武夷山常绿阔叶林木本植物小枝生物量分配

小枝是木本植物的重要组成单元,研究其生物量分配策略有利于了解不同物种对环境的响应。本文以武夷山常绿阔叶林群落中19种木本植物的小枝为研究对象,采取标准化主轴回归(standardized major axis,SMA)的方法分析小枝的叶重、叶片面积、叶柄重、茎重、叶数量等性状特征,研究在群落水平上和不同生活型上,常绿阔叶林群落木本植物的小枝生物量分配格局。结果表明:群落水平上小枝重、总叶重和茎重三者间均呈等速生长关系。同样的,在乔木和灌木不同生活型分类中,茎重和总叶重也呈等速生长关系。但是,小枝重和总叶重、茎重之间的关系却不一致:小枝重和总叶重在乔木、灌木中分别呈等速生长关系和异速生长关系;小枝重和茎重在乔木、灌木中分别呈异速生长关系和等速生长关系;此外,单叶重和出叶强度在群落水平上呈负等速生长关系。然而,这种负等速生长关系并不存在于不同生活型的植物中。小枝水平上,总叶柄重和小枝重、总叶重均呈小于1的异速生长关系,即随叶的增大,小枝需要分配更多生物量用于运输组织和支撑结构(叶柄)的构建。因此,叶柄投资是限制亚热带常绿阔叶林当年生枝和叶增大的重要因素,对于小枝和叶大小的优化选择具有重要影响。     

Slope-related variations in twig and leaf traits of Robinia pseudoacacia in the northern mountains of Lanzhou

 枝叶生长关系是植物在环境胁迫条件下提高空间资源利用能力的一种策略, 弄清枝叶生长关系对理解植物应对环境异质性的表型可塑性具有重要意义。该文利用ArcGIS建立研究区域的数字高程模型, 研究了兰州市北山不同坡向人工林刺槐(Robinia pseudoacacia)小枝茎截面积-总叶面积和出叶强度-单叶面积的生长关系。结果表明: 随着坡向由北坡向东坡、南坡和西坡转变, 刺槐林群落的郁闭度、高度和土壤含水量呈现先减小后增大的趋势, 刺槐小枝茎截面积、总叶面积和单叶面积呈现先减小后增大的趋势, 出叶强度呈现先增大后减小的趋势; 北坡、东坡、南坡和西坡4个坡向的刺槐小枝的茎截面积和总叶面积均呈显著的正相关关系(p < 0.05), 并存在显著大于1的共同斜率, 刺槐的出叶强度与单叶面积均呈显著的负相关关系(p < 0.05), 并存在接近于–1的共同斜率; 随着坡向由北坡向东坡、南坡和西坡转变, 茎截面积-总叶面积和出叶强度-单叶面积两组关系的回归方程截距呈现先减小后增大的趋势。刺槐枝叶在不同坡向上的异速生长关系反映了植物功能性状对生长环境的响应和适应, 以及植物构型构建的投资权衡机制。     

我国温带山地森林48种常见树种叶片重量-出叶强度的关系

叶片是植物的主要光合器官, 其质量与数量的权衡关系体现植物对环境的适应策略。在全球气候变化的背景下, 研究叶片质量与数量关系有助于理解植物对环境变化的响应趋势。该研究应用标准化主轴回归方法, 探讨了我国温带山地森林中48个常见树种的单叶干重与出叶强度的权衡关系。结果表明, 所有物种以及落叶阔叶林、常绿和落叶阔叶树种、单叶以及亚冠层阔叶树种的单叶干重与出叶强度表现为异速生长关系; 针叶林、针阔混交林、常绿及落叶针叶树种、复叶以及冠层阔叶树种则表现为等速生长关系。研究结果表明, 叶大小和出叶强度并无恒定的权衡关系。     

