不同海拔典型竹种枝叶大小异速生长关系
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摘要
为了分析竹子枝-叶大小间的权衡关系,本研究对武夷山不同海拔典型竹种(毛竹、箬竹、肿节少穗竹、毛竿玉山竹和武夷山玉山竹)小枝的叶片总质量、茎质量、单叶质量和出叶强度等性状进行测定.结果表明:随海拔升高,5个竹种间小枝上总叶质量与茎质量的异速生长指数呈显著下降趋势.竹种内,毛竹、箬竹和肿节少穗竹总叶质量与茎质量在不同海拔上均拥有共同异速生长指数(分别为0.94、0.85、0.84).毛竿玉山竹和武夷山玉山竹的叶茎质量也存在共同异速生长指数(0.79).除武夷山玉山竹外,竹子单叶质量与出叶强度之间均呈显著的负相关关系.5个竹种的单叶质量和出叶强度之间存在共同异速生长指数-1.12.总之,竹类植物的小枝总体上倾向于在低海拔环境中着生更多的叶片,而在高海拔生境下则投资更多的生物量到茎的构造上.尽管竹种间小枝的茎投资随海拔升高而增加,但其基于茎质量的出叶强度策略取决于叶片大小的构建而不是海拔生境差异.
To investigate the trade-off between the twig size and leaf size,we measured the total leaf mass,stem mass,individual leaf mass and leafing intensity of typical bamboo's( Phyllostachys edulis,Indocalamus tessellatus,Oligostachyum oedogonatume,Yushania hirticaulis and Yushania wuyishanensis) twigs at different altitudes in Wuyi Mountain. The results showed that the exponents of the scaling between total leaf mass and stem mass on twigs significantly decreased with increasing altitude for all the five bamboo species. The common scaling exponents of total leaf mass vs. stem mass for P. edulis,I. tessellatus and O. oedogonatum were 0.94,0. 85 and 0. 84,respectively. A common slope of 0.79 was observed in total leaf mass vs. stem mass in Y. hirticaulis and Y. wuyishanensis. There was significant negative correlation between individual leaf mass and leafing intensity among bamboo species,except Y. wuyishanensis. A common slope of-1. 12 existed between individual leaf mass and leafing intensity for five bamboo species. In conclusion,bamboos at low altitudes tended to support more leaf biomass while preferring to invest more to stem biomass at high altitudes. Although the stem mass investment of different bamboo's twig increased with altitude,the leafing intensity strategies based on stem mass were determined by the leaf size construction rather than altitude.
To investigate the trade-off between the twig size and leaf size,we measured the total leaf mass,stem mass,individual leaf mass and leafing intensity of typical bamboo's( Phyllostachys edulis,Indocalamus tessellatus,Oligostachyum oedogonatume,Yushania hirticaulis and Yushania wuyishanensis) twigs at different altitudes in Wuyi Mountain. The results showed that the exponents of the scaling between total leaf mass and stem mass on twigs significantly decreased with increasing altitude for all the five bamboo species. The common scaling exponents of total leaf mass vs. stem mass for P. edulis,I. tessellatus and O. oedogonatum were 0.94,0. 85 and 0. 84,respectively. A common slope of 0.79 was observed in total leaf mass vs. stem mass in Y. hirticaulis and Y. wuyishanensis. There was significant negative correlation between individual leaf mass and leafing intensity among bamboo species,except Y. wuyishanensis. A common slope of-1. 12 existed between individual leaf mass and leafing intensity for five bamboo species. In conclusion,bamboos at low altitudes tended to support more leaf biomass while preferring to invest more to stem biomass at high altitudes. Although the stem mass investment of different bamboo's twig increased with altitude,the leafing intensity strategies based on stem mass were determined by the leaf size construction rather than altitude.
