卧龙自然保护区华西箭竹(Fargesia nitida)生态学特征随海拔梯度的变化
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摘要
海拔梯度由于包含了温度、水分、光照等环境因子的剧烈变化而成为研究植物的环境适应性及其对全球气候变化响应的理想区域。在卧龙自然保护区沿海拔梯度研究了华西箭竹(Fargesia nitida(Mitford)T.P.Yi)分株的丛结构,比叶面积(SLA)、基径、株高、枝下高、单株地上部生物量及各器官生物量的分配。结果表明:华西箭竹丛结构表现为平均单丛面积和丛密度随着海拔升高呈先增大后降低的趋势,而单丛分株数先减少后增大;华西箭竹分株生长特征除SLA和枝下高线性降低,其他各调查变量随着海拔升高,总体呈先增大后减小的单峰变化趋势,高峰值在2 800 m附近。反映了环境因子随海拔升高的非线性变化。分析认为,高海拔地区的华西箭竹对温度较为敏感,气候变暖将有利于它的生长,因此其分布上限可能会提高。本研究结果拓展和丰富了海拔梯度上植物对异质性环境的适应和响应的规律研究,也可对地震后保护区植被的恢复与重建提供科学参考。
Global warming has been predicted to occur most rapidly at high altitude.We investigated the clump structure and growth traits of F.nitida across its altitudinal gradients from the lowest distribution boundaries(higher temperature) to the uppermost elevational limits(lower temperature) in the Wolong Nature Reserve.The present study aimed to answer the questions:(1) Do plants with or without secondary growth respond to global climate change similarly?(2) What factors mainly determine the ecological characteristics of F.nitida to elevation?(3) What is the future fate of these bamboos and the Giant Panda under global climate change in the reserve? The field investigation was carried out in April,2008.The growth are temporally stable before the growing season and thus are most suitable for altitudinal comparisons.According to nearly-equal interval of elevation,three plots at every elevation(10 m × 10 m each) were established and the gap is above 50 m among them.Number of F.nitida clumps in each plot was recorded to estimate the clump density.Three clumps in each plot were randomly selected to investigate ramet number per clump,basal diameter,height and the clear length of each ramet were measured.The clumps were then harvested,and the leaf,shoot,and stem biomass were measured separately after oven-drying to a constant weight.Fresh leaf area was scanned and analyzed using the ImageJ software and dried to a constant weight.One-way analysis of variances were used for all variables to test the differences among plots at different elevations,and multiple comparisons were conducted by Tukey's HSD test.Statistical analyses and modeling were done with SPSS 12.0 statistical software packages,and figures were done with SigmaPlot 10.0 software.The results showed that changes in the mean clump area and clump density of F.nitida populations with increasing elevation showed a convex curve,but change in the number of ramets per clump was a concave curve,both with a turning point at about 2800 m a.s.l.Except for a decreased trend of SLA and clear length,changes in other morphological traits(diameter,height,internode number) and biomass(leaf,shoot,stem and aboveground biomass) of F.nitida with increasing elevation were not linear,but formed a convex curve peaked at about 2800 m a.s.l.The high-altitude species(F.nitida) may benefit from global warming and its uppermost limit will shift upwards with further global warming.These results can also contribute to protecting the long-term survival of the Giant Panda in the reserve.
Global warming has been predicted to occur most rapidly at high altitude.We investigated the clump structure and growth traits of F.nitida across its altitudinal gradients from the lowest distribution boundaries(higher temperature) to the uppermost elevational limits(lower temperature) in the Wolong Nature Reserve.The present study aimed to answer the questions:(1) Do plants with or without secondary growth respond to global climate change similarly?(2) What factors mainly determine the ecological characteristics of F.nitida to elevation?(3) What is the future fate of these bamboos and the Giant Panda under global climate change in the reserve? The field investigation was carried out in April,2008.The growth are temporally stable before the growing season and thus are most suitable for altitudinal comparisons.According to nearly-equal interval of elevation,three plots at every elevation(10 m × 10 m each) were established and the gap is above 50 m among them.Number of F.nitida clumps in each plot was recorded to estimate the clump density.Three clumps in each plot were randomly selected to investigate ramet number per clump,basal diameter,height and the clear length of each ramet were measured.The clumps were then harvested,and the leaf,shoot,and stem biomass were measured separately after oven-drying to a constant weight.Fresh leaf area was scanned and analyzed using the ImageJ software and dried to a constant weight.One-way analysis of variances were used for all variables to test the differences among plots at different elevations,and multiple comparisons were conducted by Tukey's HSD test.Statistical analyses and modeling were done with SPSS 12.0 statistical software packages,and figures were done with SigmaPlot 10.0 software.The results showed that changes in the mean clump area and clump density of F.nitida populations with increasing elevation showed a convex curve,but change in the number of ramets per clump was a concave curve,both with a turning point at about 2800 m a.s.l.Except for a decreased trend of SLA and clear length,changes in other morphological traits(diameter,height,internode number) and biomass(leaf,shoot,stem and aboveground biomass) of F.nitida with increasing elevation were not linear,but formed a convex curve peaked at about 2800 m a.s.l.The high-altitude species(F.nitida) may benefit from global warming and its uppermost limit will shift upwards with further global warming.These results can also contribute to protecting the long-term survival of the Giant Panda in the reserve.
