雾灵香花芥开花特性与繁育系统研究

doi:10.16420/j.issn.0513-353x.2021-1133

摘要/Abstract

摘要：

为揭示野生花卉雾灵香花芥（Hesperis oreophila）的开花特性与繁育类型，对由河北塞罕坝地区引种至河北保定地区的雾灵香花芥开花动态和花器官特征、花粉活力和花粉管行为、柱头可授性、花粉/胚珠比（P/O）、杂交指数（OCI）和人工授粉效果进行了观测与研究。结果表明：引种后的雾灵香花芥群体花期为4月上旬至5月下旬，持续时间55 ~ 60 d，单花持续时间14 ~ 17 d；雌雄蕊同期成熟，花完全展开时花粉和柱头活力较高，其花粉活力最高可达69.81%；自花授粉72 h内可完成受精过程；OCI、P/O和人工授粉试验结果显示其为自交为主的混合交配系统，需要传粉者。

关键词: 雾灵香花芥, 花, 花粉活力, 柱头可授性, 繁育系统

Abstract:

To reveal the floering characteristics and breeding types of wild flower Hesperis oreophila，the flowering dynamics and floral organ characteristics，pollen vitality and pollen tube behavior，stigma receptivity，the ratio of pollen and ovule（P/O），out crossing index（OCI）and artificial pollination test of H. oreophila which were introduced to Baoding area from Saihanba area of Hebei Province were observed and studied. The results showed that the flowering period of the introduced H. oreophila population was from early April to late May and lasted 55-60 days，with a single flowering period of 14 to 17 d. The pistils and stamens matured at the same time. Pollen and stigma had the highest vitality when the flower was fully expanded，and the pollen vitality was up to 69.81%. The fertilization process could be completed within 72 hours of self pollination. OCI，P/O and artificial pollination test results showed that it was a mixed mating system dominated by selfing and needed pollinators.

Key words: Hesperis oreophila, floral, pollen vitality, stigma receptivity, breeding system

中图分类号:

S682.19

董梦宇, 吴萌, 佟冠杰, 王金鑫, 陈新娜, 赵雪瑶, 李彦慧. 雾灵香花芥开花特性与繁育系统研究[J]. 园艺学报, 2023, 50(3): 607-619.

DONG Mengyu, WU Meng, TONG Guanjie, WANG Jinxin, CHEN Xinna, ZHAO Xueyao, LI Yanhui. Studies on Flowering Characteristics and Breeding System of Hesperis oreophila[J]. Acta Horticulturae Sinica, 2023, 50(3): 607-619.

图/表 12

图1 雾灵香花芥花序

Fig. 1 Inflorescence of Hesperis oreophila

图2 雾灵香花芥单花形态及精细解剖图 A1和A2：花侧视和俯视图；B：花离析；C：花纵剖；D：子房及胚珠；E1和E2：雄蕊和花药。

Fig. 2 Flower morphology and detailed anatomy of Hesperis oreophila A1 and A2：Flower side view，top view；B：Anatomy of the flower；C：Longitudinal profile of flower；D：Ovary and ovule；E1 and E2：Stamens and anthers.

表1 雾灵香花芥花器官特征统计

Table 1 Statistics of the floral organs of Hesperis oreophila

花器官 Floral organ	长度/mm Length	花器官 Floral organ	长度/mm Length
花柄长 Length of pedicel	9.78 ± 0.06	爪长 Length of claw	10.21 ± 0.05
萼片长 Length of calyx	7.87 ± 0.09	长花药长 Length of long anther	3.13 ± 0.06
萼片宽 Width of calyx	1.96 ± 0.03	长花丝长 Length of long filament length	6.81 ± 0.08
花冠直径 Corolla diameter	25.00 ± 0.75	短花药长 Length of short anther	3.39 ± 0.04
花瓣长 Length of petal	11.82 ± 0.09	短花丝长 Length of short filament length	2.64 ± 0.05
花瓣宽 Width of petal	9.15 ± 0.10	雄蕊长 Length of stamen	6.45 ± 0.08

图3 雾灵香花芥各发育时期（P1 ~ P9）的花形态（A）和柱头可授性（B）变化

Fig. 3 Changes of flower morphology（A）and stigma receptivity（B）in different developmental stages（P1-P9）of Hesperis oreophila

图4 雾灵香花芥花粉形态 A：群体视野；B：赤道面观；C：极面观；D：外壁纹饰。

Fig. 4 Pollen morphology of Hesperis oreophila A：Group vision；B：Equatorial view；C：Polar view；D：Outer wall decoration.