引文
[1]Westoby M,Wright IJ.The leaf size-twig size spectrum and its relationship to other important spectra of variation among species.Oecologia,2003,135:621-628
[2]Niklas KJ,Enquist BJ.On the vegetative biomass partitioning of seed plant leaves,stems,and roots.American Naturalist,2002,159:482-497
[3]West GB,Brown JH,Enquist BJ.The fourth dimension of life:Fractal geometry and allometri scaling of organisms.Science,1999,284:1677-1679
[4]Brouat C,Mckey D.Leaf-stem allometry,hollow stems,and the evolution of caulinary domatia in myrmecophytes.New Phytologist,2010,151:391-406
[5]Westoby M,Falster DS,Moles AT,et al.Plant ecological strategies:Some leading dimensions of variation between species.Annual Review of Ecology and Systematics,2002,33:125-159
[6]Normand F,Bissery C,Damour G,et al.Hydraulic and mechanical stem properties affect leaf-stem allometry in mango cultivars.New Phytologist,2010,178:590-602
[7]Sun S,Jin D,Shi P.The leaf size-twig size spectrum of temperate woody species along an altitudinal gradient:An invariant allometric scaling relationship.Annals of Botany,2006,97:97-107
[8]Xiang S,Wu N,Sun SC.Within-twig biomass allocation in subtropical evergreen broad-leaved species along an altitudinal gradient:Allometric scaling analysis.Trees,2008,23:637-647
[9]Li G,Yang D,Sun S.Allometric relationships between lamina area,lamina mass and petiole mass of 93 temperate woody species vary with leaf habit,leaf form and altitude.Functional Ecology,2008,22:557-564
[10]Li M(李曼),Zheng Y(郑媛),Guo Y-R(郭英荣),et al.Scaling relationships between twig size and leaf of Pinus hwangshanensis along an altitudinal gradient.Chinese Journal of Applied Ecology(应用生态学报),2017,28(2):537-544(in Chinese)
[11]Kleiman D,Aarssen LW.The leaf size/number trade-off in trees.Journal of Ecology,2007,95:376-382
[12]Ogawa K.The leaf mass/number trade-off of Kleiman and Aarssen implies constancy of leaf biomass,its density and carbon uptake in forest stands:Scaling up from shoot to stand level.Journal of Ecology,2008,96:188-191
[13]Milla R.The leafing intensity premium hypothesis tested across clades,growth forms and altitudes.Journal of Ecology,2009,97:972-983
[14]Yan ER,Wang XH,Scoot XC,et al.Scaling relationships among twig size,leaf size and leafing intensity in a successional series of subtropical forests.Tree Physiology,2013,33:609-617
[15]Chen H,Niklas KJ,Yang D,et al.The effect of twig architecture and seed number on seed size variation in subtropical woody species.New Phytologist,2009,183:1212-1221
[16]Dombroskie SL,Tracey AJ,Aarssen LW.Leafing intensity and the fruit size/number trade-off in woody angiosperms.Journal of Ecology,2016,104:1759-1767
[17]Lzaro A,Larrinaga AR.A multi-level test of the seed number/size trade-off in two Scandinavian communities.PLo S One,2018,13(7):e0201175
[18]Niklas KJ,Cobb ED,Niinemets,et al.“Diminishing returns”in the scaling of functional leaf traits across and within species groups.Proceedings of the National Academy of Sciences of the United States of America,2007,104:8891-8896
[19]Sun J,Fan R,Niklas KJ,et al.“Diminishing returns”in the scaling of leaf area vs.dry mass in Wuyi Mountain bamboos,Southeast China.American Journal of Botany,2017,104:993-998
[20]Pan S,Liu C,Zhang W,et al.The scaling relationships between leaf mass and leaf area of vascular plant species change with altitude.PLo S One,2013,8(10):e76872
[21]Niinemets,Portsmuth A,Tobias M.Leaf size modifies support biomass distribution among stems,petioles and mid-ribs in temperate plants.New Phytologist,2010,171:91-104
[22]Sack L,Frole K.Leaf structural diversity is related to hydraulic capacity in tropical rain forest trees.Ecology,2006,87:483-491
