. 2018. Ensuring sufficient intracellular ATP supplying and friendly extracellular ATP signaling attenuates stresses,delays senescence and maintains quality in horticultural crops during postharvest life. Trends in Food Science & Technology, 76:67-81.
[2]	Ahmadi N, Nergi D, Ali M. 2013. Effects of 1-MCP and ethylene on postharvest quality and expression of senescence-associated genes in cut rose cv. Sparkle. Scientia Horticulturae, 166:78-83. doi: 10.1016/j.scienta.2013.12.015 URL
[3]	Alice T, Antonio F, Paolo V, Giovanni S. 2011. Effects of abscisic acid on ethylene biosynthesis and perception in Hibiscus rosa-sinensis L. flower development. Journal of Experimental Botany, 62 (15):5437-5452. doi: 10.1093/jxb/err218 URL
[4]	Anuj S A, Gajera H P, Hirpara D G, Golakiya B A. 2019. Bacterial membrane destabilization with cationic particles of nano-silver to combat efflux-mediated antibiotic resistance in Gram-negative bacteria. Life Sciences, 230:178-187. doi: 10.1016/j.lfs.2019.05.072 URL
[5]	Azad A K, Ishikawa T, Ishikawa T, Sawa Y, Shibata H. 2008. Intracellular energy depletion triggers programmed cell death during petal senescence in tulip. Journal of Experimental Botany, 59 (8):2085-2095. doi: 10.1093/jxb/ern066 URL
[6]	Bradford M M. 1976. A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of Protein-Dye Binding. Analytical Biochemistry, 72:248-254. doi: 10.1016/0003-2697(76)90527-3 URL
[7]	Cai Lei, Zhang Xiao-hong, Shen Hong-xiang, Gao Jun-ping. 2002. Effects of ethylene and its inhibitors on flower opening and senescence of cut roses. Acta Horticulturae Sinica, 29 (5):467-472. (in Chinese)
	蔡蕾, 张晓红, 沈红香, 高俊平. 2002. 乙烯对不同切花月季品种开花和衰老的影响. 园艺学报, 29 (5):467-472.
[8]	Cao Xue-ling, Liu Fa-xian, Jin Li. 2014. Research of nano-silver colloids prepared by microwave-assisted synthesis method and its fresh-keeping of strawberry. Science and Technology of Food Industry, 35 (5):327-364. (in Chinese)
	曹雪玲, 刘发现, 金丽. 2014. 纳米银胶的制备及对草莓的保鲜性能研究. 食品工业科技, 35 (5):327-364.
[9]	Chen C X, Hussain N, Wang Y R, Li M T, Liu L, Qin M Z, Ma N, Gao J P, Sun X M. 2020. An ethylene-inhibited NF-YC transcription factor RhNF-YC 9 regulates petal expansion in rose. Horticultural Plant Journal, 6 (6):419-427. doi: 10.1016/j.hpj.2020.11.007 URL
[10]	Cheng Gui-ping, Li Fang, Li Wan-ping, Liu Jun, Xian Xi-jin, He Sheng-gen. 2013. Effects of nano-silver plus pulse treatment on some physiological and biochemical indexes of cut roses during vase period. Hubei Agricultural Sciences, 52 (6):1338-1342. (in Chinese)
	程桂平, 李芳, 黎婉萍, 刘军, 冼锡金, 何生根. 2013. 纳米银加脉冲处理对切花月季瓶插期间生理生化指标的影响. 湖北农业科学, 52 (6):1338-1342.
[11]	Concepcion M, Lizada C, Yang S F. 1979. A simple and sensitive assay for 1-aminocyclopropane-1-carboxylic acid. Analytical Biochemistry, 100 (1):140-145. doi: 10.1016/0003-2697(79)90123-4 URL
[12]	Doorn V W G. 2004. Is petal senescence due to sugar starvation? Plant Physiology, 134 (1):35-42. doi: 10.1104/pp.103.033084 URL
[13]	Dong Shuan-quan, Xiong Qian, Wang Chun-xing, Wang Hong-fei, Shao Xing-Feng, Li He-Sheng, Xu Feng. 2016. Sucrose maintain the energy and antioxidant activity in delaying the yellowing of broccoli. Acta Horticulturae Sinica, 43 (9):1825-1833. (in Chinese)
	董栓泉, 熊茜, 王春幸, 王鸿飞, 邵兴锋, 李和生, 许凤. 2016. 蔗糖在延缓青花菜黄化过程中维持其能量和抗氧化力. 园艺学报, 43 (9):1825-1833.