高寒退化草地狼毒种群个体大小与茎、叶的异速生长

叶片和茎的生长规律决定植物对光截取的模式以及与外界环境的物质交换面积,其生物量分配是研究植物生活史对策的一个重要途径.在石羊河上游高寒退化草地,采用标准化主轴估计(standardized major axis estimation,SMA)方法,研究了狼毒(Stellera chamaejasme)种群茎大小对茎与叶生物量分配的影响,以及对叶面积支持效率的影响.结果表明:狼毒种群茎干重与比叶重(LMA)、单叶面积、叶干重呈异速生长关系,SMA斜率分别为0.781(95％的置信区间,CI=O.71 ～0.94)、0.824(CI=0.77 ～0.95)和0.856(CI=0.79 ～0.99).狼毒种群中较大茎的狼毒植株具有较高的叶生物量分配比例和叶面积支持效率,同时具有较高的比叶重,说明较大的狼毒采取以提高资源利用效率为主的适应策略,而相对较小的狼毒采取以快速生长为主的适应策略.     

亚热带常绿林不同冠层小枝叶面积-叶生物量关系研究

叶面积与叶生物量的关系对于理解植物叶片的碳收益和投资权衡策略具有重要意义。收益递减假说认为植物的叶面积与叶生物量成显著异速生长关系，其异速生长指数<1.0，但该假说是否适用于不同生活型（常绿与落叶）亚热带木本植物不同冠层高度（上下冠层）当年生小枝的叶片仍不清楚。以江西亚热带常绿阔叶林的69种常绿与落叶木本植物当年生小枝上的叶为研究对象，采用标准化主轴回归估计（standardized major axis estimation，SMA）方法检验不同冠层高度和生活型叶面积与叶生物量的异速生长关系。结果显示：（1）当年生小枝叶生物量在不同冠层高度和生活型的植物中无显著差异（P>0.05），叶面积在常绿和落叶植物中有显著差异（P<0.05），常绿和落叶植物的比叶重存在显著差异（P<0.05），而落叶植物的比叶重在不同冠层高度存在显著差异（P<0.05），同一冠层，常绿植物比叶重显著高于落叶植物（P<0.05）；（2）69种植物的叶面积与叶生物量异速生长指数具有物种特异性，60.9%的物种叶面积与叶生物量呈等速生长关系；（3）不同冠层和生活型植物的叶面积与叶生物量呈等速生长关系，但其异速生长常数在不同冠层高度与生活型间存在差异。这些结果表明冠层高度和生活型未改变叶面积-生物量之间的等速生长关系，不支持"收益递减"假说。     

EFFECTS OF TWIG SIZE ON BIOMASS ALLOCATION WITHIN TWIGS AND ON LAMINA AREA SUPPORTING EFFICIENCY IN RHODODENDRON: AL. LOMETRIC SCALING ANALYSES

 当年生小枝是多年生植物体上最活跃的部分之一, 其生物量分配是植物生活史对策研究的一个重要内容。该文采用标准化主轴估计(Standardized major axis estimation, SMA)和系统独立比较分析(Phylogenetically independent contrast analysis, PIC)的方法, 研究了杜鹃花属(Rhododendron)植物一年生小枝的大小对小枝叶片、叶柄和茎的生物量分配的影响, 以及对叶面积支持效率(即单位质量小枝支持的叶面积)的影响。结果显示: 1)小枝大小对叶片生物量分配比率的影响不显著, SMA斜率为1.040 (95%的置信区间(CI)=0.998～1.085); 但是, 小枝越大, 叶柄生物量分配比例越高(SMA斜率为1.245, 显著大于1.0, 呈显著的异速生长关系)。2)小枝越小, 单叶面积越小(支持Corner法则), 单位质量小枝所支持的叶面积越大, 即具有较小枝条和较小叶片的物种可能具有较高的叶面积支持效率。这些结果有助于我们更好地理解亲缘关系十分接近的杜鹃花属植物, 在不同生境条件下叶片大小的差异, 以及为什么在胁迫生境条件下小叶物种更为常见。     

