水分异质性影响两种根茎型克隆植物赖草和假苇拂子茅的水分存储能力

doi:10.3724/SP.J.1258.2013.00044

摘要/Abstract

摘要：

水分在自然系统中呈异质性分布。有关水分异质性对克隆植物生长、形态和生理影响的研究已有大量的工作, 但是水分异质性对克隆植物存储能力, 尤其是水分存储能力影响的研究却十分缺乏。该文将两种根茎型克隆植物赖草(Leymus secalinus)和假苇拂子茅(Calamagrostis pseudophragmites)进行水分异质性和同质性实验处理, 探讨水分异质性对克隆植物水分存储能力、生长和形态的影响。在异质性水分处理下, 两种克隆植物的间隔子、枝和根的含水量均显著增加。两种克隆植物对水分异质性分布的适应策略有所不同, 赖草通过降低单个克隆分株的生长、提高芽的数量以应对水分异质性, 而假苇拂子茅通过增强整个分株种群的地下部分(根状茎、根和芽)生长来应对水分资源的异质性分布。水分储存能力的增强可以提高克隆植物适应水分异质性的能力。

关键词: 适应策略, 克隆储存, 植物含水量, 分株种群, 水分异质性

Abstract:

Aims Essential resources for plant growth are heterogeneously distributed in natural systems, and clonal plants can efficiently use heterogeneously distributed resources. Many studies have addressed effects of heterogeneous water supply on the growth, morphology and physiology of clonal plants, but few have tested effects of water heterogeneity on the ability of clonal plants to store water.
Methods We grew ramets of two rhizomatous clonal plants Leymus secalinus and Calamagrostis pseudophragmites under both homogeneous and heterogeneous water supply. The experiment was conducted from 15 August to 14 October 2010 under a rain shelter at Ordos Sandland Ecological Station (39°29′ N, 110°11′ E). Water content and biomass of roots, shoots, spacers (rhizomes) and buds were measured.
Important findings Spacer water content, shoot water content and root water content were significantly higher under the heterogeneous water condition than under the homogeneous water condition in both species, suggesting that clonal plants could increase the ability to store water when growing in heterogeneous water conditions. Leymus secalinus decreased single ramet biomass and increased bud number to adapt to water heterogeneity, while C. pseudophragmites increased belowground biomass, including rhizome biomass, root biomass and bud biomass, to deal with water heterogeneity. Increasing water storage may increase the ability of clonal plants to adapt to unpredictable water conditions.

Key words: adaptive strategy, clonal storage, plant water content, ramet population, water heterogeneity

叶学华,胡宇坤,刘志兰,高树琴,董鸣. 水分异质性影响两种根茎型克隆植物赖草和假苇拂子茅的水分存储能力. 植物生态学报, 2013, 37(5): 427-435. DOI: 10.3724/SP.J.1258.2013.00044

YE Xue-Hua,HU Yu-Kun,LIU Zhi-Lan,GAO Shu-Qin,DONG Ming. Water heterogeneity affects water storage in two rhizomatous clonal plants Leymus secalinus and Calamagrostis pseudophragmites. Chinese Journal of Plant Ecology, 2013, 37(5): 427-435. DOI: 10.3724/SP.J.1258.2013.00044

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https://www.plant-ecology.com/CN/Y2013/V37/I5/427

