白姜花二倍体与四倍体切花形态与显微结构变化观察

doi:10.16420/j.issn.0513-353x.2021-0921

摘要/Abstract

摘要：

以白姜花（Hedychium coronarium）二倍体和四倍体切花为试验材料，观察其发育过程中的变化特征。结果表明：以小花花冠管发生弯颈和花瓣开始萎蔫为标志，小花发育进程可划分为6个阶段：苞裂期、初开期、盛开期、弯颈期、初萎期和萎蔫期。小花花冠管弯颈发生早于花瓣衰老，花冠管细胞衰老早于花瓣细胞。四倍体小花花冠管直径极显著大于二倍体，花冠管伸出苞片的比例显著低于二倍体。在瓶插期间，四倍体切花处于净吸水状态的时间比二倍体延长，花茎基部导管的堵塞频率变化较为平稳。四倍体花冠管弯颈发生、花冠管细胞和花瓣细胞的线粒体损伤均晚于二倍体。总体上，小花弯颈发生是白姜花切花衰老开启的标志；白姜花四倍体切花寿命显著长于二倍体，可能与四倍体具有较粗的花冠管、苞片对小花提供较强的支撑作用、花冠管和花瓣细胞的衰老推迟以及花茎具有较强的吸水能力和抗堵塞能力有关。

关键词: 白姜花, 四倍体, 切花, 衰老, 花冠管

Abstract:

With diploid and tetraploid Hedychium coronarium as test material，the developmental process of cut flowers were observed. Micrographic changes of the flower stem base，corolla tube and petals were identified by scanning and transmission electron microscopy. With the bent neck of corolla tube and initial wilting of petal as markers，the developmental process of florets was divided into six stages：bracts cracking stage，initial opening stage，blooming stage，neck bending stage，initial wilting stage，and wilting stage. Neck bending of the floret corolla tube preceded petal wilting，and the senescence of corolla tube cells arose earlier than the petal cells. The diameter of the corolla tube of the tetraploid was extremely significantly more than that of the diploid，and the ratio of the corolla tube length out of bracts to the total length of the tetraploid was significantly less than that of the diploid. During the whole vase life，the cut flowers of tetraploid were in the state of net water absorption for longer time and the basal vessel of cut flower stem were blocked more slightly than those of the diploid. The neck bending of the tetraploid floret arose later than that of the diploid，and damages of mitochondria of corolla tube and petal cells were posterior to those of the diploid. In total，neck bending of the corolla tube was the landmark for the beginning of senescence of H. coronarium floret. The significant longer longevity of the tetraploid cut flowers may be due to the thicker corolla tube and stronger support from bracts to florets，the postponed senescence of corolla tube and petal cells，and stronger water absorption and anti-blocking ability of the flower stems.

Key words: Hedychium coronarium, tetraploid, cut flower, senescence, corolla tube

中图分类号:

S682.19

张爱玲, 涂红艳, 肖望, 钟晓晴, 陆秋婵, 成丽萍, 林晓萍, 麦钰玲. 白姜花二倍体与四倍体切花形态与显微结构变化观察[J]. 园艺学报, 2023, 50(2): 345-358.

ZHANG Ailing, TU Hongyan, XIAO Wang, ZHONG Xiaoqing, LU Qiuchan, CHENG Liping, LIN Xiaoping, MAI Yuling. Morphology and Microstructure Characteristics of Diploid and Tetraploid Hedychium coronarium Cut Flowers[J]. Acta Horticulturae Sinica, 2023, 50(2): 345-358.

图/表 14

图1 白姜花小花花冠管弯颈部位

Fig. 1 The bent neck of corolla tube of floret of Hedychium coronarium

图2 白姜花花序（A）和小花（B）解剖图 a：穗状花序；b：苞片；c：初生苞片；d：小花花蕾；e：小花；f：花冠管；g：唇瓣；h：侧生退化雄蕊（翼瓣）；i：花冠裂片；j：柱头；k：花柱；l：子房；m：花药；n：花丝。

Fig. 2 Anatomic structure of inflorescence（A）and floret（B）of Hedychium coronarium a：Spike；b：Bract；c：Primary bracts；d：Floret bud；e：Floret；f：Corolla tube；g：Labellum；h：Lateral staminodes（Wing）；i：Corolla lobes；j：Stigma；k：Style；l：Ovary；m：Anther；n：Filament.