[14]	Fernandez-Maculet J C, Yang S F. 1992. Extraction and partial characterization of the ethylene-forming enzyme from apple fruit. Plant Physiology, 99 (2):751-754. doi: 10.1104/pp.99.2.751 URL
[15]	Gao Jun-ping, Zhang Xiao-hong, Huang Mian-jia, Ye Xin-min, Sun Zi-ran. 1997. A preliminary study on change patterns of ethylene production during flower opening and senescence in cut roses. Acta Horticulturae Sinica, 24 (3):274-278. (in Chinese)
	高俊平, 张晓红, 黄绵佳, 叶新民, 孙自然. 1997. 月季切花开花和衰老进程中乙烯变化类型初探. 园艺学报, 24 (3):274-278.
[16]	Gao Shu-lin, Zhang Chao, Du Dan-ni, Liu Ai-qing, Dong Li. 2015. Effect of glucose and ethylene on flower color and anthocyanin biosynthesis in tree peony‘Luoyang Hong’cut flower. Acta Horticulturae Sinica, 42 (7):1356-1366. (in Chinese)
	高树林, 张超, 杜丹妮, 刘爱青, 董丽. 2015. 乙烯和葡萄糖处理对‘洛阳红’牡丹切花花色和花青素苷合成的影响. 园艺学报, 42 (7):1356-1366.
[17]	Gao Yong, Wu Shao-jin. 1988. Effects of ethylene on the senescence and longevity of cutting flower. Plant Physiology Communication,(4):5-10. (in Chinese)
	高勇, 吴绍锦. 1988. 乙烯与切花的衰老及保鲜. 植物生理学通讯,(4):5-10.
[18]	Gong B, Huang S, Ye N, Yuan X, Ma H L. 2018. Pre-harvest ethylene control affects vase life of cut rose‘Carola’by regulating energy metabolism and antioxidant enzyme activity. Horticulture,Environment,and Biotechnology, 59 (6):835-845. doi: 10.1007/s13580-018-0053-8 URL
[19]	Hoppen C, Müller L, Albrecht A C, Groth G. 2019. The NOP- 1 peptide derived from the central regulator of ethylene signaling EIN2 delays floral senescence in cut flowers. Scientific Reports, 9 (1):1-8.
[20]	Huang Xin-min, Lin Qi-ling, Xian Xi-jin, Liu Ji-ping, Li Hong-mei, He Sheng-gen. 2012. Nano-silver treatments alleviated the harmful effects of exogenous ethylene on cut Roses. Acta Horticulturae Sinica, 39 (04):735-742. (in Chinese)
	黄新敏, 林启灵, 冼锡金, 刘季平, 李红梅, 何生根. 2012. 纳米银对瓶插月季切花乙烯作用的拮抗效应. 园艺学报, 39 (4):735-742.
[21]	Jiang Yue-ming. 2002. Response of postharvest fruits and vegetables to inhibitors of ethylene binding sites in relation to the technology of storage and transportation. Guihaia, 22 (6):547-552. (in Chinese)
	蒋跃明. 2002. 采后果蔬对乙烯受体抑制剂的响应及贮运保鲜技术的研究. 广西植物, 22 (6):547-552.
[22]	Jiang Yue-ming, Wang Hui, Yi Chun, Qu Hong-xia, Duan Xue-wu. 2016. Advance in energy generation and control of harvested horticultural crop. Journal of Guangzhou University(Natural Science Edition), 15 (4):1-7. (in Chinese)
	蒋跃明, 王慧, 易春, 屈红霞, 段学武. 2016. 采后园艺产品能量代谢与调控的研究进展. 广州大学学报(自然科学版), 15 (4):1-7.