图/表 6

参考文献 43

[1]	Alpert P ( 1999a). Clonal integration in Fragaria chiloensis differs between populations: ramets from grassland are selfish. Oecologia, 120, 69-76. DOI URL PMID
[2]	Alpert P ( 1999b). Effects of clonal integration on plant plasticity in Fragaria chiloensis. Plant Ecology, 141, 99-106.
[3]	Chen YF, Yu FH, Zhang CY, Dong M ( 2001). Role of clonal growth of the rehizomatous grass Psammochloa villosa in patch dynamics of Mu US sandy land. Acta Ecologica Sinica, 21, 1745-1750. (in Chinese with English abstract)
	[ 陈玉福, 于飞海, 张称意, 董鸣 ( 2001). 根茎禾草沙鞭的克隆生长在毛乌素沙地斑块动态中的作用. 生态学报, 21, 1745-1750.]
[4]	Coelho FF, Capelo C, Ribeiro LC, Figueira JEC ( 2008). Reproductive modes in Leiothrix(Eriocaulaceae) in South- eastern Brazil: the role of microenvironmental heterogeneity. Annals of Botany, 101, 353-360. URL PMID
[5]	de Kroon H, Fransen B, van Rheenen JWA, van Dijk A, Kreulen R ( 1996). High levels of inter-ramet water translocation in two rhizomatous Carex species, as quantified by deuterium labelling. Oecologia, 106, 73-84. URL PMID
[6]	Dong BC, Yu GL, Guo W, Zhang MX, Dong M, Yu FH ( 2010a). How internode length, position and presence of leaves affect survival and growth of Alternanthera philoxeroides after fragmentation? Evolutionary Ecology, 24, 1447-1461.
[7]	Dong BC, Zhang MX, Alpert P, Lei GC, Yu FH ( 2010b). Effects of orientation on survival and growth of small fragments of the invasive, clonal plant Alternanthera philoxeroides. PloS One, 5, e13631. DOI URL PMID
[8]	Dong M ( 1996a). Plant clonal growth in heterogeneous habitats: risk-spreading. Acta Phytoecologica Sinica, 20, 543-548. (in Chinese with English abstract)
	[ 董鸣 ( 1996a). 异质性生境中的植物克隆生长: 风险分摊. 植物生态学报, 20, 543-548.]
[9]	Dong M ( 1996b). Clonal growth in plants in relation to resource heterogeneity: foraging behavior. Acta Botanica Sinica, 38, 828-835. (in Chinese with English abstract)
	[ 董鸣 ( 1996b). 资源异质性环境中的植物克隆生长: 觅食行为. 植物学报, 38, 828-835.]
[10]	Dong M ( 1999). Effects of severing rhizome on clonal growth in rhizomatoua grass species Psammochloa villosa and Leymus secalinus. Acta Botanica Sinica, 41, 194-198. (in Chinese with English abstract)
	[ 董鸣 ( 1999). 切断根茎对根茎禾草沙鞭和赖草克隆生长的影响. 植物学报, 41, 194-198.]
[11]	Dong M (2011). Clonal Plant Ecology. Sciences Press, Beijing. (in Chinese)
	[ 董鸣 (2011). 克隆植物生态学. 科学出版社, 北京.]
[12]	Dong M, Alaten B ( 1999). Clonal plasticity in response to rhizome severing and heterogeneous resource supply in the rhizomatous grass Psammochloa villosa in an Inner Mongolian dune, China. Plant Ecology, 141, 53-58.
[13]	Dong M, de Kroon H ( 1994). Plasticity in morphology and biomass allocation in Cynodon dactylon, a grass species forming stolons and rhizomes. Oikos, 70, 99-106.
[14]	Huang CX ( 1991). The changes of the physical environment of the southern Maowushu Sandy Land during Holocene. Geographical Research, 10(2), 52-59. (in Chinese with English abstract)
	[ 黄赐璇 ( 1991). 毛乌素沙地南缘全新世自然环境. 地理研究, 10(2), 52-59.]
[15]	Hutchings MJ, de Kroon H ( 1994). Foraging in plants: the role of morphological plasticity in resource acquisition. Advances in Ecological Research, 25, 159-238.
[16]	Jackson RB, Caldwell MM ( 1993). The scale of nutrient heterogeneity around individual plants and its quantification with geostatistics. Ecology, 74, 612-614.
[17]	Kotliar NB, Wiens JA ( 1990). Multiple scales of patchiness and patch structure: a hierarchical framework for the study of heterogeneity. Oikos, 59, 253-260.
[18]	Li R ( 1998). Clonal Growth in the Giant Bamboo Phyllostachys pubescens. PhD dissertation, Utrecht University, Utrecht.
[19]	Liu F, Chen JM, Wang QF ( 2009). Trade-offs between sexual and asexual reproduction in a monoecious species Sagittaria pygmaea(Alismataceae): the effect of different nutrient levels. Plant Systematics and Evolution, 277, 61-65.
[20]	Liu FH, Yu FH, Liu WS, Krüsi BO, Cai XH, Schneller JJ, Dong M ( 2007). Large clones on cliff faces: expanding by rhizomes through crevices. Annals of Botany, 100, 51-54. DOI URL PMID
[21]	Liu J, Zhu XW, Yu FH, Dong M, Zhang SM, Wang RQ ( 2003). Spatial heterogeneity of Ulmus pumila open forest ecosystem in Otindag Sandy Land. Environmental Science, 24, 29-34. (in Chinese with English abstract)
	[ 刘建, 朱选伟, 于飞海, 董鸣, 张淑敏, 王仁卿 ( 2003). 浑善达克沙地榆树疏林生态系统的空间异质性. 环境科学, 24, 29-34.]
[22]	Mutoh N, Yoshida KH, Yokoi Y, Kimura M, Hogetsu K ( 1968). Studies on the production processes and net production of Miscanthus sacchariflorus community. Japanese Journal of Botany, 20, 67-92.
