不同水陆生境下入侵种喜旱莲子草表型可塑性变异的发生与植物激素信号的关系
Relationship between the Development of Phenotypically Plastic Variation and Phytohormone Signaling of Alternanthera philoxeroides (Mart.) Griseb. under Different Water Habitats
DOI: 10.12677/BR.2014.34020, PDF, HTML,  被引量    国家自然科学基金支持
作者: 高乐旋：上海辰山植物园，上海
关键词: 喜旱莲子草；表型可塑性变异；植物激素；Alternanthera philoxeroides (Mart.) Griseb.； Phenotypically Plastic Variation； Phytohormone
摘要: 喜旱莲子草(Alternanthera philoxeroides (Mart.) Griseb.)是一种恶性入侵植物，依赖极强的表型可塑性成功入侵不同水陆生境。为揭示喜旱莲子草响应生境条件变异发生表型可塑性反应的相关信号通路，作者在模拟不同水陆生境的同质园环境下，利用多种植物激素及其抑制剂设计了一系列“正反”处理实验，比较喜旱莲子草形态特征可塑性变异式样在不同处理下的异同。结果表明：从陆地到水生环境后，喜旱莲子草响应环境变异发生快速的节间伸长、髓腔扩大的适应性可塑性反应；节间伸长和髓腔扩大的形态可塑性反应都需要乙烯的参与，而赤霉素仅参与节间伸长的形态可塑性反应，生长素仅参与髓腔扩大的形态可塑性反应。这些结果说明，环境因子主要通过乙烯、赤霉素和生长素信号通路介导了水陆生境变异下喜旱莲子草表型可塑性变异的发生。
Abstract: Alternanthera philoxeroides (Mart.) Griseb. is a malignant invasive weed in China. It has successfully invaded both aquatic and terrestrial habitats through phenotypic plasticity. To understand the signaling pathways potentially responsible for the development of phenotypically plastic variation in response to changing environment, A. philoxeroides were treated by a variety of phytohormones and their inhibitors under two common gardens, which respectively simulated the typical aquatic and terrestrial habitats colonized by A. philoxeroides in natural environments, then morphological variations among plants under different treatments were compared. The results showed that pond habitat promoted rapid elongation of internodes and further expansion of stem pith cavity in A. philoxeroides, contributing to their adaptation to submergence; ethylene was involved in both internode elongation and stem pith cavity expansion process, gibberellin was only involved in the internode elongation process, and auxin was only involved in the stem pith cavity expansion process. These results suggest that the ethylene, gibberellin and auxin signaling pathways are potentially responsible for the development of phenotypically plastic variation of A. philoxeroides in responding to changing water habitat.
文章引用：高乐旋. 不同水陆生境下入侵种喜旱莲子草表型可塑性变异的发生与植物激素信号的关系[J]. 植物学研究, 2014, 3(4): 155-163. http://dx.doi.org/10.12677/BR.2014.34020

1. 引言

外来种入侵目前已成为一个世界性的生态问题，不仅破坏入侵地生态系统的结构和功能、影响土著种的生存、造成区域生物多样性丧失、威胁生态安全，而且在全球范围内造成了巨大的经济损失[1] -[3] 。因此，世界各国都已将外来种入侵列入亟待解决的重大问题之一，投入大量人力和资金寻找控制生物入侵的策略，但迄今为止尚未发展出切实有效的控制和管理办法，其主要原因之一是对入侵种的入侵机制及其生物学基础缺乏深入了解。通常认为，广泛的生态适应性是入侵生物的普遍特征，因此，研究入侵种主要适应性特征的发生机制是揭示入侵机制的关键，也是有针对性的发展外来种控制方法的重要基础。

喜旱莲子草(Alternanthera philoxeroides (Mart.) Griseb.)又名空心莲子草、水花生、革命草，原产于南美洲，目前其入侵范围已扩大到世界各国的温带和热带地区[4] 。于20世纪30年代作为军马饲料引入中国，50年代后作为猪羊饲料被广泛引种到南方许多地区，自80年代开始快速的自然逸生扩散，广泛分布于长江流域和南方各省的河流、湿地、农田和公共绿地等多种生境，成为一种恶性入侵杂草，对入侵地的生态和经济造成极大负面影响[5] 。喜旱莲子草在入侵地很少产生有活力的种子，主要通过茎段或根段进行营养繁殖，因此遗传多样性很低[6] [7] 。喜旱莲子草适应性广、抗逆性强，可以在从水体到陆地多种不同水分生境中生存，表现“水陆两栖”的特点[5] -[8] 。前人基于形态和发育式样的比较研究发现喜旱莲子草能够随生境条件的改变发生适应性表型变异，提出表型可塑性能力是决定其适应性和入侵力的关键特征[5] [7] -[9] ，但对其表型可塑性变异的发生机制和快速适应不同生境的机制缺乏深入研究，这在一定程度上阻碍了对其入侵机制的了解。