[23]	Jin P, Zhu H, Wang J, Chen J J, Wang X L, Zheng Y H. 2013. Effect of methyl jasmonate on energy metabolism in peach fruit during chilling stress. Journal of the Science of Food and Agriculture, 93 (8):1827-1832. doi: 10.1002/jsfa.5973 URL
[24]	Kato M, Hyodo H. 1999. Purification and characterization of ACC oxidase and increase in its activity during ripening of pear fruit. Journal of the Japanese Society for Horticultural Science, 68 (3):551-557.
[25]	Kim J S, Kuk E, Yu K N, Kim J H. 2007. Antimicrobial effects of silver nanoparticles. Nanomedicine:Nanotechnology,Biology,and Medicine,(3):95-101.
[26]	Le Pan. 2014. Nano-silver composite agents on preservation of nanfeng citruses. Nanchang:Nanchang University. (in Chinese)
	乐攀. 2014. 纳米银复合保鲜剂对南丰蜜桔的保鲜研究. 南昌:南昌大学.
[27]	Li Hong-mei, Lin Yan-fei, Liu Chang-zhen, Huang Xin-min, Zhou Hou-gao, He Sheng-gen. 2012. Freshness-preserving effects of Nano-silver pre-treatments on cut lily flowers. Northern Horticulture,(8):166-169. (in Chinese)
	李红梅, 林燕飞, 刘昌镇, 黄新敏, 周厚高, 何生根. 2012. 纳米银预处理对麝香百合切花的保鲜效应研究. 北方园艺,(8):166-169.
[28]	Li Liu-ze-mu. 2016. Micropropagation of Paeonia‘Bartzella’-a hybrid between herbaceous and tree peony. Beijing:Beijing Forestry University. (in Chinese)
	李刘泽木. 2016. 牡丹与芍药组间‘Bartzella’的离体快繁技术研究. 北京: 北京林业大学.
[29]	Li Mei-ling, Lin Yu-zhao, Wang Hui, Lin Yi-xiong, Lin He-tong. 2019. Recent advances in the role and regulation of energy status in senescence of harvested fruits and vegetables. Food Science, 40 (9):290-295. (in Chinese)
	李美玲, 林育钊, 王慧, 林毅雄, 林河通. 2019. 能量状态在果蔬采后衰老中的作用及其调控研究进展. 食品科学, 40 (9):290-295.
[30]	Lin X H, Li H M, Lin S Q, Xu M L, Liu J P, Li Y, He S G. 2019. Improving the postharvest performance of cut spray‘Prince’carnations by vase treatments with nano-silver and sucrose. Journal of Pomology and Horticultural Science, 94 (4):513-521.
[31]	Liu Chang-yu, Chen Xun, Long Yu-qing, Chen Ya, Liu Xiang-dan, Zhou Ri-bao. 2019. Research advances in genes involved in ethylene biosynthesis and signal transduction during flower senescence. Biotechnology Bulletin, 35 (3):177-188. (in Chinese)
	刘畅宇, 陈勋, 龙雨青, 陈娅, 刘湘丹, 周日宝. 2019. 乙烯生物合成及信号转导途径中介导花衰老相关基因的研究进展. 生物技术通报, 35 (3):177-188.
[32]	Liu H, Jiang Y, Luo Y. 2006. A simple and rapid determination of ATP,ADP and AMP concentrations in pericarp tissue of litchi fruit by high performance liquid chromatography. Food Technology and Biotechnology, 44 (4):531-534.
[33]	Liu J P, He S G, Zhang Z Q, Cao J P, Lv P T, He S D, Cheng G P, Daryl C J. 2009. Nano-silver pulse treatment inhibit stem-end bacteria on cut gerbera cv. Ruikou flowers. Postharvest Biology and Technoiogy, 54:59-62.