不同气候带间成熟林植物叶性状间异速生长关系随功能型的变异

为研究不同生物气候带内植物叶片大小与叶柄干重间的异速生长关系, 探讨不同植物功能型对叶内异速生长关系的效应, 在黑龙江呼中、吉林长白山、北京东灵山、浙江古田山、湖北神农架和四川都江堰6个地区, 选择典型地带性成熟林进行主要木本植物的叶片和叶柄性状的测定与统计分析。结果表明: 不同功能型和气候带植物叶片干重、面积、体积均与叶柄干重之间存在着显著的异速生长关系, 共同斜率分别为0.82、0.70和0.80, 均显著小于1.0。在相同叶柄干重下, 灌木较乔木支持更大的叶片体积, 但它们支持的叶片干重与叶片面积无显著差异; 常绿植物在给定叶柄干重下较落叶植物支持更高的叶片干重与体积, 但其支持的叶片面积小于落叶植物; 除神农架地区外, 在给定叶柄干重下, 亚热带的古田山、都江堰地区的植物较温带地区的植物支持更大的叶片干重、面积与体积, 而亚热带神农架地区的植物叶柄支持的叶片大小(面积、体积、干重)与温带地区相近。结果表明, 叶柄限制了叶片的不断增大(包括面积、体积和干重), 叶片和叶柄之间的异速生长关系受功能型、气候带及生境条件的影响。     

上海辰山植物园不同生活型木本植物枝叶大小关系的比较

植物枝叶生长普遍存在显著的正相关关系，决定了植物构型塑造和生物量分配。以上海辰山植物园149种木本植物为对象，通过测定顶枝上70 cm长枝条的直径、叶面积及其生物量，比较相同生境条件下不同生活型木本植物的枝叶大小关系。结果表明，枝条截面积与总叶面积间呈异速生长关系（a=1.148 6，CI=1.000 6~1.302 3），枝条干重与叶干重间呈等速生长关系（a=1.054 2，CI=0.921 3~1.205 6），不同生活型均具有相同的斜率系数a。不同生活型的异速生长常量b（y轴截距）存在显著差异，相同枝条截面积下落叶乔木比常绿乔木和常绿灌木具有更大的叶面积，相同枝条干重下常绿乔木和落叶乔木比常绿灌木具有更大的叶干重。这可能与不同生活型木本植物水分竞争效率和叶建成成本差异有关。     

常绿和落叶阔叶木本植物小枝内生物量分配关系研究：异速生长分析

生物量分配是植物净碳获取的重要驱动因素,当年生小枝内部的生物量分配是植物生活史对策研究的一个重要内容。本文采用标准化主轴估计(Standardized major axis estimation,SMA)和系统独立比较分析(Phy1ogenetica11y independent contrast analysis,PIC)的方法,研究了贡嘎山常绿和落叶阔叶木本植物当年生小枝内各组分生物量分配之间的关系。结果显示：小枝干重与茎干重、叶(含叶柄)干重和叶片(不含叶柄)干重,以及茎干重与叶(含叶柄)干重之间均呈极显著的等速生长关系,表明分配到叶或者叶片中的生物量独立于小枝生物量;叶柄干重与叶片(不含叶柄)干重和小枝干重呈极显著的异速生长关系,显示叶柄对小枝内的生物量分配具有重要的影响;在某一给定的小枝干重或叶片干重时,常绿物种比落叶物种具有更大的叶柄干重,即更大的支撑投资。这些结果表明叶柄作为叶片生物量最大化的一个不利因素,影响了小枝内的生物量分配,并且叶片与其支撑结构之间的异速生长关系随叶片生活型的变化而变化。     

