常绿和落叶阔叶物种当年生小枝茎长度和茎纤细率对展叶效率的影响
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摘要
枝和叶是组成树冠的两个最主要的结构单元,枝条的结构特征及其与叶性状之间的变化关系决定了植物对光照、空间等资源的利用和适应策略。该研究以小枝茎长度和茎纤细率表征茎结构,以叶面积比(即单位小枝茎干质量的总叶面积)、叶密度(即单位茎长度的叶数量)和叶茎生物量比(即单位小枝茎干质量的总叶干质量)表征展叶效率,以浙江省清凉峰自然保护区25个常绿阔叶物种和60个落叶阔叶物种为对象,探讨小枝茎结构变化对展叶效率的影响。结果显示,展叶效率无论是用叶面积比、叶密度还是用叶茎生物量比表示,常绿和落叶物种小枝的茎长结构性状均与展叶效率呈显著的负相关关系,即随着小枝茎长度和茎纤细率的增加,展叶效率逐渐降低,可能反映了机械安全和光照限制的作用。同时,尽管茎长度-展叶效率的斜率在常绿和落叶物种间无显著差异,并且常绿物种叶密度与小枝茎结构性状之间的关系截距显著大于落叶物种,即常绿物种具有较高的出叶强度。但由于单叶面积和比叶面积显著小于落叶物种,在某一给定的小枝茎长结构下,常绿物种每单位质量的茎生物量投资获得的总叶面积和总叶质量都比落叶物种少,即常绿物种的展叶效率比落叶物种低。这可能反映了常绿物种高消耗慢收益的保守型策略。这些结果表明,小枝茎长结构对展叶效率具有显著的影响,并将随叶习性的不同而改变,与植物对环境的适应策略密切相关。
Aims Branches and leaves are the two main structural units of tree crown composition. Among the adaptive strategies of plants, the functional traits of branches and the relationships between branch traits and leaf traits determine the capacity of trees to access light and space. In this study, our objective is to test the hypothesis that leaf display efficiency is affected by the stem length to stem slender ratio within current-year twigs. Methods The stem length to stem slender ratios of current-year twigs were used as the proxy of stem structure traits. Leaf area ratio(total leaf area per stem mass), leaf density(leaf number per stem length) and leaf/stem mass ratio(total leaf mass per stem mass) were used as the proxies of leaf display efficiency. The relationship between stem structure traits and leaf display efficiency within current-year twigs were studied for 25 evergreen and 60 deciduous broadleaved woody species in Qingliang Mountain, Zhejiang, China. The standardized major axis estimation method was used to examine the scaling relationship between stem structural traits and leaf display efficiency within current-year twigs. Important findings The proxies of leaf display efficiency, measured by leaf area ratio, leaf density or leaf/stem mass ratio, were all significantly and negative correlated with stem length to stem slender ratio within current-year twigs in both evergreen and deciduous broadleaved woody species. This suggested that leaf display efficiency decreased with stem length to stem slender ratios within current-year twigs, which may reflect the role ofmechanical safety and light within twigs. The slope of the relationship between leaf display efficiency and stem long-dimension structure traits in evergreen species was not significantly different from the one in deciduous species. In contrast, the y-intercept of the relationship between leaf density and stem long-dimension structure traits was significantly larger in evergreen species than in deciduous species, i.e. the leafing intensity of evergreen species was higher than that of deciduous species. Individual leaf area and specific leaf area were smaller in evergreen species than in deciduous species, which resulted in deciduous species have a larger leaf area per stem mass and leaf mass per stem mass at a given stem length to stem slender ratio compared to evergreen species. It may reflect the conservative adaptive strategy of high consumption and slow benefit in evergreen species. Our results demonstrated that leaf display efficiency could be affected by stem length, and would change with leaf life-span(deciduous versus evergreen).
Aims Branches and leaves are the two main structural units of tree crown composition. Among the adaptive strategies of plants, the functional traits of branches and the relationships between branch traits and leaf traits determine the capacity of trees to access light and space. In this study, our objective is to test the hypothesis that leaf display efficiency is affected by the stem length to stem slender ratio within current-year twigs. Methods The stem length to stem slender ratios of current-year twigs were used as the proxy of stem structure traits. Leaf area ratio(total leaf area per stem mass), leaf density(leaf number per stem length) and leaf/stem mass ratio(total leaf mass per stem mass) were used as the proxies of leaf display efficiency. The relationship between stem structure traits and leaf display efficiency within current-year twigs were studied for 25 evergreen and 60 deciduous broadleaved woody species in Qingliang Mountain, Zhejiang, China. The standardized major axis estimation method was used to examine the scaling relationship between stem structural traits and leaf display efficiency within current-year twigs. Important findings The proxies of leaf display efficiency, measured by leaf area ratio, leaf density or leaf/stem mass ratio, were all significantly and negative correlated with stem length to stem slender ratio within current-year twigs in both evergreen and deciduous broadleaved woody species. This suggested that leaf display efficiency decreased with stem length to stem slender ratios within current-year twigs, which may reflect the role ofmechanical safety and light within twigs. The slope of the relationship between leaf display efficiency and stem long-dimension structure traits in evergreen species was not significantly different from the one in deciduous species. In contrast, the y-intercept of the relationship between leaf density and stem long-dimension structure traits was significantly larger in evergreen species than in deciduous species, i.e. the leafing intensity of evergreen species was higher than that of deciduous species. Individual leaf area and specific leaf area were smaller in evergreen species than in deciduous species, which resulted in deciduous species have a larger leaf area per stem mass and leaf mass per stem mass at a given stem length to stem slender ratio compared to evergreen species. It may reflect the conservative adaptive strategy of high consumption and slow benefit in evergreen species. Our results demonstrated that leaf display efficiency could be affected by stem length, and would change with leaf life-span(deciduous versus evergreen).
