閉花受精是一種植物在關閉的花朵內完成自花授粉,並產生種子有性繁殖方式[1]閉鎖花(Cleistogamous Flower, CL)是閉花受精的關鍵,這類花朵不開放,授粉過程在花朵內部自發完成[2]。閉花受精現象分佈在50個228個的約693種植物中,常見於堇菜科鳳仙花科禾本科遠志科以及豆科等植物類群[3]

毛葉黃堇Viola pubescens)的(a)開放花與(b)閉鎖花

分類

編輯

閉花受精植物通常分為以下三類[2]

  • 完全閉花受精:所有花均為閉鎖花,植物不產生開放花。
  • 誘導閉花受精:在特定環境條件下誘發閉鎖花的產生。
  • 兩型閉花受精(Dimorphic Cleistogamy):同一植物在不同部位或不同時期同時產生閉鎖花和開放花(Chasmogamous Flower, CH)。

閉鎖花通常具有減少的花冠雄蕊數量[4]。相較於開放花鮮艷的花瓣和特殊氣味,閉鎖花花瓣發育不全或完全缺失,通常無色,且雄蕊數量減少,有時雄蕊退化為絲狀或原基狀[5][6][7]

假說

編輯

關於兩型閉花受精現象,存在多種假說[2]

  • 繁殖保障:閉鎖花是一種應對開放花授粉失敗的備份機制。在授粉不成功的情況下,閉鎖花可以通過自交保障種子的生產,延長生長季節的有性生殖時間[8][9][10][11]
  • 防止自交衰退:自交後若有害等位基因未能被清除,將增加群體遺傳漂變與自交衰退的風險[12][13]。閉鎖花主要依靠自交繁殖,而開放花通過異交產生更多具遺傳變異的後代,從而維持後代的遺傳多樣性,以適應環境變化[9][14][10][15]
  • 能量分配:閉鎖花的能量消耗較低,因為它們節省了產生更多花粉蜜腺體與花瓣的成本;而開放花較大,通常具有吸引傳粉昆蟲的功能,因此需要更多能量[10][16]。兩型閉花受精可以通過不同花型的比例調整能量分配,以優化其生殖效益[9][17][18][19]
  • 環境異質性:環境因素(如光周期光強溫度、土壤營養和水分等)決定了閉鎖花與開放花的產生,環境異質性促進了兩種花型的分化與選擇[6][20][21][15]