[34]	Liu Jian-xin, Yu Xiao-nan. 2015. Grey correlation analysis between blooming stage and related phenophases and prediction model establishment of blooming stage herbaceous peony introduced in Beijing. Journal of Plant Resources and Environment, 24 (4):108-110. (in Chinese)
	刘建鑫, 于晓南. 2015. 北京地区引进观赏芍药花期与相关物候期的灰色关联分析及其花期预测模型建立. 植物资源与环境学报, 24 (4):108-110.
[35]	Liu Ya-jie, Chang Ping, Tang Dong-qin. 2015. Endogenous hormone contents of cut freesia flowers during vase holding. Bulletin of Botanical Research, 35 (2):218-224. (in Chinese)
	刘亚杰, 常苹, 唐东芹. 2015. 香雪兰切花瓶插过程中内源激素含量变化的研究. 植物研究, 35 (2):218-224.
[36]	Mahendra R, Alka Y, Aniket G. 2009. Silver nanoparticles as a new generation of antimicrobials. Biotechnology Advances, 27:76-83. doi: 10.1016/j.biotechadv.2008.09.002 URL
[37]	Marieke D, Lisa V D B, Dirk L. 2018. The Pivotal role of ethylene in plant growth. Trends in Plant Science, 23 (4):311-323. doi: 10.1016/j.tplants.2018.01.003 URL
[38]	Ma Xiang-Long, Wu Jing-xu, Liu Shao-hua. 2018. The development status and prospect of Itoh peony. China Flowers and Horticulture,(16):28-31. (in Chinese)
	马翔龙, 吴敬需, 刘少华. 2018. 伊藤牡丹发展现状与展望. 中国花卉园艺,(16):28-31.
[39]	Mo Yi-wei, Jin Wen-long, Liu Kai-dong. 2013. Determination of ATP,ADP and AMP concentrations in several tropical fruits by high performance liquid chromatography. Journal of Fruit Science, 30 (6):1077-1082. (in Chinese)
	莫亿伟, 金文龙, 刘锴栋. 2013. 高效液相色谱法测定几种热带水果ATP、ADP 和AMP 含量. 果树学报, 30 (6):1077-1082.
[40]	Nian Lin-ke, Meng Hai-yan, Su Xiao-lin, Shi Guo-an. 2017. Effects of adding chlorine dioxide to vase solution on fresh-keeping of tree peony cut flower. Plant Physiology Journal, 53 (11):2022-2030. (in Chinese)
	年林可, 孟海燕, 苏笑林, 史国安. 2017. 瓶插液添加二氧化氯对牡丹切花的保鲜效果. 植物生理学报, 53 (11):2022-2030.
[41]	Pang Li-zheng, Cheng Fang-yun. 2011. Grafting propagation of distant hybrids(Iton Hybrids)between Tree and Herbaceous peonies. China Flowers and Horticulture,(22):30-32. (in Chinese)
	庞利铮, 成仿云. 2011. 牡丹与芍药组间远缘杂种(伊藤杂种)的嫁接繁殖. 中国花卉园艺,(22):30-32.
[42]	Peng Gang-zhi, Li Wei, Gao Ben-wang, Song Zheng-jiang, Zhang Hai-ling. 2018. Comparative study on biological characteristics of ornamental peony introduced to Yichang. Hubei Forestry Science and Technology, 47 (1):18-21. (in Chinese)
	彭刚志, 李薇, 高本旺, 宋正江, 张海玲. 2018. 宜昌引种观赏牡丹生物学特性研究. 湖北林业科技, 47 (1):18-21.
[43]	Ruan Y L. 2014. Sucrose metabolism:gateway to diverse carbon use and sugar signaling. Annual Review of Plant Biology, 65:33-67. doi: 10.1146/annurev-arplant-050213-040251 URL
[44]	Schaller G E, Binder B M. 2017. SInhibitors of ethylene biosynthesis and signalingS//Binder B,Schaller E G. Ethylene signaling:methods and protocols,methods in molecular biology. New York:Humana Press.