[23]Sack L,Scoffoni C.Leaf venation:Structure,function,development,evolution,ecology and applications in the past,present and future.New Phytologist,2013,198:983-1000
[24]Chen S-L(陈双林).Study on Effects of Altitude on the Structure and Physio-ecological Characteristic of Moso Bamboo Forest.PhD Thesis.Nanjing:Nanjing Forestry University,2009(in Chinese)
[25]Song L-X(宋利霞),Tao J-P(陶建平),Wang W(王微),et al.The ramet population structures of the clonal bamboo Fargesia nitida in different canopy conditions of sulbalpine dark coniferous forest in Wolong Nature Reserve,China.Acta Ecologica Sinica(生态学报),2006,26(3):730-736(in Chinese)
[26]Montti L,Villagra M,Campanello PI,et al.Functional traits enhance invasiveness of bamboos over co-occurring tree saplings in the semideciduous Atlantic forest.Acta Oecologica,2014,54:36-44
[27]Gagnon PR,Platt WJ,Moser EB.Response of a native bamboo[Arundinaria gigantea,(Walt.)Muhl.]in a wind-disturbed forest.Forest Ecology and Management,2007,241:288-294
[28]Shi J-M(施建敏),Ye X-H(叶学华),Chen F-S(陈伏生),et al.Adaptation of bamboo to heterogeneous habitat:Phenotypic plasticity.Acta Ecologica Sinica(生态学报),2014,34(20):5687-5695(in Chinese)
[29]Yang SJ,Sun M,Zhang YJ,et al.Strong leaf morphological,anatomical,and physiological responses of a subtropical woody bamboo(Sinarundinaria nitida)to contrasting light environments.Plant Ecology,2014,215:97-109
[30]Zhu Q-G(朱强根),Jin A-W(金爱武),Wang Y-K(王意锟),et al.Biomass allocation of branches and leaves in Phyllostachys heterocycla‘Pubescens’under different management modes:Allometric scaling analysis.Chinese Journal of Plant Ecology(植物生态学报),2013,37(9):811-819(in Chinese)
[31]Tong L(童龙),Li B(李彬),Chen L-J(陈丽洁),et al.Effects of truncation intensity on the biomass allocation and physiological characteristics of Dendrocalamus latiflorus.Chinese Journal of Applied Ecology(应用生态学报),2016,27(10):3137-3144(in Chinese)
[32]Liu J(刘骏),Yang Q-P(杨清培),Song Q-N(宋庆妮),et al.Strategy of fine root expansion of Phyllostachys pubescens population into evergreen broadleaved forest.Chinese Journal of Plant Ecology(植物生态学报),2013,37(3):230-238(in Chinese)
[33]Song QN,Ouyang M,Yang QP,et al.Degradation of litter quality and decline of soil nitrogen mineralization after moso bamboo(Phyllostachys pubscens)expansion to neighboring broadleaved forest in subtropical China.Plant and Soil,2016,404:113-124
[34]Takahashi K,Suzuki UI,Hara T.Effects of understory dwarf bamboo on soil water and the growth of overstory trees in a dense secondary Betula ermanii forest,Northern Japan.Ecological Research,2003,18:767-774
[35]Gaku K,Yukihiro A,Buho H,et al.Invasion of dwarf bamboo into alpine snow-meadows in Northern Japan:Pattern of expansion and impact on species diversity.Ecology&Evolution,2011,1:85-96
[36]Yang Q-P(杨清培),Guo Y-R(郭英荣),Lan W-J(兰文军),et al.Addition effects of co-expansion of two bamboos on plant diversity in broad-leaved forests.Chinese Journal of Applied Ecology(应用生态学报),2017,28(10):3155-3162(in Chinese)
[37]Liu G,Shi P,Xu Q,et al.Does the size-density relationship developed for bamboo species conform to the self-thinning rule?Forest Ecology and Management,2016,361:339-345
[38]Yang D,Li G,Sun S.The generality of leaf size versus number trade-off in temperate woody species.Annals of Botany,2008,102:623-629
[39]Warton DI,Wright IJ,Falster DS,et al.A review of bivariate line-fitting methods for allometry.Biological Reviews,2006,81:259-291
[40]Warton DI,Duursma RA,Falster D,et al.Smatr 3:An R package for estimation and inference about allometric lines.Methods in Ecology and Evolution,2012,3:257-259