兰州北山刺槐枝叶性状的坡向差异性

枝叶生长关系是植物在环境胁迫条件下提高空间资源利用能力的一种策略, 弄清枝叶生长关系对理解植物应对环境异质性的表型可塑性具有重要意义。该文利用ArcGIS建立研究区域的数字高程模型, 研究了兰州市北山不同坡向人工林刺槐(Robinia pseudoacacia)小枝茎截面积-总叶面积和出叶强度-单叶面积的生长关系。结果表明: 随着坡向由北坡向东坡、南坡和西坡转变, 刺槐林群落的郁闭度、高度和土壤含水量呈现先减小后增大的趋势, 刺槐小枝茎截面积、总叶面积和单叶面积呈现先减小后增大的趋势, 出叶强度呈现先增大后减小的趋势; 北坡、东坡、南坡和西坡4个坡向的刺槐小枝的茎截面积和总叶面积均呈显著的正相关关系(p < 0.05), 并存在显著大于1的共同斜率, 刺槐的出叶强度与单叶面积均呈显著的负相关关系(p < 0.05), 并存在接近于-1的共同斜率; 随着坡向由北坡向东坡、南坡和西坡转变, 茎截面积-总叶面积和出叶强度-单叶面积两组关系的回归方程截距呈现先减小后增大的趋势。刺槐枝叶在不同坡向上的异速生长关系反映了植物功能性状对生长环境的响应和适应, 以及植物构型构建的投资权衡机制。     

The leaf size-twig size spectrum of temperate woody species along an altitudinal gradient: an invariant allometric scaling relationship

BACKGROUND AND AIMS: The leaf size-twig size spectrum is one of the leading dimensions of plant ecological variation, and now it is under development. The purpose of this study was to test whether the relationship between leaf size and twig size is isometric or allometric, and to examine the relationship between plant allometric growth and life history strategies in the spectrum. METHODS: Leaf and stem characters-including leaf and stem mass, total leaf area, individual leaf area, stem cross-sectional area, leaf number and stem length-at the twig level for 59 woody species were investigated along an altitudinal gradient on Changbaishan Mountain in the temperate zone of China. The environmental gradient ranges from temperate broad-leaved mixed forest at low altitude, to conifer forest at middle altitude, and to sub-alpine birch forest at high altitude. The scaling relationships between stem cross-sectional area and stem mass, stem mass and leaf mass, and leaf mass and leaf area at the twig level were simultaneously determined. KEY RESULTS: Twig cross-sectional area was found to have invariant allometric scaling relationships with the stem mass, leaf mass, total leaf area and individual leaf area, all with common slopes being significantly larger than 1, for three altitudinal-zoned vegetation types under investigation. However, leaf mass was found to be isometrically related to stem mass and leaf area along the environmental gradient. Based on the predictions of previous models, the exponent value of the relationship between twig cross-sectional area and total leaf area can be inferred to be 1.5, which falls between the confidence intervals of the relationship at each altitude, and between the confidence intervals of the common slope value (1.17-1.56) of this study. This invariant scaling relationship is assumed to result from the fractural network and/or developmental constraints of plants. The allometric constants (y-intercepts) of the relationships between the stem cross-sectional area and leaf area (both total leaf area and individual leaf area) were found to decrease significantly along the altitudinal gradient. This suggests that the species would support less leaf area at a given twig cross-sectional area with increasing environmental stress. CONCLUSIONS: This study demonstrated that plants respond to the environmental gradient by changing the y-intercepts of the relationship between leaf size-twig size, while keeping the exponent value of the allometric relationship as an invariant constant. The allometric growth in the twig size-leaf size spectrum is related to many other components of plant life history strategy, including the well established life history trade-off between efficiency and safety in the hydraulic transport of water.     

The effects of leaf size,leaf habit,and leaf form on leaf/stem relationships in plant twigs of temperate woody species