[23]	Oborny B, Czárán T, Kun Á ( 2001). Exploration and exploitation of resource patches by clonal growth: a spatial model on the effect of transport between modules. Ecological Modelling, 141, 151-169. DOI URL
[24]	Roiloa SR, Retuerto R ( 2005). Presence of developing ramets of Fragaria vesca L. increases photochemical efficiency in parent ramets. International Journal of Plant Sciences, 166, 795-803. DOI URL
[25]	Roiloa SR, Retuerto R ( 2006a). Development, photosynthetic activity and habitat selection of the clonal plant Fragaria vesca growing in copper-polluted soil. Functional Plant Biology, 33, 961-971. DOI URL PMID
[26]	Roiloa SR, Retuerto R ( 2006b). Small-scale heterogeneity in soil quality influences photosynthetic efficiency and habitat selection in a clonal plant. Annals of Botany, 98, 1043-1052. URL PMID
[27]	Stuefer JF (1997). Division of Labour in Clonal Plants? On the Response of Stoloniferous Herbs to Environmental Heterogeneity. PhD dissertation, Utrecht University Press, Utrecht, The Netherlands.
[28]	Stuefer JF, During HJ, de Kroon H ( 1994). High benefits of clonal integration in two stoloniferous species, in response to heterogeneous light environments. Journal of Ecology, 82, 511-518. DOI URL
[29]	Suzuki JI, Hutchings MJ (1997). Interactions between shoots in clonal plants and the effects of stored resources on the structure of shoot populations. In: de Kroon H, van Groenendael J eds. The Ecology and Evolution of Clonal Plants. Backhuys Publishers, Leiden. 311-329.
[30]	Suzuki JI, Stuefer JF ( 1999). On the ecological and evolutionary significance of storage in clonal plants. Plant Species Biology, 14, 11-17. DOI URL
[31]	Thompson FL, Eckert CG ( 2004). Trade-offs between sexual and clonal reproduction in an aquatic plant: experimental manipulations vs. phenotypic correlations. Journal of Evolutionary Biology, 17, 581-592. DOI URL PMID
[32]	Watson MA ( 2008). Resource storage and the expression of clonal plant life histories. Evolutionary Ecology, 22, 471-475. DOI URL
[33]	Wijesinghe DK, Hutchings MJ ( 1997). The effects of spatial scale of environmental heterogeneity on the growth of a clonal plant: an experimental study with Glechoma hederacea. Journal of Ecology, 85, 17-28. DOI URL
[34]	Winkler E, Fischer M ( 1999). Two fitness measures for clonal plants and the importance of spatial aspects. Plant Ecology, 141, 191-199. DOI URL
[35]	Xiao KY, Yu D, Xu XW, Xiong W ( 2007). Benefits of clonal integration between interconnected ramets of Vallisneria spiralis in heterogeneous light environments. Aquatic Botany, 86, 76-82. DOI URL
[36]	Yang YF, Zhang BT (2004). Clone growth and its age structure of Leymus secalimus modules in the Songnen Plain of China. Chinese Journal of Applied Ecology, 15, 2109-2112. (in Chinese with English abstract) URL PMID
	[ 杨允菲, 张宝田 ( 2004). 松嫩平原赖草无性系生长及其构件的年龄结构. 应用生态学报, 15, 2109-2112.] URL PMID
[37]	Yang YF, Zheng HY ( 2000). Age structure of clone population of Calamagrostis pseudophragmites in Songnen Plain of China. Acta Prataculturae Sinica, 9(3), 8-13. (in Chinese with English abstract)
	[ 杨允菲, 郑慧莹 ( 2000). 松嫩平原假苇拂子茅无性系种群的年龄结构. 草业学报, 9(3), 8-13.]
[38]	Ye XH, Yu FH, Dong M ( 2006). A trade-off between guerrilla and phalanx growth forms in Leymus secalinus under different nutrient supplies. Annals of Botany, 98, 187-191. DOI URL PMID
[39]	Yu FH, Dong M, Krüsi B ( 2004). Clonal integration helps Psammochloa villosa survive sand burial in an inland dune. New Phytologist, 162, 697-704. DOI URL
[40]	Yu FH, Dong M, Zhang CY ( 2002). Intraclonal resource sharing and functional specialization of ramets in response to resource heterogeneity in three stoloniferous herbs. Acta Botanica Sinica, 44, 468-473.
[41]	Yu FH, Dong M, Zhang CY, Zhang SM ( 2002). Phenotypic plasticity in response to salinity gradient in a stoloniferous herb Halerpestes ruthenica. Acta Phytoecologica Sinica, 26, 140-148. (in Chinese with English abstract)
	[ 于飞海, 董鸣, 张称意, 张淑敏 ( 2002). 匍匐茎草本金戴戴对基质盐分含量的表型可塑性. 植物生态学报, 26, 140-148.]
[42]	Zhang XS (张新时) ( 1994). Principles and optimal models for development of Maowusu sandy grassland. Acta Phytoevologica Sinica, 18, 1-16. (in Chinese with English abstract)
	[ 张新时 ( 1994). 毛乌素沙地的生态背景及其草地建设的原则与优化模式. 植物生态学报, 18, 1-16.]
[43]	Zhu LY (1993). Site Quality Evaluation of Trees and Shrubs in Mu Us Sandy Land. China Forestry Publishing House, Beijing. (in Chinese)
	[ 朱灵益 (1993). 毛乌素沙地乔灌木立地质量评价. 中国林业出版社, 北京.]