参考文献

[1] 李博, 徐炳生, 陈家宽 (2001) 从上海外来杂草区系剖析植物入侵的一般特征. 生物多样性, 9, 217-219.
[2] Lodge, D.M. (1993) Biological invasions—Lessons for ecology. Trends in Ecology & Evolution, 8, 133-137.
[3] Pimentel, D., McNair, S., Janecka, J., Wightman, J., Simmonds, C., O’Connell, C., Wong, E., Russel, L., Zern, J., Aquino, T. and Tsomondo, T. (2001) Economic and environmental threats of alien plant, animal, and microbe inva- sions. Agriculture Ecosystems & Environment, 84, 1-20.
[4] Julien, M.H., Skarratt, B. and Maywald, G.F. (1995) Potential geographical-distribution of alligator weed and its bio- logical-control by Agasicles hygrophila. Journal of Aquatic Plant Management, 33, 55-60.
[5] 马瑞燕, 王韧 (2005) 喜旱莲子草在中国的入侵机理及其生物防治. 应用与环境生物学报, 11, 246-250.
[6] Julien, M.H. and Stanley, J.N. (1999) The management of a alligator weed, a challenge for the new millennium. Pro- ceedings of the 10th Biennial Noxious Weed Conference, New South Wales Department of Agriculture, Ballina, 20-22 July 1999, 2-13.
[7] Geng, Y.P., Pan, X.Y., Xu, C.Y., Zhang, W.J., Li, B., Chen, J.K., Lu, B.R. and Song, Z.P. (2007) Phenotypic plasticity rather than locally adapted ecotypes allows the invasive alligator weed to colonize a wide range of habitats. Biological Invasions, 9, 245-256.
[8] 潘晓云, 耿宇鹏, Alejandro, S.O.S.A, 张文驹, 李博, 陈家宽 (2007) 入侵植物喜旱莲子草——生物学、生态学及管理. 植物分类学报, 45, 884-900.
[9] Geng, Y.P., Pan, X.Y., Xu, C.Y., Zhang, W.J., Li, B. and Chen, J.K. (2006) Phenotypic plasticity of invasive Alternanthera philoxeroides in relation to different water availability, compared to its native congener. Acta Oe-cologica, 30, 380-385.
[10] Bradshaw, A.D. (1965) Evolutionary significance of phenotypic plasticity in plants. Advances in Genetics, 13, 115-155.
[11] van Kleunen, M. and Fischer, M. (2005) Constraints on the evolution of adaptive phenotypic plasticity in plants. New Phytologist, 166, 49-60.
[12] Grant-Downton, R.T. and Dickinson, H.G. (2006) Epigenetics and its implications for plant biology 2. The “epigenetic epiphany”: Epigenetics, evolution and beyond. Annals of Botany, 97, 11-27.
[13] Marden, J.H. (2008) Quantitative and evolutionary biology of alternative splicing: how changing the mix of alternative transcripts affects phenotypic plasticity and reaction norms. Heredity, 100, 111-120.
[14] Venturini, L., Ferrarini, A., Zenoni, S., Tornielli, G.B., Fasoli, M., Dal Santo, S., Minio, A., Buson, G., Tononi, P., Zago, E.D., Zamperin, G., Bellin, D., Pezzotti, M. and Delledonne, M. (2013) De novo transcriptome cha-racterization of Vitis vinifera cv. Corvina unveils varietal diversity. BMC Genomics, 14, 41.
[15] Landry, C.R., Oh, J., Hartl, D.L. and Cavalieri, D. (2006) Genome-wide scan reveals that genetic variation for tran- scriptional plasticity in yeast is biased towards multi-copy and dispensable genes. Gene, 366, 343-351.
[16] Smith, S., Bernatchez, L. and Beheregaray, L.B. (2013) RNA-seq analysis reveals extensive transcriptional plasticity to temperature stress in a freshwater fish species. BMC Genomics, 14, 375.
[17] Stern, S., Dror, T., Stolovicki, E., Brenner, N. and Braun, E. (2007) Genome-wide transcriptional plasticity underlies cellular adaptation to novel challenge. Molecular Systems Biology, 3, 106.
[18] Pigliucci, M. (2001) Phenotypic plasticity: Beyond nature and nurture. Johns Hopkings University Press, Baltimore.
[19] Bailey-Serres, J. and Voesenek, L. (2008) Flooding stress: Acclimations and genetic diversity. Annual Review of Plant Biology, 59, 313-339.