引文
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[6]DESPLAND E,HOULE G.Climate influences on growth and reproduction of Pinus banksiana(Pinaceae)at the limit of the species distribution in eastern North America[J].American Journal of Botany,1997,84:928-937.
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[9]JAMES J C,GRACE J,HOAD S P.Growth and photosynthesis of Pinus sylvestris at its altitudinal limit in Scotland[J].Journal of Ecology,1994,82:297-306.
[10]KRONFUSS H,HAVRANEK W M.Effects of elevation and wind on the growth of Pinus cembra L.in a subalpine afforestation[J].Phyton,1998,39:99-106.
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[13]LI M H,YANG J,KRAUCHI N.Growth responses of Picea abies and Larix decidua to elevation in subalpine areas of Tyrol,Austria[J].Canadian Journal of Forest Research,2003,33:653-662.
[14]LI M H,YANG J.Effects of microsite on growth of Pinus cembra in the subalpine zone of the Austrian Alps[J].Annuals of Forest Science,2004,61:319-325.
[15]LI M H,KRUCHI N,DOBBERTIN M.Biomass distribution of different-aged needles in young and old Pinus cembra trees at highland and lowland sites[J].Trees,2006b,20:611-618.
[16]PENUELAS J,PRIETO P,BEIER C,et al.Response of plant species richness and primary productivity in shrub lands along a north-south gradient in Europe to seven years of experimental warming and drought:reductions in primary productivity in the heat and drought year of 2003[J].Global Change Biology,2007,13:2563-2581.
[17]PREMOLI A C,RAFFAELE E,MATHIASEN P.Morphological and phenological differences in Nothofagus pumilio from contrasting elevations:evidence from a common garden[J].Austral Ecology,2007,32:515-523.
[18]SCHOETTLE A W,ROCHELLE S G.Morphologic al variation of Pinus flexilis(Pinaceae),a bird-dispersed pine,across a range of Elevations[J].Amercian Journal of Botany,2000,87,1797-1806.
[19]FARRELLY D.The Book of Bamboo:A Comprehensive Guide to This Remarkable Plant,Its Uses,and Its History[M].San Francisco:Sierra Club Books.1984:23-46.
[20]陶建平,宋利霞.亚高山暗针叶林不同林冠环境下华西箭竹的克隆可塑性[J].生态学报,2006,26(12):4019-4026.TAO Jianping,SONG Lixia.Response of clonal plasticity of Fargesia nitida to different canopy conditions of subalpine coniferous forest[J].Acta Ecologica Sinica,2006,26(12):4019-4026.
[21]周世强,黄金燕.大熊猫主食竹种的研究与进展[J].世界竹藤通讯,2005,3(1):1-6.ZHOU Shiqiang,HUANG Jinyan.Research into bamboo species as giant panda’s main diet and its progress[J].World Bamboo Rattan,2005,3(1):1-6.
[22]张万儒.中国森林土壤[M].北京:科学出版社,1986:430-440.ZHANG Wanru.Forest soil in China[M].Beijing:Science press,1986:430-440.
[23]ABRAMOFF M D,MAGALHAES P J,RAM S J.Image processing with ImageJ[J].International Biophotonies,2004,11:36-42.