性状 Trait	物种 Species			水分处理 Water treatment			交互作用 Interaction
性状 Trait	df	F	p	df	F	p	df	F	p
生物量 Biomass
总生物量 Total biomass	1, 19	15.93	0.001	1, 19	0.62	0.441	1, 19	1.30	0.269
间隔子生物量 Spacer biomass	1, 19	7.40	0.014	1, 19	2.88	0.106	1, 19	0.95	0.341
芽生物量 Bud biomass	1, 19	0.03	0.866	1, 19	7.16	0.015	1, 19	0.09	0.765
枝生物量 Shoot biomass	1, 19	42.03	<0.001	1, 19	0.10	0.752	1, 19	2.37	0.140
根生物量 Root biomass	1, 19	7.22	0.015	1, 19	0.36	0.557	1, 19	0.74	0.401
生物量分配 Biomass allocation
根冠比 Root to shoot ratio	1, 19	0.13	0.725	1, 19	0.62	0.443	1, 19	0.02	0.879
分株数量和大小 Number and size of ramets
分株数 Number of ramets	1, 19	6.89	0.017	1, 19	0.77	0.113	1, 19	0.54	0.471
单个分株生物量 Biomass per ramet	1, 19	8.86	0.008	1, 19	0.02	0.890	1, 19	8.02	0.011
芽数量和大小 Number and size of buds
芽数 Number of buds	1, 19	2.45	0.134	1, 19	24.60	<0.001	1, 19	0.43	0.522
单个芽生物量 Biomass per bud	1, 19	0.54	0.472	1, 19	<0.001	0.997	1, 19	0.12	0.733
含水量 Water content
总含水量 Total water content	1, 19	1.58	0.224	1, 19	94.36	<0.001	1, 19	2.81	0.110
芽含水量 Bud water content	1, 19	0.55	0.469	1, 19	3.12	0.093	1, 19	0.004	0.951
间隔子含水量 Spacer water content	1, 19	11.39	0.003	1, 19	35.38	<0.001	1, 19	3.01	0.099
根含水量 Root water content	1, 19	1.57	0.225	1, 19	53.71	<0.001	1, 19	12.98	0.002
枝含水量 Shoot water content	1, 19	0.30	0.591	1, 19	13.22	0.002	1, 19	0.44	0.514