图3 白姜花小花不同发育阶段

Fig. 3 Different developmental stages of floret of Hedychium coronarium

图4 白姜花二倍体和四倍体小花发育过程形态变化规律 0 h：苞裂期；蓝色箭头：弯颈期；黄色箭头：初萎期；红色箭头：萎蔫期。

Fig. 3 Morphological changes of diploid and tetraploid florets of Hedychium coronarium in different developmental stages 0 h：Bracts cracking；Blue arrow：Neck bending stage；Yellow arrow：Initial wilting stage；Red arrow：Wilting stage.

表1 白姜花二倍体和四倍体小花和花序最佳观赏期和瓶插寿命

Table 1 Comparison of different developmental stages and inflorescence longevity of diploid and tetraploid florets of Hedychium coronarium

白姜花 Hedychium coronarium	小花Florets		花序Inflorescence
白姜花 Hedychium coronarium	观赏时长/h Ornamental duration	瓶插寿命/h Longevity	观赏时长/h Ornamental duration	瓶插寿命/h Longevity
二倍体Diploid	15.3 ± 2.9 a	23.8 ± 3.3 a	58.4 ± 3.2 a	70.8 ± 6.4 a
四倍体Tetraploid	23.7 ± 4.0 b	29.5 ± 4.1 b	83.8 ± 5.3 b	106.6 ± 7.1 b

图5 白姜花二倍体和四倍体切花花序瓶插期鲜质量的变化

Fig. 5 Changes of fresh weight of diploid and tetraploid cut flowers of Hedychium coronarium during vase periods

表2 白姜花二倍体和四倍体花冠管形态比较

Table 2 Comparison of corolla tube morphology between diploid and tetraploid of Hedychium coronarium

材料 Material	直径/cm Diameter	全长/cm Total length	伸出苞片外长/cm The length out of bracts	伸出苞片外长占全长比例/% The ratio of the length out of bracts to the total length
二倍体Diploid	0.23 ± 0.05 A	7.97 ± 0.61 a	2.75 ± 0.49 A	30.0 B
四倍体Tetraploid	0.43 ± 0.04 B	8.80 ± 0.62 b	2.03 ± 0.60 B	21.6 A

图6 白姜花二倍体（D）和四倍体（T）花冠管横截面显微结构光学显微镜观察结果

Fig. 6 Microstructures of corolla tube in diploid（D）and tetraploid（T）flowers of Hedychium coronarium with light microscope

图7 白姜花二倍体（D，d）和四倍体（T，t）小花（D，T）和花冠管横切面显微结构电镜扫描结果（d，t） 1、2、3分别为盛开期、弯颈期、初萎期。

Fig. 7 Development of diploid（D，d）and tetraploid（T，t）florets（D，T）and the corresponding cross sections（d，t） with scanning electron microscope 1，2，3 represent blooming stage，neck bending stage and initial wilting stage.

图8 白姜花二倍体和四倍体花冠管不同发育阶段亚细胞结构的透射电镜观察结果

Fig. 8 Subcellular structures of corolla tube cells of diploid and tetraploid flowers in different vase periods with transmission electron microscope

图9 白姜花二倍体和四倍体花瓣表皮细胞不同发育阶段表面形态变化过程的扫描电镜结果

Fig. 9 Changes of petal epidermal cell surfaces of diploid and tetraploid flowers in different developmental periods with scanning electron microscopic

图10 白姜花二倍体和四倍体花瓣表皮细胞不同发育阶段亚细胞结构透射电镜观察结果

Fig. 10 Subcellular structure changes of petal epidermal cells of diploid and tetraploid flowers in different developmental periods with transmission electron microscope

图11 白姜花二倍体和四倍体切花花茎基部不同瓶插时期显微结构电镜扫描结果

Fig. 11 Microstructures of flower stem bases in different vase periods in diploid and tetraploid cut flowers of Hedychium coronarium with scanning electron microscope