[20] Bailey-Serres, J. and Voesenek, L. (2010) Life in the balance: A signaling network controlling survival of flooding. Current Opinion in Plant Biology, 13, 489-494.
[21] Pierik, R., Millenaar, F.F., Peeters, A.J.M. and Voesenek, L. (2005) New perspectives in flooding research: the use of shade avoidance and Arabidopsis thaliana. Annals of Botany, 96, 533-540.
[22] Carr, S.M., Seifert, M., Delabaere, B. and Jaffe, M.J. (1995) Pith autolysis in herbaceous dicotyledonous plants—A physiological ecological study of pith autolysis under native conditions with special attention to the wild plant Impa- tienscapensis Meerb. Annals of Botany, 76, 177-189.
[23] Tao, Y., Chen, F., Wan, K.Y., Li, X.W. and Li, J.Q. (2009) The structural adaptation of aerial parts of invasive Alter- nanthera philoxeroides to water regime. Journal of Plant Biology, 52, 403-410.
[24] Gao, L.X., Geng, Y.P., Li, B., Chen, J.K. and Yang, J. (2010) Genome-wide DNA methylation alterations of Alternan- thera philoxeroides in natural and manipulated habitats: Implications for epigenetic regulation of rapid responses to environmental fluctuation and phenotypic variation. Plant Cell and Environment, 33, 1820-1827.
[25] Schlichting, C.D. (1986) The evolution of phenotypic plasticity in plants. Annual Review of Ecology and Systematics, 17, 667-693.
[26] He, C.J., Morgan, P.W. and Drew, M.C. (1996) Transduction of an ethylene signal is required for cell death and lysis in the root cortex of maize during aerenchyma formation induced by hypoxia. Plant Physiology, 112, 463-472.
[27] Drew, M.C., He, C.J. and Morgan, P.W. (2000) Programmed cell death and aerenchyma formation in roots. Trends in Plant Science, 5, 123-127.
[28] Hattori, Y., Nagai, K., Furukawa, S., Song, X.J., Kawano, R., Sakakibara, H., Wu, J.Z., Matsumoto T, Yoshimura A, Kitano, H., Matsuoka, M., Mori, H. and Ashikari, M. (2009) The ethylene response factors SNORKEL1 and SNORKEL2 allow rice to adapt to deep water. Nature, 460, 1026-1031.
[29] Vreeburg, R.A.M., Benschop, J.J., Peeters, A.J.M., Colmer, T.D., Ammerlaan, A.H.M., Staal, M., Elzenga, T.M., Staals, R.H.J., Darley, C.P., McQueen-Mason, S.J. and Voesenek, L.A.C.J. (2005) Ethylene regulates fast apoplastic acidification and expansin A transcription during submergence-induced petiole elongation in Rumex palustris. Plant Journal, 43, 597-610.
[30] Fukao, T. and Bailey-Serres, J. (2008) Ethylene—A key regulator of submergence responses in rice. Plant Science, 175, 43-51.
[31] Jackson, M.B. (2008) Ethylene-promoted elongation: An adaptation to submergence stress. Annals of Botany, 101, 229-248.
[32] Shieh, M.W. and Cosgrove, D.J. (1998) Expansins. Journal of Plant Research, 111, 149-157.
[33] Vriezen, W.H., De Graaf, B., Mariani, C. and Voesenek, L. (2000) Submergence induces expansin gene expression in flooding-tolerant Rumex palustris and not in flooding-intolerant R. acetosa. Planta, 210, 956-963.
[34] Koizumi, Y., Hara, Y., Yazaki, Y., Sakano, K. and Ishizawa, K. (2011) Involvement of plasma membrane H+-ATPase in anoxic elongation of stems in pondweed (Potamogeton distinctus) turions. New Phytologist, 190, 421-430.
[35] Huberman, M., Pressman, E. and Jaffe, M.J. (1993) Pith autolysis in plants. 4. The activity of polyga-lacturonase and cellulase during drought stress-induced pith autolysis. Plant and Cell Physiology, 34, 795-801.
[36] Pierik, R., de Wit, M. and Voesenek, L. (2011) Growth-mediated stress escape: convergence of signal transduction pathways activated upon exposure to two different environmental stresses. New Phytologist, 189, 122-134.