[45]	Shi Guo-an, Guo Xiang-feng, Kong Xiang-sheng, Zhang Guo-hai, Bao Man-zhu. 2011. Respiration rate and endogenous hormone levels in relation to the flower development of tree peonies. Acta Horticulturae Sinica, 38 (2):105-112. (in Chinese)
	史国安, 郭香凤, 孔祥生, 张国海, 包满珠. 2011. 牡丹呼吸速率和内源激素含量变化与开花衰老的关系. 园艺学报, 38 (2):105-112.
[46]	Shi Guo-an, Yang Zheng-shen, Wang Chang-zhong, Jin Zhi-wei, Han Xiang-yang. 1997. Effects of temperature and chemical agents on ethylene release and storage quality of cut peony flowers. Northern Horticulture,(6):62-63. (in Chinese)
	史国安, 杨正申, 王长忠, 金志伟, 韩向阳. 1997. 温度和化学药剂对牡丹切花乙烯释放及贮藏品质影响. 北方园艺,(6):62-63.
[47]	Shi Guo-an, Guo Xiang-feng, Han Jian-guo, Sun Xian-ming, Yang Zheng-shen. 1999. A study on ethylene production and lipid peroxidization in florescence and flower senescence of Paeonia suffruticosa. Acta University Agricultural Boreali-occidentalis, 27 (5):50-53. (in Chinese)
	史国安, 郭香凤, 韩建国, 孙鲜明, 杨正申. 1999. 牡丹开花和衰老期间乙烯及脂质过氧化的研究. 西北农业大学学报, 27 (5):50-53.
[48]	Shi Guo-an, Guo Xiang-feng, Li Chung-li, Fan Bing-you, Shi jiang, Bao Man-zhu. 2010a. Changes on ethylene release and ACC content of peony flowers at different development stages. Acta Horticulturae Sinica, 37 (1):77-82. (in Chinese)
	史国安, 郭香凤, 李春丽, 范丙友, 施江, 包满珠. 2010a. 牡丹花枝不同发育时期各器官乙烯释放和ACC含量的变化. 园艺学报, 37 (1):77-82.
[49]	Shi Guo-an, Guo Xiang-feng, Zhang Guo-hai, Gao Shuang-cheng, Fan Bing-you, Bao Man-zhu. 2010b. Studies of vasing physicological characteristics of cut peony flowers in various development stages. Acta Horticulturae Sinica, 37 (3):449-456. (in Chinese)
	史国安, 郭香凤, 张国海, 高双成, 范丙友, 包满珠. 2010b. 不同发育时期牡丹切花瓶插生理特性的研究. 园艺学报, 37 (3):449-456.
[50]	Shi Guo-an, Guo Xiang-feng, Zhang Guo-hai, Bao Man-zhu. 2009. Analysis of sugar metabolism during florescence and flower senescence of tree peony petal. Acta Horticulturae Sinica, 36 (8):1184-1190. (in Chinese)
	史国安, 郭香凤, 张国海, 包满珠. 2009. 牡丹开花和衰老期间花瓣糖代谢的研究. 园艺学报, 36 (8):1184-1190.
[51]	Shi Jun-yan, Gao Li-pu, Zuo Jin-hua, Wang Qing, Fan Lin-lin, Lü Jia-yu. 2017. Effect of nano-Ag film packaging on preservation of cucumber. The food industry, 38 (1):109-112. (in Chinese)
	史君彦, 高丽朴, 左进华, 王清, 范林林, 吕佳煜. 2017. 纳米银保鲜膜包装对黄瓜保鲜效果的影响. 食品工业, 38 (1):109-112.
[52]	Shimizu-Yumoto H, Ichimura K. 2013. Postharvest characteristics of cut dahlia flowers with a focus on ethylene and effectiveness of 6-benzylaminopurine treatments in extending vase life. Postharvest Biology and Technology, 86:479-486. doi: 10.1016/j.postharvbio.2013.07.036 URL
[53]	Smith D R. 1999. Fertility in the Itoh Hybrid,yellow crown. Paeonia, 29 (4):7-8.
[54]	Smith D R. 2001. Updated hybridizers list of named and registered intersectional hybrids. Paeonia, 31 (3):3-4.