性状 Trait	物种 Species			水分处理 Water treatment			交互作用 Interaction
性状 Trait	df	F	p	df	F	p	df	F	p
生物量 Biomass
总生物量 Total biomass	1, 19	15.93	0.001	1, 19	0.62	0.441	1, 19	1.30	0.269
间隔子生物量 Spacer biomass	1, 19	7.40	0.014	1, 19	2.88	0.106	1, 19	0.95	0.341
芽生物量 Bud biomass	1, 19	0.03	0.866	1, 19	7.16	0.015	1, 19	0.09	0.765
枝生物量 Shoot biomass	1, 19	42.03	<0.001	1, 19	0.10	0.752	1, 19	2.37	0.140
根生物量 Root biomass	1, 19	7.22	0.015	1, 19	0.36	0.557	1, 19	0.74	0.401
生物量分配 Biomass allocation
根冠比 Root to shoot ratio	1, 19	0.13	0.725	1, 19	0.62	0.443	1, 19	0.02	0.879
分株数量和大小 Number and size of ramets
分株数 Number of ramets	1, 19	6.89	0.017	1, 19	0.77	0.113	1, 19	0.54	0.471
单个分株生物量 Biomass per ramet	1, 19	8.86	0.008	1, 19	0.02	0.890	1, 19	8.02	0.011
芽数量和大小 Number and size of buds
芽数 Number of buds	1, 19	2.45	0.134	1, 19	24.60	<0.001	1, 19	0.43	0.522
单个芽生物量 Biomass per bud	1, 19	0.54	0.472	1, 19	<0.001	0.997	1, 19	0.12	0.733
含水量 Water content
总含水量 Total water content	1, 19	1.58	0.224	1, 19	94.36	<0.001	1, 19	2.81	0.110
芽含水量 Bud water content	1, 19	0.55	0.469	1, 19	3.12	0.093	1, 19	0.004	0.951
间隔子含水量 Spacer water content	1, 19	11.39	0.003	1, 19	35.38	<0.001	1, 19	3.01	0.099
根含水量 Root water content	1, 19	1.57	0.225	1, 19	53.71	<0.001	1, 19	12.98	0.002
枝含水量 Shoot water content	1, 19	0.30	0.591	1, 19	13.22	0.002	1, 19	0.44	0.514