图5 雾灵香花芥花粉活力变化不同字母表示不同时期差异显著（P < 0.05）。下同。

Fig. 5 Changes of pollen vitality of Hesperis oreophila Different letters between different times meant significant differences at 0.05 level. The same below.

图6 离体培养下雾灵香花芥花粉萌发率及花粉管长度变化

Fig. 6 Changes of pollen germination and pollen tube length of Hesperis oreophila in vitro culture

图7 离体培养下雾灵香花芥花粉萌发情况

Fig. 7 Germination of pollen tube of Hesperis oreophila in vitro culture

图8 自然状态下雾灵香花芥荚果（A、B）和种子（C）

Fig. 8 Pods（A，B）and seeds（C）of Hesperis oreophila in natural state

图9 雾灵香花芥访花昆虫 A：斜斑鼓额食蚜蝇；B：狭带条胸蚜蝇；C：食蚜蝇；D：中华回条蜂；E：非洲蜂；F：胡蜂；G：菜粉蝶；H：柑橘凤蝶；I：黄钩蛱蝶。

Fig. 9 Flower-visiting insects of Hesperis oreophila flowers A：Scaeva pyrastri；B：Helophilus virgatus；C：Syrphidae；D：Habropoda sinensis；E：Africanized honeybee；F：Vespidae；G：Pieris rapae；H：Papilio xuthus；I：Polygonia c-aureum.

表2 雾灵香花芥不同传粉方式的坐果率及果实发育情况

Table 2 Fruiting rate and pods development of different pollination methods of Hesperis oreophila

处理方式 Processing method	坐果率/% Fruiting rate	荚果发育情况 Pods development	单果结实数 Number of single fruit
自然授粉（对照）Natural pollination（Control）	98.16 ± 1.14 a	饱满 Full	20 ~ 50
自花授粉Self-pollination	84.49 ± 2.05 b	较饱满 Relatively full	15 ~ 40
自然异花授粉Natural cross-pollination	34.38 ± 2.02 d	较饱满 Relatively full	10 ~ 30
无融合生殖Apomixis	2.25 ± 1.51 e	干瘪 Shriveled	4 ~ 8
同株异花授粉Strains cross-pollination	89.20 ± 3.50 b	饱满 Full	20 ~ 40
异株异花授粉Different strains cross-pollination	74.89 ± 3.33 c	饱满 Full	10 ~ 50

图10 雾灵香花芥花粉萌发、花粉管生长及受精过程 A：授粉2 h柱头表面；B：授粉24 h花柱中下部；C、D：授粉48 h子房上部；E、F：授粉72 h子房下部。PG：花粉粒；PT：花粉管；OU：胚珠。

Fig. 10 Pollen germination，pollen tube growth and fertilization process of Hesperis oreophila A：Surface of stigma after pollination in 2 h；B：Middle and lower part after pollination in 24 h；C，D：Upper ovary after pollination in 48 h；E，F：Lower ovary after pollination in 72 h. PG：Pollen grain；PT：Pollen tube；OU：Ovule.