表3 白姜花二倍体和四倍体切花瓶插期间花茎基部导管的堵塞频率

Table 3 Blocking frequency of basal vessel of cut flower stem of diploid and tetraploid Hedychium coronarium during different vase periods %

瓶插时间/h Vase time	二倍体 Diploid	四倍体 Tetraploid
0	18.6 ± 0.29 a	15.1 ± 0.38 a
24	29.5 ± 0.47 b	15.1 ± 0.26 a
48	30.9 ± 0.37 b	16.4 ± 0.25 a
72	37.1 ± 0.42 c	19.9 ± 0.30 a

参考文献 34

[1]	Asit R, Sudipta J, Biswabhusan D, Ambika S, Basudeba K, Jeetendranath P, Pratap C P, Sanghamitra N, Namita M. 2019. Hedychium coronarium extract arrests cell cycle progression,induces apoptosis,and impairs migration and invasion in HeLa cervical cancer cells. Cancer Management and Research, 11:483-500. doi: 10.2147/CMAR.S190004 pmid: 30655700
[2]	Behera S, Kamila K P, Rout K K, Durga P B, Pratap C P, Soumendra K N. 2018. An efficient plant regeneration protocol of an industrially important plant,Hedychium coronarium J. Koenig and establishment of genetic & biochemical fidelity of the regenerants. Industrial Crops and Products, 126:58-68. doi: 10.1016/j.indcrop.2018.09.058 URL
[3]	Chen H, Yue Y C, Yu R C, Fan Y P. 2019. A Hedychium coronarium short chain alcohol dehydrogenase is a player in allo-ocimene biosynthesis. Plant Molecular Biology, 101 (3):297-313. doi: 10.1007/s11103-019-00904-z pmid: 31368003
[4]	Chen Hemin, Li Zuo, Ma Nan, Xiao Wenfang, Chen Heming, Lü Fubing, Li Zongyan, Zhu Genfa. 2022. Research advances on fresh-keeping technologies of orchid cut flowers and potted plants. Acta Horticulturae Sinica, 49 (12):2743-2760. (in Chinese) doi: 10.16420/j.issn.0513-353x.2022-0488
	陈和敏, 李佐, 马男, 肖文芳, 陈和明, 吕复兵, 李宗艳, 朱根发. 2022. 兰花切花保鲜及盆花品质保持技术研究进展. 园艺学报, 49 (12):2743-2760. doi: 10.16420/j.issn.0513-353x.2022-0488
[5]	Chiang C E W, Wong S K. 2015. Phytochemistry and pharmacology of ornamental gingers,Hedychium coronarium and Alpinia purpurata:a review. Journal of Integrative Medicine, 13 (6):368-379. doi: 10.1016/S2095-4964(15)60208-4 pmid: 26559362
[6]	Gantait S, Mandal N, Bhattachacharyya S, Kanti Das P. 2011. Induction and identification of tetraploids using in vitro colchicine treatment of Gerbera jamesonii Bolus cv. Sciella. Plant Cell,Tissue and Organ Culture, 106:485-493. doi: 10.1007/s11240-011-9947-1 URL
[7]	Hu Xiu, Wu Yong-qing, Ji Bing-bing, Huang Jia-qi, Zhou Jing, Liu Nian. 2018. A new Hedychium cultivar‘Jianbian’. Acta Horticulturae Sinica, 45 (3):607-608. (in Chinese)
	胡秀, 吴永清, 姬兵兵, 黄嘉琦, 周静, 刘念. 2018. 姜花新品种‘渐变’. 园艺学报, 45 (3):607-608. doi: 10.16420/j.issn.0513-353x.2017-0679
[8]	Hu Xiu, Zhong Yucheng, Tan Jiachuan. 2021. A new Hedychium cultivar‘Hanyue’. Acta Horticulturae Sinica, 48 (S2):2925-2926. (in Chinese)
	胡秀, 钟玉成, 谭嘉川. 2021. 姜花新品种‘寒月’. 园艺学报, 48 (S2):2925-2926.
[9]	Ke Y G, Abbas F, Zhou Y W, Yu R C, Yue Y C, Li X Y, Yu Y Y, Fan Y P. 2019. Genome-wide analysis and characterization of the Aux/IAA family genes related to floral scent formation in Hedychium coronarium. International Journal of Molecular Sciences, 20 (13):3235. doi: 10.3390/ijms20133235 URL