引文
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Coley PD(1988).Effects of plant growth rate and leaf lifetime on the amount and type of anti-herbivore defense.Oecologia,74,531-536.
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Day ME,Greenwood MS,Diazsala C(2002).Age-and sizerelated trends in woody plant shoot development:Regulatory pathways and evidence for genetic control.Tree Physiology,22,507-513.
Dean TJ,Long JN(1986).Validity of constant-stress and elastic-instability principles of stem formation in Pinus contorta and Trifolium pratense.Annals of Botany,58,833-840.
Diemer M(1998).Life span and dynamics of leaves of herbaceous perennials in high-elevation environments:“News from the elephant’s leg”.Functional Ecology,12,413-425.
Dong L,Jia GX,Su XH(2003).Change of the leaf tissue structure of evergreen broad-leaf plants during overwintering.Acta Horticulture Sinica,30,59-64.(in Chinese with English abstract)[董丽,贾桂霞,苏雪痕(2003).常绿阔叶植物越冬期间叶片组织结构的适应性变化.园艺学报,30,59-64.]
Falster DS,Warton DI,Wright IJ(2006).User’s guide to SMATR:Standardised major axis tests and routines version 2.0.http://www.bio.mq.edu.au/ecology/SMATR.Cited:2015-3-11.
Gartner BL(1995).Plant Stems:Physiology and Functional Morphology.Academic Press,San Diego,USA.
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Gregory RA(1980).Annual cycle of shoot development in sugar maple.Canadian Journal of Forest Research,10,316-326.
Harvey PH,Pagel MD(1991).The Comparative Method in Evolutionary Biology.Oxford University Press,Oxford,UK.239.
Honda H,Fisher JB(1978).Tree branch angle:Maximizing effective leaf area.Science,199,888-890.
Horn HS(1971).The adaptive geometry of trees.Botanical Gazette,142,394-401.
Kikuzawa K(1991).A cost-benefit analysis of leaf habit and leaf longevity of trees and their geographical pattern.The American Naturalist,138,1250-1263.
Kikuzawa K(1995).Leaf phenology as an optimal strategy for carbon gain in plants.Canadian Journal Botany,73,158-163.
King D(1981).Tree dimensions:Maximizing the rate of height growth in dense stands.Oecologia,51,351-356.
King D,Loucks OL(1978).The theory of tree bole and branch form.Radiation and Environmental Biophysics,15,141-165.
King DA(1986).Tree form,height growth,and susceptibility to wind damage in Acer saccharum.Ecology,67,980-990.
Kleiman D,Aarssen LW(2007).The leaf size/number trade-off in trees.Journal of Ecology,95,376-382.
Koike F(1989).Foliage-crown development and interaction in Quercus gilva and Q.acuta.Journal of Ecology,77,92-111.
Li G,Yang D,Sun S(2008).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,22,557-564.
Li YX,Liu YC(1996).The neighbourhood modular interference effects of seedlings of Gordonia acuminata.Journal of Southwest China Normal University(Natural Science),21,271-275.(in Chinese with English abstract)[黎云祥,刘玉成(1996).四川大头茶苗木构件水平的邻体干扰效应.西南师范大学学报自然科学版,21,271-275.]
Liu YF(2015).The Study on Anatomical Structures Character and Its Environmental Adaptation of Leaves from BroadLeaf Woody Plants in Gongga Mountain.Master degree dissertation,Southwest University,Chongqing.(in Chinese with English abstract)[刘艳芳(2015).贡嘎山阔叶木本植物叶片解剖结构特征及其环境适应研究.硕士学位论文,西南大学,重庆.]
Long JN,Smith FW,Scott DRM(1981).The role of Douglasfir stem sapwood and heartwood in the mechanical and physiological support of crowns and development of stem form.Canadian Journal of Forest Research,11,459-464.
Maruyama K(1983).Shoot characteristics as a function of the bud length on Japanese beech trees.Journal of the Japanese Forestry Society,65,43-51.
Mcmahon T(1973).Size and shape in biology.Science,179,1201-1204.
Niklas KJ,Kerchner V(1984).Mechanical and photosynthetic constraints on the evolution of plant shape.Paleobiology,10,79-101.
Niklas KJ(1988).The role of phyllotactic pattern as a“developmental constraint”on the interception of light by leaf surfaces.Evolution,42,1-16.
Pan SA,Peng GQ,Yang DM(2015).Biomass allocation strategies within a leaf:Implication for leaf size optimization.Chinese Journal of Plant Ecology,39,971-979.(in Chinese with English abstract)[潘少安,彭国全,杨冬梅(2015).从叶内生物量分配策略的角度理解叶大小的优化.植物生态学报,39,971-979.]
Pickup M,Westoby M,Basden A(2005).Dry mass costs of deploying leaf area in relation to leaf size.Functional Ecology,19,88-97.
Pitman E(1939).A note on normal correlation.Biometrika,31,9-12.
Reich P,Walters M,Ellsworth D(1992).Leaf life-span in relation to leaf,plant,and stand characteristics among diverse ecosystems.Ecological Monographs,62,365-392.
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