参考文献 36

[1]	Bian Li-ming, Huang Dou, Zhang Xue-feng, Tong Xin-lei, Ye Dai-quan, Shi Ji-sen. 2020. Analysis on flowering phenology and synchronization indexes of Chinese fir clonal archive. Journal of Nanjing Forestry University(Natural Sciences Edition), 44 (6):207-212. (in Chinese)
	边黎明, 黄豆, 张学峰, 童鑫磊, 叶代全, 施季森. 2020. 杉木优树收集区无性系花期物候与同步性分析. 南京林业大学学报(自然科学版), 44 (6):207-212.
[2]	Bie Peng-fei, Tang Ting, Hu Jin-yao, Jiang Wei. 2018. Flowering phenology and breeding system of an endangered and rare species Urophysa rockii (Ranunculaceae). Acta Ecologica Sinica, 38 (11):3899-3908. (in Chinese)
	别鹏飞, 唐婷, 胡进耀, 蒋炜. 2018. 珍稀濒危植物距瓣尾囊草(Urophysa rochkii)的开花物候和繁育系统特性. 生态学报, 38 (11):3899-3908.
[3]	Cheng Min, Peng Yan. 2009. The influence of using ABA to high temperature dormancy characteristics of Agrostis stolonifera. Northern Horticulture,(9):157-160. (in Chinese)
	程敏, 彭燕. 2009. 施用外源脱落酸激素对匍茎翦股颖高温休眠特性的影响. 北方园艺,(9):157-160.
[4]	Cruden R W. 1977. Pollen ovule ratios:a conservative indicator of breeding systems in flowering plants. Evolution, 31 (1):32246.
[5]	Dafni A. 1992. Pollination ecology:a practical approach. New York: Oxford University Press.
[6]	Duan Shao-feng, Zhu Chun-yan, Yu Cheng-hua, Liang Yan-li. 2021. Study on the biological characteristics of pollination of Gentiana rigescens Franch. Journal of Yunnan Agricultural University(Natural Science), 36 (4):660-665. (in Chinese)
	段绍凤, 朱春艳, 余成华, 梁艳丽. 2021. 滇龙胆传粉生物学特征研究. 云南农业大学学报(自然科学), 36 (4):660-665.
[7]	Feng Xiao-lu, Chen Jun-qiang, You Jie, Dou Xiao, Dong Zhang-kai. 2021. Introduction adaptability and stress resistance of Hibiscus syriacus:research progress. Chinese Agricultural Science Bulletin, 37 (13):63-68. (in Chinese)
	冯小璐, 陈俊强, 游捷, 窦霄, 董章凯. 2021. 木槿引种适应性及抗逆性研究进展. 中国农学通报, 37 (13):63-68. doi: 10.11924/j.issn.1000-6850.casb2020-0134
[8]	Han De-liang, Liu Chun-xia, Han Lie-bao. 2009. Application of wide clover plant in landscaping. Northern Horticulture,(3):122-126. (in Chinese)
	韩德梁, 刘春霞, 韩烈保. 2009. 野生地被植物在园林绿化中的应用. 北方园艺,(3):122-126.
[9]	Han Yu-na, Liang Dong, Shen Yu-xian, Zhang Qing-hua, Zhang Su-ting, Jin Xiao-ming. 2016. Study on pollen viability and stigma receptivity of three alfalfa cultivars. Chinese Journal of Grassland, 38 (5):8-12. (in Chinese)
	韩玉娜, 梁冬, 申玉贤, 张菁华, 张苏婷, 金晓明. 2016. 3个苜蓿材料花粉活力和柱头可授性比较. 中国草地学报, 38 (5):8-12.
[10]	He Ya-ping, Liu Jian-quan. 2003. A review on recent advances in the studies of plant breeding system. Acta Phytoecologica Sinica,(2):151-163. (in Chinese)
	何亚平, 刘建全. 2003. 植物繁育系统研究的最新进展和评述. 植物生态学报,(2):151-163. doi: 10.17521/cjpe.2003.0024
[11]	Ji Qian-wen, Zheng Cheng-yang, Zhang Lei, Zeng Fa-xu. 2020. Stem radial growth dynamics of Pinus sylvestris var. mongolica and their relationship with meteorological factor in Saihanba,Hebei,China. Chinese Journal of Plant Ecology, 44 (3):257-265. (in Chinese) doi: 10.17521/cjpe.2019.0340
	季倩雯, 郑成洋, 张磊, 曾发旭. 2020. 河北塞罕坝樟子松径向生长动态变化及其与气象因子的关系. 植物生态学报, 44 (3):257-265. doi: 10.17521/cjpe.2019.0340
[12]	Jiang Hong-bo, Ding Qiong, Jia Gui-xia, Xie Hai-yan. 2004. The resources of herbaceous wild flowers in longtou mountain of Hebei Province and their utilization in landscape gardening. Scientia Silvae Sinicae,(6):102-109. (in Chinese)
	姜洪波, 丁琼, 贾桂霞, 谢海燕. 2004. 河北省龙头山区野生草本花卉植物资源及园林应用. 林业科学,(6):102-109.