[10]	Li D M, Zhao C Y, Zhu G F, Xu Y C. 2019. Complete chloroplast genome sequence of Hedychium coronarium. Mitochondrial DNA Part B, 4 (2):2806-2807. doi: 10.1080/23802359.2019.1659114 URL
[11]	Li H B, Liu J P, Luo Z H, Joyce D, He S G. 2017. Nano-silver treatments reduced bacterial colonization and biofilm formation at the stem-ends of cut gladiolus‘Eerde’spikes. Postharvest Biology and Technology, 123:102-111. doi: 10.1016/j.postharvbio.2016.08.014 URL
[12]	Li He-ping. 2009. Plant microscope technology. Beijing: Science Press:67-89,228-231. (in Chinese)
	李和平. 2009. 植物显微技术. 北京: 科学出版社:67-89,228-231.
[13]	Lü Guo-sheng. 2011. Studies on the relationship between lignin and bent neck and aphid resistance in spray cut Chrysanthemum[M. D. Dissetation]. Nanjing: Nanjing Agricultural University. (in Chinese)
	吕国胜. 2011. 木质素与多头切花菊弯颈及蚜虫抗性的相关性研究[硕士论文]. 南京: 南京农业大学.
[14]	McCarthy E W, Landis J B, Kurti A, Lawhorn A J, Chase M W, Knapp S, Le Comber S C, Leitch A R, Litt A. 2019. Early consequences of allopolyploidy alter floral evolution in Nicotiana(Solanaceae). BMC Plant Biology, 19 (1):162. doi: 10.1186/s12870-019-1771-5
[15]	Mitter R. 2016. ROS are good. Trends in Plant Science, 22:11-19. doi: 10.1016/j.tplants.2016.08.002 URL
[16]	Mohanty P, Behera S, Swain S S, Barik D P, Naik S K. 2013. Micropropagation of Hedychium coronarium J. Koenig through rhizome bud. Physiology and Molecular Biology of Plants Physiolgy, 19 (4):605-610.
[17]	Perik R R J, Rezé D, Ferrante A, Van Doorn W G. 2014. Stem bending in cut Gerbera jamesonii flowers:effects of a pulse treatment with sucrose and calcium ions. Postharvest Biology and Technology, 98:7-13. doi: 10.1016/j.postharvbio.2014.06.008 URL
[18]	Perik R R J, Rezé D, Harkema H, Zhong Y, van Doorn W G. 2012. Bending in cut Gerbera jamesonii flowers relates to adverse water relations and lack of stem sclerenchyma development,not to expansion of the stem central cavity or stem elongation. Postharvest Biology and Technology, 74:11-18. doi: 10.1016/j.postharvbio.2012.06.009 URL
[19]	Panigrahy S K, Kumar A, Bhatt R J. 2020. Hedychium coronarium Rhizomes: Promising Antidiabetic and Natural Inhibitor of alpha-Amylase and alpha-Glucosidase. Journal of Dietary Supplements, 17 (1):81-87. doi: 10.1080/19390211.2018.1483462 pmid: 30325249
[20]	Ray A, Jena S, Dash B, Kar B, Halder T, Chatterjee T, Ghosh B, Panda Chander P, Nayak S, Mahapatra N. 2018. Chemical diversity,antioxidant and antimicrobial activities of the essential oils from Indian populations of Hedychium coronarium Koen. Industrial rops and Products, 112:352-362.
[21]	Rogers H J. 2013. From models to ornamentals;How is flower senescence regulated. Plant Molecular Biology, 82:563-574. doi: 10.1007/s11103-012-9968-0 URL
[22]	Shahri W, Tahir I. 2011. Flower senescence: Strategies and some associated events. Botanical Review, 77:152-184. doi: 10.1007/s12229-011-9063-2 URL
[23]	Shahri W, Tahir I. 2014. Flower senescence: some molecular aspects. Planta, 239 (2):277-297. doi: 10.1007/s00425-013-1984-z pmid: 24178586