參考文獻

編輯
  1. ^ Kuhn M. Bemerkungen über Vandellia und den Blüten dimorphismus. Bot. Zeitung. 1867, 25: 65–67. 
  2. ^ 2.0 2.1 2.2 Li, Qiaoxia; Li, Youlong; Li, Jigang; Chen, Chenlong; Sun, Kun. Effects of photoperiods on the development of chasmogamous and cleistogamous flowers in <i>Viola monbeigii</i> and <i>V. dissecta</i>. Biodiversity Science. 2024, 32 (6): 23484. doi:10.17520/biods.2023484. 
  3. ^ Culley, Theresa M.; Klooster, Matthew R. The Cleistogamous Breeding System: A Review of Its Frequency, Evolution, and Ecology in Angiosperms. The Botanical Review. 2007-01, 73 (1): 1–30. doi:10.1663/0006-8101(2007)73[1:TCBSAR]2.0.CO;2. 
  4. ^ Uphof, J. C. Th. Cleistogamic flowers. The Botanical Review. 1938-01, 4 (1): 21–49. doi:10.1007/BF02869833. 
  5. ^ Wang, Yunjing; Ballard, Harvey E.; McNally, R. Ryan; Wyatt, Sarah E. Gibberellins are involved but not sufficient to trigger a shift between chasmogamous-cleistogamous flower types in Viola pubescens 1. The Journal of the Torrey Botanical Society. 2013-01, 140 (1): 1–8. doi:10.3159/TORREY-D-12-00044.1. 
  6. ^ 6.0 6.1 Li, Qiaoxia; Huo, Qingdi; Wang, Juan; Zhao, Jing; Sun, Kun; He, Chaoying. Expression of B-class MADS-box genes in response to variations in photoperiod is associated with chasmogamous and cleistogamous flower development in Viola philippica. BMC Plant Biology. 2016-12, 16 (1). doi:10.1186/s12870-016-0832-2. 
  7. ^ Li, Q; Li, J; Zhang, L; Pan, C; Yang, N; Sun, K; He, C. Gibberellins are required for dimorphic flower development in Viola philippica.. Plant science : an international journal of experimental plant biology. 2021-02, 303: 110749. PMID 33487338. doi:10.1016/j.plantsci.2020.110749. 
  8. ^ Le Corff, Josiane. Effects of light and nutrient availability on chasmogamy and cleistogamy in an understory tropical herb, Calathea micans (Marantaceae). American Journal of Botany. 1993-12, 80 (12): 1392–1399. doi:10.1002/j.1537-2197.1993.tb15383.x. 
  9. ^ 9.0 9.1 9.2 Culley, Theresa M. Reproductive Biology and Delayed Selfing in Viola pubescens (Violaceae), an Understory Herb with Chasmogamous and Cleistogamous Flowers. International Journal of Plant Sciences. 2002-01, 163 (1): 113–122. doi:10.1086/324180. 
  10. ^ 10.0 10.1 10.2 Culley, Theresa M.; Klooster, Matthew R. The Cleistogamous Breeding System: A Review of Its Frequency, Evolution, and Ecology in Angiosperms. The Botanical Review. 2007-01, 73 (1): 1–30. doi:10.1663/0006-8101(2007)73[1:TCBSAR]2.0.CO;2. 
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  12. ^ Masuda, Michiko; Yahara, Tetsukazu; Maki, Masayuki. Evolution of floral dimorphism in a cleistogamous annual, Impatiens noli‐tangere L. occurring under different environmental conditions. Ecological Research. 2004-11, 19 (6): 571–580. doi:10.1111/j.1440-1703.2004.00673.x. 
  13. ^ Ginwal, H. S. Inbreeding depression in Eucalyptus tereticornis Sm. due to cleistogamous flowering. New Forests. 2010-09, 40 (2): 205–212. doi:10.1007/s11056-010-9194-z. 
  14. ^ Ansaldi, Beth H.; Weber, Jennifer J.; Goodwillie, Carol; Franks, Steven J. Low levels of inbreeding depression and enhanced fitness in cleistogamous progeny in the annual plant Triodanis perfoliata. Botany. 2019-07, 97 (7): 405–415. doi:10.1139/cjb-2019-0022. 
  15. ^ 15.0 15.1 Sternberger, AL; Ruhil, AVS; Rosenthal, DM; Ballard, HE; Wyatt, SE. Environmental impact on the temporal production of chasmogamous and cleistogamous flowers in the mixed breeding system of Viola pubescens.. PloS one. 2020, 15 (3): e0229726. PMID 32160228. doi:10.1371/journal.pone.0229726. 
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  17. ^ Parra-Tabla, V; Munguía-Rosas, M; Campos-Navarrete, MJ; Ramos-Zapata, JA. Effects of flower dimorphism and light environment on arbuscular mycorrhizal colonisation in a cleistogamous herb.. Plant biology (Stuttgart, Germany). 2015-01, 17 (1): 163–8. PMID 25077675. doi:10.1111/plb.12223. 
  18. ^ Zhang, Li-Hui; Sun, Qi; Zhao, Ji-Min; Zhang, Yan-Wen. Plasticity in the reproductive strategy of a clonal cleistogamous species, Pseudostellaria heterophylla. Plant Ecology. 2018-12, 219 (12): 1493–1502. doi:10.1007/s11258-018-0896-1. 
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  20. ^ Seguí, J.; Lázaro, A.; Traveset, A.; Salgado-Luarte, C.; Gianoli, E. Phenotypic and reproductive responses of an Andean violet to environmental variation across an elevational gradient. Alpine Botany. 2018-04, 128 (1): 59–69. doi:10.1007/s00035-017-0195-9. 
  21. ^ Furukawa, T; Itagaki, T; Murakoshi, N; Sakai, S. Inherited dimorphism in cleistogamous flower production in Portulaca oleracea: a comparison of 16 populations growing under different environmental conditions.. Annals of botany. 2020-03-09, 125 (3): 423–431. PMID 31630158. doi:10.1093/aob/mcz167.