[24]刘兴良,向性明.王朗林区人工缺苞箭竹亚层数量特征及其模型[J].林业科学,1994,30(5):471-477.LIU Xingliang,XIANG Xingming.The quantitative characteristics and growth pattern of arrow bamboo plantation in Wanglang forest region[J].Scientia Silvae Sinicae,1994,30(5):471-477.
[25]O'BRIEN M J,O'HARA K L,ERBILGIN N,et al.Overstory and shrub effects on natural regeneration processes in native Pinus radiata Stands[J].Forest Ecology and Management,2007,240:178-185.
[26]PARKER W C,DEY D C.Influence of overstory density on ecophysiology of red oak(Quercus rubra)and sugar maple(Acer saccharum)seedlings in central Ontario shelterwoods[J].Tree Physiology,2008,28:797-804.
[27]TAKAHASHI K,UEMURA S,SUZUKI J I,et al.Effects of understory dwarf bamboo on soil water and the growth of overstory trees in a dense secondary Betula ermanii forest,northern Japan[J].Ecological Research,2003,18:767-774.
[28]SILVERTOWN J.Plant coexistence and the niche[J].Trends in Ecology and Evolution,2004,19:605-611.
[29]鲁叶江,王开运,杨万勤,等.缺苞箭竹群落密度对土壤养分库的影响[J].应用生态学报,2005,16(6):996-1001.LU Yejiang,WANG Kaiyun,YANG Wanqin,et al.Effects of Fargesia denudata density on soil nutrient pool[J].Chinese Journal of Applied Ecology,2005,16(6):996-1001.
[30]LUO J X,ZANG R G,LI C Y.Physiological and morphological variations of Picea asperata populations originating from different altitudes in the mountains of southwestern China[J].Forest Ecology and Management,2006,221:285-290.
[31]KRNER C.The nutritional status of plants from high altitudes:A worldwide comparison[J].Oecologia,1989,81:379-391.
[32]KRNER C.The use of‘altitude’in ecological research[J].Trends in Ecology and Evolution,2007,22:569-574.
[33]WRIGHT I J,REICH P B,CORNELISSEN J H C,et al.Modulation of leaf economic traits and trait relationships by climate[J].Global Ecology and Biogeography,2005,14:411-421.
[34]李文华,罗天祥.中国冷云杉林生产力格局及其数学模型[J].生态学报,1997,17(5):511-518.LI Wenhua,LUO Tianxiang.Productivity patterns and mathematical models of spruce-fir forests in china[J].Acta Ecologica Sinica,1997,17(5):511-518.
[35]LEVESQUE E,HENRY G H R,SVOBODA J.Phenological and growth responses of Papaver radicatum along altitudinal gradients in the Canadian High Arctic[J].Global Change Biology,1997,3:125-145.
[36]LUO T X,PAN Y D,OUYANG H,et al.Leaf area index and net primary productivity along subtropical to alpine gradients in the Tibetan Plateau[J].Global Ecology and Biogeography,2004,13:345-358.
[37]RAICH J W,RUSSELL A E,VITOUSEK P M.Primary productivity and ecosystem development along an elevational gradient on Mauna Loa,Hawaii[J].Ecology,1997,78:707-721.
[38]刘兴良,刘世荣,宿以明,等.巴郎山川滇高山栎灌丛地上生物量及其对海拔梯度的响应[J].林业科学,2006,42(6):1-7.LIU Xingliang,LIU Shirong,SU Yiming,et al.Aboveground biomass of Quercus aquifolioides shrub community and its responses to altitudinal gradients in Balangshan Mountain,Sichuan Province[J].Scientia Silvae Sinicae,2006,42(6):1-7.
[39]DELUCIA E H,CALLAWAY R M,SCHLESINGER W H.Offsetting changes in biomass allocation and photosynthesis in ponderosa pine(Pinus ponderosa)in response to climate change[J].Tree Physiology,1994,14:669-677.
[40]WANG R Z,GAO Q.Morphological responses of Leymus chinensis(Poaceae)to the large-scale climatic gradient along the North-east China Transect(NECT)[J].Diversity Distribution,2004,10:65-73.Ecological responses of Fargesia nitida to altitude in the Wolong Nature Reserve
[2]KRNER C.Alpine Plant Life:Functional Plant Ecology of High Mountain Ecosystems[M].2th ed.Berlin Heidelberg,New York:Springer,2003:1-30.
[3]TRANQUILLINI W.Physiological Ecology of the Alpine Timberline:Tree Existence at High Altitude with Special Reference to the European Alps,Ecological Studies[M].New York:Springer,1979.