Question: Do thick‐twigged/large‐leaf species have an advantage in leaf display over their counterparts, and what are the effects of leaf habit and leaf form on the leaf‐stem relationship in plant twigs of temperature broadleaf woody species? Location: Gongga Mountain, southwest China. Methods: (1) We investigated stem cross‐sectional area and stem mass, leaf area and leaf/lamina mass of plant twigs (terminal branches of current‐year shoots) of 89 species belonging to 55 genera in 31 families. (2) Data were analyzed to determine leaf‐stem scaling relationships using both the Model type II regression method and the phylogenetically independent comparative (PIC) method. Results: (1) Significant, positive allometric relationships were found between twig cross‐sectional area and total leaf area supported by the twig, and between the cross‐sectional area and individual leaf area, suggesting that species with large leaves and thick twigs could support a disproportionately greater leaf area for a given twig cross‐sectional area. (2) However, the scaling relationships between twig stem mass and total leaf area and between stem mass and total lamina mass were approximately isometric, which indicates that the efficiency of deploying leaf area and lamina mass was independent of leaf size and twig size. The results of PIC were consistent with these correlations. (3) The evergreen species were usually smaller in total leaf area for a given twig stem investment in terms of both cross‐sectional area and stem mass, compared to deciduous species. Leaf mass per area (LMA) was negatively associated with the stem efficiency in deploying leaf area. (4) Compound leaf species could usually support a larger leaf area for a given twig stem mass and were usually larger in both leaf size and twig size than simple leaf species. Conclusions: Generally, thick‐twigged/large‐leaf species do not have an advantage over their counterparts in deploying photosynthetic compartments for a given twig stem investment. Leaf habit and leaf form types can modify leaf‐stem scaling relationships, possibly because of contrasting leaf properties. The leaf size‐twig size spectrum is related to the LMA‐leaf life span dimension of plant life history strategies.     

Testing of corner’s rules across woody plants in Tiantong region,Zhejiang Province: effects of micro-topography

Aims The size and quantity relationships between twigs and leaves can be used to describe the hydraulic properties of plants in response to environmental stresses. The objective of this study was to examine how twig-leaf relationship would vary with changes in micro-habitat conditions.
Methods The study site is located in the Tiantong National Forest Park (29.87° N, 121.65° E), Zhejiang Province. We measured twig cross-sectional area (twig size), sub-twig cross-sectional area (sub-twig size), individual leaf area, total leaf area (leaf size per twig), the number of twigs at a given twig size (twig intensity), and the number of leaves at a given twig size (leafing intensity) across individual woody plants on 10 plots in each of the convex and concave habitats within an evergreen broad-leaved forest. The standardized major axis (SMA) analysis was conducted to determine the scaling relationships between twig size and leaf size, between sub-twig size and twig intensity, and between leaf size and leafing intensity.
Important findings Significant, positive allometric relationships between twig cross-sectional area and total leaf area were found in plants in both types of micro-habitats (p < 0.001). There was no significant difference between the two micro-habitats in the slope of the regression between twig cross-sectional area and total leaf area, and the common slope of the regressions was significantly greater than 1 (p < 0.001). The intercept was significantlygreater in plants of the concave habitat than in plants of the convex habitat (p < 0.001), indicating that plants in a concave habitat support greater total leaf area at a given twig size than in a convex habitat. Significant, negative allometric scaling relationships were found between twig size and twig intensity in plants in both micro-habitats. There was also no significant difference between the two habitats in the slope of the regression between twig size and twig intensity, and the common slope of the regressions was significantly less than –1 (p < 0.001). The similar intercept in the regression relationship of twig area and twig intensity between the two habitats suggests that plants deploy similar amount of sub-twigs per twig size in both types of habitat. In addition, significant, negative allometric scaling relationships between leaf size and leafing intensity were found to be consistently conserved across micro-habitat types, with the common slope being smaller than –1. A higher value of y-intercept in the scaling relationships of leaf area vs. leafing intensity for plants in the concave habitat indicates that at a given leaf area, more leaves were supported by plants in a concave habitat than in a convex habitat. Overall, plants in a concave habitat tend to deploy more large leaves per twig size than those in a convex habitat. This study demonstrated that both the Corner’s rules and the leaf size-number trade-offs could be generalized to apply at the small local spatial scales. The magnitude and quantitative adjustment of twig-leaf deployment manifests a selection preference of hydraulic properties of plants in coping with changes in water availability between concave and convex habitats.     