[41]Warton DI,Weber NC.Common slope tests for bivariate structural relationships.Biometrical Journal,2002,44:161-174
[42]Corner EJH.The durian theory or the origin of the modern tree.Annals of Botany,1949,13:367-414
[43]Niklas KJ.Gravity-induced effects on material properties and size of leaves on horizontal shoots of Acer saccharum(Aceraceae).American Journal of Botany,1992,79:820-827
[44]Dong L-N(董丽娜).Studies on Developmental Anatomy of Elongated Growth about Bamboo Culms.Master Thesis.Nanjing:Nanjing Forestry University,2007(in Chinese)
[45]Díaz S,Cabido M,Casanoves F.Plant functional traits and environmental filters at a regional scale.Journal of Vegetation Science,1998,9:113-122
[46]Díaz S,Cabido M,Zak M,et al.Plant functional traits,ecosystem structure and land-use history along a climatic gradient in central-western Argentina.Journal of Vegetation Science,1999,10:651-660
[47]Yue CL,Wang KH,Zhu YM.Morphological plasticity of clonal plant Phyllostachys praecox f.prevernalis(Poaceae)in response to nitrogen availability.Annales Botanici Fennici,2005,42:123-127
[48]Yue CL,Chang J,Wang KH,et al.Response of clonal growth in Phyllostachys praecox f.prevernalis to changing light intensity.Australian Journal of Botany,2004,52:171-174
[49]Fajardo A.Are trait-scaling relationships invariant across contrasting elevations in the widely distributed treeline species Nothofagus pumilio?American Journal of Botany,2016,103:821-829
[50]Huang Y,Lechowicz MJ,Price CA,et al.The underlying basis for the trade-off between leaf size and leafing intensity.Functional Ecology,2015,30:1-7
[51]Whitman T,Aarssen LW.The leaf size/number trade-off in herbaceous angiosperms.Journal of Plant Ecology,2010,3:49-58
[52]Xie X,Hu Y,Pan X,et al.Biomass allocation of stoloniferous and rhizomatous plant in response to resource availability:A phylogenetic meta-analysis.Frontiers in Plant Science,2016,7:1-11
[2]Niklas KJ,Enquist BJ.On the vegetative biomass partitioning of seed plant leaves,stems,and roots.American Naturalist,2002,159:482-497
[3]West GB,Brown JH,Enquist BJ.The fourth dimension of life:Fractal geometry and allometri scaling of organisms.Science,1999,284:1677-1679
[4]Brouat C,Mckey D.Leaf-stem allometry,hollow stems,and the evolution of caulinary domatia in myrmecophytes.New Phytologist,2010,151:391-406
[5]Westoby M,Falster DS,Moles AT,et al.Plant ecological strategies:Some leading dimensions of variation between species.Annual Review of Ecology and Systematics,2002,33:125-159
[6]Normand F,Bissery C,Damour G,et al.Hydraulic and mechanical stem properties affect leaf-stem allometry in mango cultivars.New Phytologist,2010,178:590-602
[7]Sun S,Jin D,Shi P.The leaf size-twig size spectrum of temperate woody species along an altitudinal gradient:An invariant allometric scaling relationship.Annals of Botany,2006,97:97-107
[8]Xiang S,Wu N,Sun SC.Within-twig biomass allocation in subtropical evergreen broad-leaved species along an altitudinal gradient:Allometric scaling analysis.Trees,2008,23:637-647
[9]Li G,Yang D,Sun S.Allometric relationships between lamina area,lamina mass and petiole mass of 93 temperate woody species vary with leaf habit,leaf form and altitude.Functional Ecology,2008,22:557-564
[10]Li M(李曼),Zheng Y(郑媛),Guo Y-R(郭英荣),et al.Scaling relationships between twig size and leaf of Pinus hwangshanensis along an altitudinal gradient.Chinese Journal of Applied Ecology(应用生态学报),2017,28(2):537-544(in Chinese)
[11]Kleiman D,Aarssen LW.The leaf size/number trade-off in trees.Journal of Ecology,2007,95:376-382