物种 Species	性状 Trait	df	t	p
赖草 Leymus secalinus	总生物量 Total biomass	10	0.204	0.842
	间隔子生物量 Spacer biomass	10	-0.417	0.685
	芽生物量 Bud biomass	10	-1.564	0.149
	枝生物量 Shoot biomass	10	0.682	0.877
	根生物量 Root biomass	10	0.159	0.511
	根冠比 Root to shoot ratio	10	-0.581	0.574
	分株数 Number of ramets	10	-1.529	0.157
	单个分株生物量 Biomass per ramet	10	2.565	0.028
	芽数 Number of buds	10	-2.795	0.019
	单个芽生物量 Biomass per bud	10	0.206	0.841
	总含水量 Total water content	10	-5.385	<0.001
	芽含水量 Bud water content	10	-1.491	0.167
	间隔子含水量 Spacer water content	10	-4.445	0.001
	根含水量 Root water content	10	-3.647	0.004
	枝含水量 Shoot water content	10	-3.307	0.008
假苇拂子茅 Calamagrostis pseudophragmites	总生物量 Total biomass	10	-2.171	0.055
	间隔子生物量 Spacer biomass	10	-3.313	0.008
	芽生物量 Bud biomass	10	-2.696	0.022
	枝生物量 Shoot biomass	10	-2.664	0.209
	根生物量 Root biomass	10	-1.342	0.024
	根冠比 Root to shoot ratio	10	-0.360	0.726
	分株数 Number of ramets	10	-0.412	0.689
	单个分株生物量 Biomass per ramet	5.317	-1.833	0.123
	芽数 Number of buds	10	-4.982	0.001
	单个芽生物量 Biomass per bud	6.323	-0.085	0.935
	总含水量 Total water content	10	-5.156	0.000
	芽含水量 Bud water content	10	-1.439	0.181
	间隔子含水量 Spacer water content	10	-4.733	0.001
	根含水量 Root water content	4.528	-5.647	0.003
	枝含水量 Shoot water content	10	-2.296	0.045

物种 Species	性状 Trait	df	t	p
赖草 Leymus secalinus	总生物量 Total biomass	10	0.204	0.842
	间隔子生物量 Spacer biomass	10	-0.417	0.685
	芽生物量 Bud biomass	10	-1.564	0.149
	枝生物量 Shoot biomass	10	0.682	0.877
	根生物量 Root biomass	10	0.159	0.511
	根冠比 Root to shoot ratio	10	-0.581	0.574
	分株数 Number of ramets	10	-1.529	0.157
	单个分株生物量 Biomass per ramet	10	2.565	0.028
	芽数 Number of buds	10	-2.795	0.019
	单个芽生物量 Biomass per bud	10	0.206	0.841
	总含水量 Total water content	10	-5.385	<0.001
	芽含水量 Bud water content	10	-1.491	0.167
	间隔子含水量 Spacer water content	10	-4.445	0.001
	根含水量 Root water content	10	-3.647	0.004
	枝含水量 Shoot water content	10	-3.307	0.008
假苇拂子茅 Calamagrostis pseudophragmites	总生物量 Total biomass	10	-2.171	0.055
	间隔子生物量 Spacer biomass	10	-3.313	0.008
	芽生物量 Bud biomass	10	-2.696	0.022
	枝生物量 Shoot biomass	10	-2.664	0.209
	根生物量 Root biomass	10	-1.342	0.024
	根冠比 Root to shoot ratio	10	-0.360	0.726
	分株数 Number of ramets	10	-0.412	0.689
	单个分株生物量 Biomass per ramet	5.317	-1.833	0.123
	芽数 Number of buds	10	-4.982	0.001
	单个芽生物量 Biomass per bud	6.323	-0.085	0.935
	总含水量 Total water content	10	-5.156	0.000
	芽含水量 Bud water content	10	-1.439	0.181
	间隔子含水量 Spacer water content	10	-4.733	0.001
	根含水量 Root water content	4.528	-5.647	0.003
	枝含水量 Shoot water content	10	-2.296	0.045