[4]CHAPIN F S III,BLOOM A J,FIELD C B,et al.Plant Responses to Multiple Enviromental Factors[J].Bioscience,1987,37(1):49-57.
[5]CORDELL S,GOLDSTEIN G,MULELLER-DOMBOIS D,et al.Physiological and morphological variation in Metrosideros polymorpha,a dominant Hawaiian tree species,along an altitudinal gradient:role of phonotypic plasticity[J].Oecologia,1998,113:188-196.
[6]DESPLAND E,HOULE G.Climate influences on growth and reproduction of Pinus banksiana(Pinaceae)at the limit of the species distribution in eastern North America[J].American Journal of Botany,1997,84:928-937.
[7]GRATANI L,GHIA E.Adaptive strategy at the leaf level of Arbutus unedo L.to cope with Mediterranean climate[J].Flora,2002,197:275-284.
[8]HULTINE K R,MARSHALL J D.Altitude trends in conifer leaf morphology and stable carbon isotope composition[J].Oecologia,2000,123:32-40.
[9]JAMES J C,GRACE J,HOAD S P.Growth and photosynthesis of Pinus sylvestris at its altitudinal limit in Scotland[J].Journal of Ecology,1994,82:297-306.
[10]KRONFUSS H,HAVRANEK W M.Effects of elevation and wind on the growth of Pinus cembra L.in a subalpine afforestation[J].Phyton,1998,39:99-106.
[11]LI B,SUZUKI J,HARA T.Latitudinal variation in plant size and relative growth rate in Arabidopsis thaliana[J].Oecologia,1998,115:291-301.
[12]LI C Y,ZHANG X J,LIU X L,et al.Leaf morphological and physical responses of Quercus aquifolioides along an altitudinal gradient[J].Silva Fennica,2006a,40(1):5-13.
[13]LI M H,YANG J,KRAUCHI N.Growth responses of Picea abies and Larix decidua to elevation in subalpine areas of Tyrol,Austria[J].Canadian Journal of Forest Research,2003,33:653-662.
[14]LI M H,YANG J.Effects of microsite on growth of Pinus cembra in the subalpine zone of the Austrian Alps[J].Annuals of Forest Science,2004,61:319-325.
[15]LI M H,KRUCHI N,DOBBERTIN M.Biomass distribution of different-aged needles in young and old Pinus cembra trees at highland and lowland sites[J].Trees,2006b,20:611-618.
[16]PENUELAS J,PRIETO P,BEIER C,et al.Response of plant species richness and primary productivity in shrub lands along a north-south gradient in Europe to seven years of experimental warming and drought:reductions in primary productivity in the heat and drought year of 2003[J].Global Change Biology,2007,13:2563-2581.
[17]PREMOLI A C,RAFFAELE E,MATHIASEN P.Morphological and phenological differences in Nothofagus pumilio from contrasting elevations:evidence from a common garden[J].Austral Ecology,2007,32:515-523.
[18]SCHOETTLE A W,ROCHELLE S G.Morphologic al variation of Pinus flexilis(Pinaceae),a bird-dispersed pine,across a range of Elevations[J].Amercian Journal of Botany,2000,87,1797-1806.
[19]FARRELLY D.The Book of Bamboo:A Comprehensive Guide to This Remarkable Plant,Its Uses,and Its History[M].San Francisco:Sierra Club Books.1984:23-46.
[20]陶建平,宋利霞.亚高山暗针叶林不同林冠环境下华西箭竹的克隆可塑性[J].生态学报,2006,26(12):4019-4026.TAO Jianping,SONG Lixia.Response of clonal plasticity of Fargesia nitida to different canopy conditions of subalpine coniferous forest[J].Acta Ecologica Sinica,2006,26(12):4019-4026.
[21]周世强,黄金燕.大熊猫主食竹种的研究与进展[J].世界竹藤通讯,2005,3(1):1-6.ZHOU Shiqiang,HUANG Jinyan.Research into bamboo species as giant panda’s main diet and its progress[J].World Bamboo Rattan,2005,3(1):1-6.
[22]张万儒.中国森林土壤[M].北京:科学出版社,1986:430-440.ZHANG Wanru.Forest soil in China[M].Beijing:Science press,1986:430-440.