[55]	Song L L, Peng Y H. 2004. Effect of cold storage on sensitivity of cut lily to ethylene. Journal of Horticultural Science & Biotechnology, 79 (5):723-728.
[56]	Song L L, Jiang Y M, Gao H Y, Li C T, Liu H, You Y L, Sun J. 2006. Effects of adenosine triphosphate on browning and quality of harvested litchi fruits. American Journal of Food Technology, 1 (2):173-178. doi: 10.3923/ajft.2006.173.178 URL
[57]	Song L L, Liu H, You Y L, Sun J, Yi C, Li Y B, Jiang Y M, Wu J S. 2014. Quality deterioration of cut carnation flowers involves in antioxidant systems and energy status. Scientia Horticulturae, 170:45-52. doi: 10.1016/j.scienta.2014.02.035 URL
[58]	Sun Ju-fang, Cheng Fang-yun. 2007. Preliminary report on the introduction of intersectional hybrids between tree and herbaceous peonies. Chinese Landscape Architecture, 23 (5):51-54. (in Chinese)
	孙菊芳, 成仿云. 2007. 芍药与牡丹组间杂种引种栽培初报. 中国园林, 23 (5):51-54.
[59]	Sun Ju-zhi, Xiang Ji-qian, Yin Hong-qing, Chen Fei-fei, Li Hong-ying, Yang Yong-kang. 2019. Interactions between silver nanoparticle and plants:uptake,translocation,accumulation,transformation,bioeffect and underlying mechanism. Plant Physiology Joural, 55 (11):1570-1578. (in Chinese)
	孙举志, 向极钎, 殷红清, 陈菲菲, 李红英, 杨永康. 2019. 纳米银与植物的相互作用:吸收、迁移、累积、转化、生物效应及潜在机制. 植物生理学报, 55 (11):1570-1578.
[60]	Tang Ye, Xue Yu-qian, Zhang Xiu-xin, Wang Shun-li, Wu Zhao-hai, Xue Jing-qi. 2018. Study on the effect of sucrose and mannitol treatment on the quality of cut flower of paeonia lactiflora after harvest. Bulletin of Agricultural Science and Technology, 557 (5):153-155. (in Chinese)
	唐也, 薛玉前, 张秀新, 王顺利, 吴朝海, 薛璟祺. 2018. 不同浓度蔗糖及甘露醇处理对芍药切花采后品质影响的研究. 农业科技通讯, 557 (5):153-155.
[61]	Thao N P, Khan M I R, Thu N B A, Hoang X L T, Asgher M, Khan N A, Tran L S P. 2015. Role of ethylene and its cross talk with other signaling molecules in plant responses to heavy metal stress. Plant Physiology, 169:73-84. doi: 10.1104/pp.15.00663 URL
[62]	Wang H, Qian Z J, Ma S M, Zhou Y C, Patrick J W, Duan X W, Jiang Y M, Qu H X. 2013. Energy status of ripening and postharvest senescent fruit of litchi(Litchi chinensis Sonn.). BMC Plant Biology, 13 (1):55-70. doi: 10.1186/1471-2229-13-55 URL
[63]	Wang Kai-xuan, Wang Yi, Shi Tian, Ding Xi-ning, Yuan Jun-hui, Shi Guo-an, Hu Yong-hong. 2020. Physiological effets of rapamycin pretreatment on delaying the senescence of cut flower in tree peony ‘Luoyanghong’. Acta Horticulturae Sinica, 47 (10):1956-1968. (in Chinese)
	王凯轩, 王依, 史田, 丁熙柠, 袁军辉, 史国安, 胡永红. 2020. 雷帕霉素预处理延缓牡丹‘洛阳红’切花衰老的生理效应. 园艺学报, 47 (10):1956-1968.
[64]	Wang Yan-jie. 2013. Effect of glucose on ethylene biosynthesis and signal transduction in tree penoy cut flowers during postharvest senescence[Ph.D. Dissertation]. Beijing:Beijing Forstry University.