[24]	Su J C, Nie Y, Zhao G, Cheng D, Wang R, Chen J, Zhang S H, Shen W B. 2019. Endogenous hydrogen gas delays petal senescence and extends the vase life of lisianthus cut flowers. Postharvest Biology and Technology, 147:148-155. doi: 10.1016/j.postharvbio.2018.09.018 URL
[25]	Suchitr K P, Awanish K, Renu B. 2020. Hedychium coronarium rhizomes:promising antidiabetic and natural inhibitor of α-amylase and α-glucosidase. Journal of Dietary Supplements, 17 (1):81-87. doi: 10.1080/19390211.2018.1483462 URL
[26]	Trivellini A, Cocetta G, Hunter D A, Vernieri P, Ferrante A. 2016. Spatial and temporal transcriptome changes occurring during flower opening and senescence of the ephemeral hibiscus flower,Hibiscus rosa-sinensis. Journal of Experimental Botany, 67 (20):5919-5931. pmid: 27591432
[27]	Tsai J Y, Wang T T, Huang P L, Do Y Y. 2021. Effects of developmental stages on postharvest performance of White Crane Orchid(Calanthe triplicata)inflorescences. Scientia Horticulturae, 281:1-9. https://doi.org/10.1016/j.scienta.2021.109988.
[28]	Tu H Y, Zhang A L, Xiao W, Lin Y R, Shi J H, Wu YW, Wu S T, Zhong C H, Mo S X. 2018. Induction and identification of tetraploid Hedychium coronarium through thin cell layer culture. Plant Cell Tissue and Organ Culture, 135 (3):395-406. doi: 10.1007/s11240-018-1472-z
[29]	Wagstaff C, Leverantz M K, Griffiths G, Thomas B, Chanasut U, Stead A D, Rogers H J. 2002. Cysteine protease gene expression and proteolytic activity during senescence of Alstroemeria petals. Journal of Experimental Botany, 367:233-240.
[30]	Wang Yi, Wang Kaixuan, Hu Siyuan, Zhou Shuang, Shi Guoan. 2021. Effects of ethylene metabolism and energy status on vase quality of cut Itoh peony‘Bartzella’flowers. Acta Horticulturae Sinica, 48 (6):1135-1149. (in Chinese)
	王依, 王凯轩, 胡思源, 周爽, 史国安. 2021. 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. 园艺学报, 48 (6):1135-1149. doi: 10.16420/j.issn.0513-353x.2020-0094
[31]	Xiao Wang, Tu Hong-yan, Zhang Ai-ling. 2016. Embryogenic callus induction and plant regeneration from Hedychium coronarium via somatic embryogenesis. Acta Horticulturae Sinica, 43 (8):1605-1612. (in Chinese)
	肖望, 涂红艳, 张爱玲. 2016. 白姜花胚性愈伤组织的诱导与植株再生体系的建立. 园艺学报, 43 (8):1605-1612. doi: 10.16420/j.issn.0513-353x.2015-0943
[32]	Yue Y C, Yu R C, Fan Y P. 2015. Transcriptome profiling provides new insights into the formation of floral scent in Hedychium coronarium. BMC Genomics, 16 (1):470. doi: 10.1186/s12864-015-1653-7 URL
[33]	Zhao D Q, Luan Y T, Xia X, Shi W B, Tang Y H, Tao J. 2020. Lignin provides mechanical support to herbaceous peony(Paeonia lactiflora Pall)stems. Horticulture Research, 7 (1):213. doi: 10.1038/s41438-020-00451-5
[34]	Zhou Yiwei, Xu Guoyu, Wang Qin, Yan Fulong, Yu Yunyi, Yu Rangcai, Fan Yanping. 2021. Genetic analysis and development of associated SSR markers of the flower color in F₁population of Hedychium coronarium‘COR01’× H.‘Jin’. Acta Horticulturae Sinica, 48 (10):1921-1933. (in Chinese) doi: 10.16420/j.issn.0513-353x.2021-0624
	周熠玮, 许国宇, 王琴, 严福龙, 玉云祎, 余让才, 范燕萍. 2021. ‘白姜花’×‘金姜花’杂交F₁代花色遗传分析及其相关SSR分子标记开发. 园艺学报, 48 (10):1921-1933. doi: 10.16420/j.issn.0513-353x.2021-0624