[13]	Kudo G. 2003. Anther arrangement influences pollen deposition and removal in hermaphrodite flowers. Functional Ecology, 17:349-355. doi: 10.1046/j.1365-2435.2003.00736.x URL
[14]	Li Sen, Liu Juan, Gong Feifei, Wang Nan, Du Wei, Zhang Lingling, Kang Xiuping, Xing Guoming. 2021. Studies on flowering characteristics and breeding system of Hemerocallis citrina‘Datong Huanghua’. Acta Horticulturae Sinica, 48 (8):1541-1551. (in Chinese) doi: 10.16420/j.issn.0513-353x.2020-1024
	李森, 刘娟, 公菲菲, 王楠, 杜崴, 张玲玲, 亢秀萍, 邢国明. 2021. 黄花菜‘大同黄花’开花特性与繁育系统研究. 园艺学报, 48 (8):1541-1551. doi: 10.16420/j.issn.0513-353x.2020-1024
[15]	Liu Fen, Li Quan-jian, Wang Cai-xia, Lian Jing-jing, Tian Min. 2013. Floral characteristics and breeding systems of an endangered species Cypripedium japonicum. Scientia Silvae Sinicae, 49 (1):53-60. (in Chinese)
	刘芬, 李全健, 王彩霞, 连静静, 田敏. 2013. 濒危植物扇脉杓兰的花部特征与繁育系统. 林业科学, 49 (1):53-60.
[16]	Ma Sheng-jun, Tan Dun-yan. 2007. Phenology and sex expression of Junngar desert ephemerals Neotorularia korolkovii and Isatis violascens(Brassicaceae). Acta Ecologica Sinica,(2):486-496. (in Chinese)
	马生军, 谭敦炎. 2007. 短命植物甘新念珠芥(Neotorularia korolkovii)和宽翅菘蓝(Isatis violascens)的物候与性表达特征. 生态学报,(2):486-496.
[17]	Mai Jin-mei, Niu Dou, Tang Xiao-gang, Liu Hang, Zhang Ai-qin. 2019. Adaptability of floral syndrome and pollination pattern of Descurainia sophia. Arid Zone Research, 36 (1):169-175. (in Chinese)
	买金梅, 牛豆, 汤小港, 刘航, 张爱勤. 2019. 播娘蒿花部综合征与传粉模式的适应性研究. 干旱区研究, 36 (1):169-175.
[18]	Majetic C J, Raguso R A, Ashman T L. 2009. The sweet smell of success:floral scent affects pollinator attraction and seed fitness in Hesperis matronalis. Functional Ecology, 23 (3):480-487. doi: 10.1111/fec.2009.23.issue-3 URL
[19]	Peng De-li, Zhang Zhi-qiang, Niu Yang, Song Bo, Sun Hang, Li Zhi-min. 2012. Advances in the studies of reproductive strategies of alpine plants. Biodiversity Science, 20 (3):286-299. (in Chinese) doi: 10.3724/SP.J.1003.2012.14039
	彭德力, 张志强, 牛洋, 杨扬, 宋波, 孙航, 李志敏. 2012. 高山植物繁殖策略的研究进展. 生物多样性, 20 (3):286-299. doi: 10.3724/SP.J.1003.2012.14039
[20]	Ren Ming-xun. 2009. Intrafloral stamen differentiations and their adaptive significances. Chinese Journal of Plant Ecology, 33 (1):222-236. (in Chinese) doi: 10.3773/j.issn.1005-264x.2009.01.024
	任明迅. 2009. 花内雄蕊分化及其适应意义. 植物生态学报, 33 (1):222-236. doi: 10.3773/j.issn.1005-264x.2009.01.024
[21]	Sun Yi, Li Xi, Ding Suqin, Shi Yimin, Tang Dongqin. 2018. Studies on flowering characteristics and breeding system of Freesia hybrida. Acta Horticulturae Sinica, 45 (2):299-308.
	孙忆, 李玺, 丁苏芹, 史益敏, 唐东芹. 2018. 小苍兰开花特性与繁育系统研究. 园艺学报, 45 (2):299-308. (in Chinese) doi: 10.16420/j.issn.0513-353x.2017-0393 URL
[22]	Susko D J, Clubb M. 2008. Pollination effects on patterns of ovule fate in Hesperis matronalis(Brassicaceae). Botany-botanique, 86 (5):466-474. doi: 10.1139/B08-017 URL
[23]	Tang Qiang, Peng Yan. 2008. Summer dormancy in wild Poapratensis L. in subtropical regions. Northern Horticulture,(7):159-161. (in Chinese)
	汤强, 彭燕. 2008. 亚热带地区野生草地早熟禾夏季休眠特性研究. 北方园艺,(7):159-161.
[24]	Tian Lin-lin, Liu Jia, Li Yan-hui. 2018. Effects of different treatments on seed germination characteristics of Hesperis oreophila. Seed, 37 (8):30-35. (in Chinese)
	田琳琳, 刘佳, 李彦慧. 2018. 不同处理对雾灵香花芥种子萌发特性的影响. 种子, 37 (8):30-35.
[25]	Wang He-dong, Sun Zhu-biao, Yin Feng-qin. 2007. Investigation and introduction and domestication of wild flowers suitable for flower borders in Chengde area. The Journal of Hebei Forestry Science and Technology,(3):20-24. (in Chinese)