	王彦杰. 2013. 葡萄糖对牡丹切花采后乙烯生物合成及信号转导的影响[博士论文]. 北京: 北京林业大学.
[65]	Wang Zhe, Shi Guo-an, Ma Xue-qing, Fan Bing-you. 2014. The ethylene metabolism in flowers of chinese peony‘taohua feixue’during opening and senescence. Acta Horticulturae Sinica, 41 (11):2268-2274. (in Chinese)
	王哲, 史国安, 马雪情, 范丙友. 2014. 芍药‘桃花飞雪’开花衰老期间乙烯代谢生理机制的研究. 园艺学报, 41 (11):2268-2274.
[66]	Wu F, Zhang C, Wang X Q, Guo J, Dong L. 2017. Ethylene-influenced development of tree peony cut flowers and characterization of genes involved in ethylene biosynthesis and perception. Postharvest Biology and Technology, 125:150-160. doi: 10.1016/j.postharvbio.2016.11.014 URL
[67]	Wu Guo-xin, Cui Ling-hua, Liu Shao-hua, Yao-Fang, Si Shou-xia, Ren Shu-hui. 2011. Preliminary studing report of cultivating and demonstrating itoh hybrids introduced from abroad. Northern Horticulture,(24):67-71. (in Chinese)
	吴国新, 崔玲华, 刘少华, 姚方, 司守霞, 任叔辉. 2011. 国外伊藤杂种牡丹引进栽培示范研究. 北方园艺,(24):67-71.
[68]	Wu L, Ma N, Jia Y C, Feng M, Jiang C Z, Ma C, Gao J P. 2017. An ethylene-induced regulatory module delays flower senescence by regulating cytokinin content. Plant Physiology, 173:853-862. doi: 10.1104/pp.16.01064 URL
[69]	Xue Jing-qi, Yang Feng, Zuo Zhi-rui, Gao Jun-ping. 2011. Effect of ethylene and 1-MCP on ethylene production and related gene expression in floral tissues of roses. Acta Horticulturae Sinica, 38 (2):311-316. (in Chinese)
	薛璟褀, 杨丰, 左志锐, 高俊平. 2011. 乙烯和1-MCP的竞争性作用对月季花器官乙烯生成量及相关基因表达的影响. 园艺学报, 38 (2):311-316.
[70]	Yang Ruo-wen, Zhang Ping, Xue Yu-qian, Xue Jing-qi, Wang Shunli, Zhang Xiu-xin. 2018. Effects of ethylene on opening and senescence process of herbaceous peony cut flowers‘Banker Hill’and‘Yangfei Chuyu’. Acta Horticulturae Sinica, 45 (8):152-163. (in Chinese)
	杨若雯, 张萍, 薛玉前, 薛璟祺, 王顺利, 张秀新. 2018. 乙烯对‘班克海尔’和‘杨妃出浴’芍药切花开放和衰老的影响. 园艺学报, 45 (8):152-163.
[71]	Zhang C, Liu M, Fu J X, Wang Y J, Dong L. 2012. Exogenous sugars involvement in senescence and ethylene production of tree peony‘Luoyang Hong’cut flowers. Korean Journal of Horticultural Science and Technology, 30 (6):718-724. doi: 10.7235/hort.2012.12089 URL
[72]	Zhao D Q, Cheng M L, Tang W H, Liu D, Zhou S Y, Meng J S, Tao J. 2018. Nano-silver modifies the vase life of cut herbaceous peony(Paeonia lactiflora Pall.).flowers. Protoplasma, 255:1001-1013. doi: 10.1007/s00709-018-1209-1 URL
[73]	Zhuang Qian, Zhu Song-yan, Du Xiao-qi, Zhao Li-qun. 2011. Introduction of interspecific hybrids of Paeonia to northeast China. Journal of Northeast Forestry University, 39 (4):21-23. (in Chinese)
	庄倩, 朱松岩, 杜晓琪, 赵立群. 2011. 芍药属组间杂种引进东北地区栽培试验. 东北林业大学学报, 39 (4):21-23.