Within-twig biomass allocation in subtropical evergreen broad-leaved species along an altitudinal gradient: allometric scaling analysis

We studied the effects of twig size and altitude on biomass allocation within plant twigs (terminal branches of current-year shoots), to determine whether species with large twigs/leaves or living at low altitude allocate a higher proportion of biomass to laminas than their counterparts with small twigs/leaves or living at high altitude. Stem mass, lamina mass and area, and petiole mass were measured for terminal branches of current-year shoots in 80 subtropical evergreen broad-leaved species belonging to 38 genera from 24 families along an altitudinal gradient of Mt. Emei, Southwest China. The scaling relationships between the biomass allocations of within-twig components were determined using model type II regression method. Isometric relationships were found between leaf mass and twig mass and between lamina mass and twig mass, suggesting that the biomass allocation to either leaves or laminas was independent of twig mass. Petiole mass disproportionally increased with both lamina mass and twig mass, indicating the importance of leaf petioles to the within-twig biomass allocation. These cross-species correlations were consistent with those among evolutionary divergences. In addition, species at low altitude tended to have a greater leaf and lamina mass but a smaller stem mass at a given twig mass than at middle and high altitudes. This is possibly due to the high requirement in physical support and the low efficiency of eco-physiological transport for the species living at high altitude. In general, within-twig biomass allocation pattern was not significantly affected by twig size but was greatly modulated by altitude.     

亚热带常绿阔叶林木本植物幼树阶段抽枝展叶的权衡关系

亚热带常绿阔叶林植物幼树阶段适应林内生境并开枝散叶是其长成大树的一个重要过程, 植物一年内多次抽枝的现象及其在抽枝展叶过程中小枝伸长、枝茎增粗与叶面积的增加优先顺序及其内在驱动机制还有待进一步研究。该研究对青城山常绿阔叶林木本植物多次抽枝发生比例进行了调查, 并以茶(Camellia sinensis)、细枝柃(Eurya loquaiana)、短刺米槠(Castanopsis carlesii var. spinulosa)、润楠(Machilus nanmu)和大叶山矾(Symplocos grandis) 5种植物的幼树为研究对象, 比较分析了植物在多次抽枝中小枝和叶片生长动态及适应策略的差异。结果显示: 1)一、二次抽枝分别开始于春季(4月)和夏末(8月下旬), 小枝水平上二次抽枝率乔木小于灌木, 常绿植物小于落叶植物。2)一次抽枝小枝枝长、单叶面积, 小枝直径和叶片数量(除大叶山矾外)均高于二次抽枝, 但二次抽枝单叶面积相对生长速率均高于一次抽枝, 二次抽枝叶片比叶质量(LMA)的增长速率高于一次抽枝。3)一次抽枝小枝枝长、叶片数量、小枝直径(除细枝柃和短刺米槠外)和总叶面积(除短刺米槠外)最大相对生长速率均高于二次抽枝, 且大部分物种最大相对生长速率出现在抽枝开始的第一、二周。4)两次抽枝中, 物种先侧重于叶片的生长, 其次是小枝枝长的生长, 最后是小枝直径的增粗。单叶面积和总叶面积皆随着小枝枝长和小枝直径的增加呈显著的异速生长关系, 表明叶片的增长速度大于小枝。单叶面积与叶片数呈显著大于1的异速生长关系, 暗示单叶面积的增长速度大于叶片数的增加速度。小枝枝长与小枝直径也呈显著大于1的异速生长关系, 揭示小枝枝长的增长速度大于小枝直径。综上所述, 两次抽枝过程中, 植物枝叶的优先生长顺序反映了植物为获取更多的资源(尤其是光源)而形成特定的抽枝展叶策略; 二次抽枝单叶面积相对生长速率和LMA增长速率高于一次抽枝, 这可能与植物即将面临的昆虫取食和气温降低压力有关。因此, 了解植物抽枝策略对于理解物种生态适应机制, 揭示物种生活史过程中存在的权衡关系具有重要的理论意义。     