[23]ABRAMOFF M D,MAGALHAES P J,RAM S J.Image processing with ImageJ[J].International Biophotonies,2004,11:36-42.
[24]刘兴良,向性明.王朗林区人工缺苞箭竹亚层数量特征及其模型[J].林业科学,1994,30(5):471-477.LIU Xingliang,XIANG Xingming.The quantitative characteristics and growth pattern of arrow bamboo plantation in Wanglang forest region[J].Scientia Silvae Sinicae,1994,30(5):471-477.
[25]O'BRIEN M J,O'HARA K L,ERBILGIN N,et al.Overstory and shrub effects on natural regeneration processes in native Pinus radiata Stands[J].Forest Ecology and Management,2007,240:178-185.
[26]PARKER W C,DEY D C.Influence of overstory density on ecophysiology of red oak(Quercus rubra)and sugar maple(Acer saccharum)seedlings in central Ontario shelterwoods[J].Tree Physiology,2008,28:797-804.
[27]TAKAHASHI K,UEMURA S,SUZUKI J I,et al.Effects of understory dwarf bamboo on soil water and the growth of overstory trees in a dense secondary Betula ermanii forest,northern Japan[J].Ecological Research,2003,18:767-774.
[28]SILVERTOWN J.Plant coexistence and the niche[J].Trends in Ecology and Evolution,2004,19:605-611.
[29]鲁叶江,王开运,杨万勤,等.缺苞箭竹群落密度对土壤养分库的影响[J].应用生态学报,2005,16(6):996-1001.LU Yejiang,WANG Kaiyun,YANG Wanqin,et al.Effects of Fargesia denudata density on soil nutrient pool[J].Chinese Journal of Applied Ecology,2005,16(6):996-1001.
[30]LUO J X,ZANG R G,LI C Y.Physiological and morphological variations of Picea asperata populations originating from different altitudes in the mountains of southwestern China[J].Forest Ecology and Management,2006,221:285-290.
[31]KRNER C.The nutritional status of plants from high altitudes:A worldwide comparison[J].Oecologia,1989,81:379-391.
[32]KRNER C.The use of‘altitude’in ecological research[J].Trends in Ecology and Evolution,2007,22:569-574.
[33]WRIGHT I J,REICH P B,CORNELISSEN J H C,et al.Modulation of leaf economic traits and trait relationships by climate[J].Global Ecology and Biogeography,2005,14:411-421.
[34]李文华,罗天祥.中国冷云杉林生产力格局及其数学模型[J].生态学报,1997,17(5):511-518.LI Wenhua,LUO Tianxiang.Productivity patterns and mathematical models of spruce-fir forests in china[J].Acta Ecologica Sinica,1997,17(5):511-518.
[35]LEVESQUE E,HENRY G H R,SVOBODA J.Phenological and growth responses of Papaver radicatum along altitudinal gradients in the Canadian High Arctic[J].Global Change Biology,1997,3:125-145.
[36]LUO T X,PAN Y D,OUYANG H,et al.Leaf area index and net primary productivity along subtropical to alpine gradients in the Tibetan Plateau[J].Global Ecology and Biogeography,2004,13:345-358.
[37]RAICH J W,RUSSELL A E,VITOUSEK P M.Primary productivity and ecosystem development along an elevational gradient on Mauna Loa,Hawaii[J].Ecology,1997,78:707-721.
[38]刘兴良,刘世荣,宿以明,等.巴郎山川滇高山栎灌丛地上生物量及其对海拔梯度的响应[J].林业科学,2006,42(6):1-7.LIU Xingliang,LIU Shirong,SU Yiming,et al.Aboveground biomass of Quercus aquifolioides shrub community and its responses to altitudinal gradients in Balangshan Mountain,Sichuan Province[J].Scientia Silvae Sinicae,2006,42(6):1-7.
[39]DELUCIA E H,CALLAWAY R M,SCHLESINGER W H.Offsetting changes in biomass allocation and photosynthesis in ponderosa pine(Pinus ponderosa)in response to climate change[J].Tree Physiology,1994,14:669-677.
[40]WANG R Z,GAO Q.Morphological responses of Leymus chinensis(Poaceae)to the large-scale climatic gradient along the North-east China Transect(NECT)[J].Diversity Distribution,2004,10:65-73.Ecological responses of Fargesia nitida to altitude in the Wolong Nature Reserve
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