[12]Ogawa K.The leaf mass/number trade-off of Kleiman and Aarssen implies constancy of leaf biomass,its density and carbon uptake in forest stands:Scaling up from shoot to stand level.Journal of Ecology,2008,96:188-191
[13]Milla R.The leafing intensity premium hypothesis tested across clades,growth forms and altitudes.Journal of Ecology,2009,97:972-983
[14]Yan ER,Wang XH,Scoot XC,et al.Scaling relationships among twig size,leaf size and leafing intensity in a successional series of subtropical forests.Tree Physiology,2013,33:609-617
[15]Chen H,Niklas KJ,Yang D,et al.The effect of twig architecture and seed number on seed size variation in subtropical woody species.New Phytologist,2009,183:1212-1221
[16]Dombroskie SL,Tracey AJ,Aarssen LW.Leafing intensity and the fruit size/number trade-off in woody angiosperms.Journal of Ecology,2016,104:1759-1767
[17]Lzaro A,Larrinaga AR.A multi-level test of the seed number/size trade-off in two Scandinavian communities.PLo S One,2018,13(7):e0201175
[18]Niklas KJ,Cobb ED,Niinemets,et al.“Diminishing returns”in the scaling of functional leaf traits across and within species groups.Proceedings of the National Academy of Sciences of the United States of America,2007,104:8891-8896
[19]Sun J,Fan R,Niklas KJ,et al.“Diminishing returns”in the scaling of leaf area vs.dry mass in Wuyi Mountain bamboos,Southeast China.American Journal of Botany,2017,104:993-998
[20]Pan S,Liu C,Zhang W,et al.The scaling relationships between leaf mass and leaf area of vascular plant species change with altitude.PLo S One,2013,8(10):e76872
[21]Niinemets,Portsmuth A,Tobias M.Leaf size modifies support biomass distribution among stems,petioles and mid-ribs in temperate plants.New Phytologist,2010,171:91-104
[22]Sack L,Frole K.Leaf structural diversity is related to hydraulic capacity in tropical rain forest trees.Ecology,2006,87:483-491
[23]Sack L,Scoffoni C.Leaf venation:Structure,function,development,evolution,ecology and applications in the past,present and future.New Phytologist,2013,198:983-1000
[24]Chen S-L(陈双林).Study on Effects of Altitude on the Structure and Physio-ecological Characteristic of Moso Bamboo Forest.PhD Thesis.Nanjing:Nanjing Forestry University,2009(in Chinese)
[25]Song L-X(宋利霞),Tao J-P(陶建平),Wang W(王微),et al.The ramet population structures of the clonal bamboo Fargesia nitida in different canopy conditions of sulbalpine dark coniferous forest in Wolong Nature Reserve,China.Acta Ecologica Sinica(生态学报),2006,26(3):730-736(in Chinese)
[26]Montti L,Villagra M,Campanello PI,et al.Functional traits enhance invasiveness of bamboos over co-occurring tree saplings in the semideciduous Atlantic forest.Acta Oecologica,2014,54:36-44
[27]Gagnon PR,Platt WJ,Moser EB.Response of a native bamboo[Arundinaria gigantea,(Walt.)Muhl.]in a wind-disturbed forest.Forest Ecology and Management,2007,241:288-294
[28]Shi J-M(施建敏),Ye X-H(叶学华),Chen F-S(陈伏生),et al.Adaptation of bamboo to heterogeneous habitat:Phenotypic plasticity.Acta Ecologica Sinica(生态学报),2014,34(20):5687-5695(in Chinese)
[29]Yang SJ,Sun M,Zhang YJ,et al.Strong leaf morphological,anatomical,and physiological responses of a subtropical woody bamboo(Sinarundinaria nitida)to contrasting light environments.Plant Ecology,2014,215:97-109
[30]Zhu Q-G(朱强根),Jin A-W(金爱武),Wang Y-K(王意锟),et al.Biomass allocation of branches and leaves in Phyllostachys heterocycla‘Pubescens’under different management modes:Allometric scaling analysis.Chinese Journal of Plant Ecology(植物生态学报),2013,37(9):811-819(in Chinese)
[31]Tong L(童龙),Li B(李彬),Chen L-J(陈丽洁),et al.Effects of truncation intensity on the biomass allocation and physiological characteristics of Dendrocalamus latiflorus.Chinese Journal of Applied Ecology(应用生态学报),2016,27(10):3137-3144(in Chinese)