胡杨当年生小枝茎构型对展叶效率的影响

枝叶异速生长关系对理解荒漠植物生物量分配和生活史策略具有重要意义。该研究以小枝茎长度、茎纤细率和茎体积表征茎构型, 以叶密度(单位茎长度的叶数量)、叶面积比(单位小枝茎干质量的总叶面积)和叶茎质量比(单位小枝茎干质量的总叶干质量)表征展叶效率, 采用标准化主轴回归(SMA)方法研究胡杨(Populus euphratica)当年生小枝茎构型对展叶效率的影响及二者沿地下水埋深(GWD)梯度的权衡策略。结果显示, 胡杨当年生小枝茎直径、展叶效率和比叶面积、叶大小随GWD增加而降低, 小枝茎长度、茎纤细率及其上的叶数量则增大。小枝茎构型性状均与展叶效率呈显著负相关关系, 即随小枝茎长度、茎纤细率和茎体积的增加, 展叶效率逐渐降低, 这可能是枝叶大小、水分传导与机械支撑间的权衡结果。小枝茎构型与展叶效率的异速生长指数(斜率)随GWD增加而增大, 是由于单位小枝茎投资获得的叶面积或者叶质量降低所致, 反映出随GWD增加胡杨采取了高消耗低收益的保守型策略。胡杨应对环境压力时, 倾向于在长枝上着生数量较多的小叶, 短枝上着生数量较少的大叶, 体现出胡杨小枝的资源利用策略与枝叶大小的权衡机制。综上所述, GWD显著影响胡杨小枝茎构型-展叶效率的权衡关系, 低展叶效率是胡杨应对日益旱化荒漠环境的适应策略。     

武夷山不同海拔黄山松枝叶大小关系

对武夷山自然保护区不同海拔黄山松叶片面积、叶片数量、小枝长度及小枝直径等性状进行测定,分析不同海拔黄山松枝-叶大小间的权衡关系.结果表明: 随海拔升高,黄山松小枝的叶片数量、小枝长度、小枝直径、出叶强度及茎截面积逐渐增大,单叶面积呈逐渐减小趋势;不同海拔黄山松小枝出叶强度与单叶面积均呈显著负相关,不同海拔黄山松小枝茎截面积与总叶面积呈显著正相关;不同海拔黄山松小枝长度、小枝直径与出叶强度呈显著负相关,与单叶面积、叶片数量及总叶面积均呈显著正相关.为提高竞争优势或是资源利用效率,低海拔黄山松倾向于在短枝上着生量少但单叶面积大的针叶,而高海拔黄山松趋向于在长枝上着生量大但单叶面积小的针叶,这体现出不同海拔梯度黄山松小枝的资源利用策略及枝叶间生物量分配的权衡机制.     

Effect of stem length to stem slender ratio of current-year twigs on the leaf display efficiency in evergreen and deciduous broadleaved trees

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 of mechanical 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).     

清凉峰不同海拔木本植物小枝内叶大小-数量权衡关系

权衡关系是生活史对策理论的基础, 叶大小-数量的权衡关系对理解叶大小进化具有重要的意义。该研究以单叶面积和单叶片干重表示叶大小, 用小枝干重和小枝茎干重表示小枝大小, 采用标准化主轴估计(standardized major axis estimation, SMA)和系统独立比较分析(phylogenetically independent contrast analysis, PIC)的方法, 对浙江省清凉峰自然保护区3个不同海拔落叶阔叶木本植物当年生小枝内的叶大小与数量间的关系进行研究。结果显示, 无论叶大小和小枝大小是用面积或干重表示, 在每个海拔, 叶大小与出叶强度均存在显著的等速负相关关系, 表明在落叶阔叶木本植物中发现的叶大小与出叶强度之间的权衡关系在不同生境物种中是普遍存在的, 植物在叶大小方面的种间变化, 可能不是自然选择的直接产物, 而是叶片数量变化权衡关系的一个副产物。不同海拔间的比较显示, 高海拔物种的叶面积或干重与出叶强度相关关系的y轴截距比中、低海拔物种小, 表明在出叶强度一定的情况下, 高海拔物种比低海拔物种具有更小的叶大小。与高海拔物种相比, 中海拔物种的共同斜率沿共同主轴有一个向上的位移, 表明中海拔物种比高海拔物种具有更大的叶大小, 但出叶强度更小。这些结果表明生境对叶大小-数量的权衡关系具有显著的影响, 中海拔生境具有更适宜植物生长的气候及养分条件, 而高海拔的低温等不利影响使得叶片变小。