	王贺东, 孙柱彪, 尹凤琴. 2007. 承德地区花境适用野生花卉调查及引种驯化. 河北林业科技,(3):20-24.
[26]	Weeks E L, Frey F M. 2007. Seed production and insect visitation rates in Hesperis matronalis are not affected by floral symmetry. International Journal Plant Sciences, 168 (5):611-617.
[27]	Wu Xue-lian, Tan Dun-yan. 2007. Floral characters and breeding systems in the dimorphic annual plant Diptychocarpus strictus(Cruciferae). Acta Phytotaxonomica Sinica,(4):538-550. (in Chinese)
	吴雪莲, 谭敦炎. 2007. 异果芥的花部综合征及其繁育系统. 植物分类学报,(4):538-550.
[28]	Xu Li-jun. 2019. Study on introduction and propagation techniques of new landscape plants[M. D. Dissertation]. Beijing: Chinese Academy of Forestry. (in Chinese)
	许丽君. 2019. 新型园林植物引种及其繁殖技术研究[硕士论文]. 北京: 中国林业科学研究院.
[29]	Yang J Q, Fan Y L, Jiang X F, Li Q J, Zhu X F. 2018. Correlation between the timing of autonomous selfing and floral traits:a comparative study from three selfing Gentianopsis species(Gentianaceae). Scientific Reports, 8 (1):1-7.
[30]	Yang Jiahui, Wu Ting, Zhu Jun, Yang Shuhua, Zhao Xin, Ge Hong, Wu Ronghua, Jia Ruidong. 2021. Studies on flowering characteristics and breeding system of Paphiopedilum dianthum. Acta Horticulturae Sinica, 48 (5):1002-1012. (in Chinese) doi: 10.16420/j.issn.0513-353x.2020-0809 URL
	杨佳慧, 吴婷, 朱俊, 杨树华, 赵鑫, 葛红, 武荣花, 贾瑞冬. 2021. 长瓣兜兰开花特性与繁育系统研究. 园艺学报, 48 (5):1002-1012. doi: 10.16420/j.issn.0513-353x.2020-0809 URL
[31]	Yu Cheng-hua, Zhu Chun-yan, Duan Shao-feng, Yang Qiu-xiong, Han Jun, Liang Yan-li. 2021. Flowering phenology and mating system of Gentiana rigescens. Chinese Journal of Tropical Crops, 42 (6):1639-1645. (in Chinese)
	余成华, 朱春艳, 段绍凤, 杨秋雄, 韩俊, 梁艳丽. 2021. 滇龙胆开花物候与交配系统研究. 热带作物学报, 42 (6):1639-1645.
[32]	Yu Qin-lan, Liu Shou-zan, Xu Renximou, Pan Chenhang, Yan Yiming, Wang Yiying, Xia Guohua. 2021. Population structure and breeding systems of rare and endangered Gymnospermium kiangnanense(Berberidaceae). Acta Horticulturae Sinica, 48 (3):539-552. (in Chinese)
	虞钦岚, 刘守赞, 徐韧析谋, 潘晨航, 颜忆铭, 王义英, 夏国华. 2021. 珍稀濒危植物江南牡丹草种群结构和繁育系统研究. 园艺学报, 48 (3):539-552. doi: 10.16420/j.issn.0513-353x.2020-0474 URL
[33]	Zhang C, Zhou G Y, Yang Y P, Duan Y W. 2014. Better than nothing:evolution of autonomous selfing under strong inbreeding depression in an alpine annual from the Qinghai-Tibet Plateau. Journal of Systematics and Evolution, 52 (3):363-367. doi: 10.1111/jse.12082 URL
[34]	Zhang Chan, An Yu-meng, Yun JÄSCHKE, Wang Lin-lin, Zhou Zhi-li, Wang Li-ping, Yang Yong-ping, Duan Yuan-wen. 2020. Processes on reproductive ecology of plant species in the Qinghai-Xizang Plateau and adjacent highlands. Chinese Journal of Plant Ecology, 44 (1):1-21. (in Chinese) doi: 10.17521/cjpe.2019.0296
	张婵, 安宇梦, Yun JÄSCHKE, 王林林, 周知里, 王力平, 杨永平, 段元文. 2020. 青藏高原及周边高山地区的植物繁殖生态学研究进展. 植物生态学报, 44 (1):1-21. doi: 10.17521/cjpe.2019.0296
[35]	Zhang Li-ming. 2010. On the significance and plant allocation principles in the landscape construction in the urban ecological system. Value Engineering, 29 (4):157. (in Chinese)
	张黎明. 2010. 论城市生态系统中园林建设的重要性及植物配置原则. 价值工程, 29 (4):157.
[36]	Zhai Xin-ying, Ma Deng-tan, Wang Cheng-zhang, Yan Xue-bing. 2012. Effects of altitude on the mating system of alpine plant. Pratacultural Science, 29 (4):640-645. (in Chinese)
	翟新莹, 马登坦, 王成章, 严学兵. 2012. 海拔对高山植物交配系统的影响. 草业科学, 29 (4):640-645.