[32]Liu J(刘骏),Yang Q-P(杨清培),Song Q-N(宋庆妮),et al.Strategy of fine root expansion of Phyllostachys pubescens population into evergreen broadleaved forest.Chinese Journal of Plant Ecology(植物生态学报),2013,37(3):230-238(in Chinese)
[33]Song QN,Ouyang M,Yang QP,et al.Degradation of litter quality and decline of soil nitrogen mineralization after moso bamboo(Phyllostachys pubscens)expansion to neighboring broadleaved forest in subtropical China.Plant and Soil,2016,404:113-124
[34]Takahashi K,Suzuki UI,Hara T.Effects of understory dwarf bamboo on soil water and the growth of overstory trees in a dense secondary Betula ermanii forest,Northern Japan.Ecological Research,2003,18:767-774
[35]Gaku K,Yukihiro A,Buho H,et al.Invasion of dwarf bamboo into alpine snow-meadows in Northern Japan:Pattern of expansion and impact on species diversity.Ecology&Evolution,2011,1:85-96
[36]Yang Q-P(杨清培),Guo Y-R(郭英荣),Lan W-J(兰文军),et al.Addition effects of co-expansion of two bamboos on plant diversity in broad-leaved forests.Chinese Journal of Applied Ecology(应用生态学报),2017,28(10):3155-3162(in Chinese)
[37]Liu G,Shi P,Xu Q,et al.Does the size-density relationship developed for bamboo species conform to the self-thinning rule?Forest Ecology and Management,2016,361:339-345
[38]Yang D,Li G,Sun S.The generality of leaf size versus number trade-off in temperate woody species.Annals of Botany,2008,102:623-629
[39]Warton DI,Wright IJ,Falster DS,et al.A review of bivariate line-fitting methods for allometry.Biological Reviews,2006,81:259-291
[40]Warton DI,Duursma RA,Falster D,et al.Smatr 3:An R package for estimation and inference about allometric lines.Methods in Ecology and Evolution,2012,3:257-259
[41]Warton DI,Weber NC.Common slope tests for bivariate structural relationships.Biometrical Journal,2002,44:161-174
[42]Corner EJH.The durian theory or the origin of the modern tree.Annals of Botany,1949,13:367-414
[43]Niklas KJ.Gravity-induced effects on material properties and size of leaves on horizontal shoots of Acer saccharum(Aceraceae).American Journal of Botany,1992,79:820-827
[44]Dong L-N(董丽娜).Studies on Developmental Anatomy of Elongated Growth about Bamboo Culms.Master Thesis.Nanjing:Nanjing Forestry University,2007(in Chinese)
[45]Díaz S,Cabido M,Casanoves F.Plant functional traits and environmental filters at a regional scale.Journal of Vegetation Science,1998,9:113-122
[46]Díaz S,Cabido M,Zak M,et al.Plant functional traits,ecosystem structure and land-use history along a climatic gradient in central-western Argentina.Journal of Vegetation Science,1999,10:651-660
[47]Yue CL,Wang KH,Zhu YM.Morphological plasticity of clonal plant Phyllostachys praecox f.prevernalis(Poaceae)in response to nitrogen availability.Annales Botanici Fennici,2005,42:123-127
[48]Yue CL,Chang J,Wang KH,et al.Response of clonal growth in Phyllostachys praecox f.prevernalis to changing light intensity.Australian Journal of Botany,2004,52:171-174
[49]Fajardo A.Are trait-scaling relationships invariant across contrasting elevations in the widely distributed treeline species Nothofagus pumilio?American Journal of Botany,2016,103:821-829
[50]Huang Y,Lechowicz MJ,Price CA,et al.The underlying basis for the trade-off between leaf size and leafing intensity.Functional Ecology,2015,30:1-7
[51]Whitman T,Aarssen LW.The leaf size/number trade-off in herbaceous angiosperms.Journal of Plant Ecology,2010,3:49-58
[52]Xie X,Hu Y,Pan X,et al.Biomass allocation of stoloniferous and rhizomatous plant in response to resource availability:A phylogenetic meta-analysis.Frontiers in Plant Science,2016,7:1-11