笋瓜节间长发育逆转及其遗传模型研究

分别以短蔓型和长蔓型笋瓜材料为亲本构建6世代群体,测量各群体不同节位节间长的变化,对节间长性状进行遗传规律分析。研究结果表明:节间长性状受核基因控制,F1群体的节间长性状在生长过程中存在发育逆转现象,逆转时期发生在6~15节位之间。遗传模型分析结果显示,在幼苗期和逆转期,笋瓜的节间长受到2对主效基因控制,符合E1模型,主基因方差能解释F2群体方差的70%以上;生长后期则受到主效单基因控制,符合D4模型,但主效基因效应不强,只能解释F2群体方差的10%,且环境因素对植株生长后期节间长具有明显影响。     

小兴安岭不同功能型阔叶植物的柄叶权衡

在叶水平上, 叶柄(支撑结构)与叶片(同化结构)的权衡关系受多种因素的影响, 研究不同功能型植物柄叶性状之间的权衡关系有助于更好地理解植物的生长特性与生活史策略。该研究选定小兴安岭地区典型阔叶红松(Pinus koraiensis)林林内乔木、灌木、草本植物, 采用最小显著差异法比较植物叶片性状与叶柄性状在不同生活型间的变异, 并用标准化主轴法从生活型、叶型、耐阴性3个方面研究叶片与叶柄性状之间的权衡关系。结果表明: (1)不同生活型、叶型植物以及不同耐阴性乔木叶片性状与叶柄干质量之间具有显著的异速生长关系, 且斜率均小于1; (2)随着叶柄干质量增加, 乔木叶片鲜质量、叶片干质量增长速度比灌木、草本更快, 但相同叶柄干质量, 乔木叶柄所能支撑的叶片面积最小; (3)单叶植物叶片面积-叶柄干质量的回归斜率显著大于复叶植物, 叶片鲜质量-叶柄干质量的回归斜率小于复叶植物, 并且相同叶柄干质量下, 单叶植物的叶片干质量总是大于复叶植物; (4)比起喜光树种, 相同叶柄干质量, 耐阴树种的叶柄能够支撑的叶片面积与叶片鲜质量更大, 并且叶柄生物量分配比例(叶柄干质量/叶干质量)与叶片性状的回归斜率均表现为喜光树种>0, 耐阴树种<0。该研究结果表明叶片大小(叶片面积、叶片鲜质量、叶片干质量)与叶柄性状之间存在典型的权衡关系, 其在不同生活型、叶型植物及不同耐阴性乔木内的差异有助于揭示不同功能型植物的生长特性与生活史策略。     

树冠位置对香樟叶形态性状和异速生长关系的影响

为深入了解树冠位置对植物叶形态性状的影响,在常绿乔木香樟树冠上下2层和东南西北4个方位开展调查取样,系统分析了不同树冠位置间叶形态性状(叶长、叶宽、叶厚、叶柄长、叶柄直径和叶形指数)及其异速生长关系的差异性。结果表明,叶形态性状在不同树冠方位间均差异显著,但上下2层变化趋势不完全一致。在树冠上层,除叶形指数和叶炳长外,其余4个性状均表现为东侧最大。在树冠下层,除叶形指数外,其余5个性状指标均表现为东侧最小。在同一方位上,叶形态性状在上下2层间也存在一定差异,其中叶形指数多为下层高于上层,而其他形态性状多呈相反趋势。此外,树冠层次和方位的交互作用对叶片长、叶片厚、叶柄长和叶柄直径有显著影响。各层次和各方位叶形态性状间多为异速生长关系(即异速生长指数不等于1),且多无显著差异。在所有树冠层次和树冠方位,叶宽与叶厚、叶宽与叶炳长、叶长与叶厚及叶长与叶柄长之间均呈异速生长关系。可见,树冠位置对香樟叶形态性状的影响较大,但形态性状间的异速生长关系相对稳定,这是香樟叶形态性状表型可塑性和内在关系稳定性的重要体现。     

葎草种群自疏过程中幼苗构件性状及生物量分配变化

以雌雄异株攀援草本植物葎草为材料,通过每10 d测量1次,连续6次,测定幼苗期葎草种群的密度和高度、个体构件性状和生物量分配等参数,分析种群自疏过程中种群密度与个体构件性状及生物量分配的关系,研究葎草种群的自疏规律。结果表明:幼苗期葎草种群存在显著的自疏现象,种群密度60 d内下降了71%;幼苗期葎草由直立生长向横向生长时,种群密度和株高显著降低;自疏过程中存留植株的茎性状有显著变化,变化大小为节间长主茎长茎直径,节间长增加,叶性状变化大小为叶面积叶柄长叶厚叶宽叶长总叶数保留叶片数,根性状变化大小为总根长根体积根数根长最大根长;自疏过程中存留植株的构件生物量、单株生物量显著增加,而单位面积累积生物量呈阶段性下降;留存植株的地上生物量分配比相对稳定(P0.05),根茎比和叶茎比有极显著变化(P0.01);叶、茎、叶柄生物量与根生物量和地上生物量之间均呈极显著的异速关系(P0.01),茎随地上生物量增长呈等速生长,而叶、叶柄和根随地上生物量增长呈异速生长,地上生物量与叶、茎、叶柄及根生物量极显著相关(P0.01);茎生物量与密度的异速关系遵循最终产量恒定法则,叶、叶柄和根生物量并不满足-3/2或-4/3或-1自疏法则;地上和单株总生物量与密度极显著相关(P0.01),存留单株的地上生物量和总生物量与密度的异速关系遵循最终产量恒定法则。     

樟子松人工林一级枝条基径和枝长模型的研究

以东北林业大学帽儿山实验林场樟子松人工林为研究对象,采用枝解析的方法,于2002年、2003年测定了53株林木（年龄17～38a,直径8．61～21．5cm,树高7．48～18．24m）的树冠变量,建立了基于总着枝深度（DINC）和树冠内一级枝条基径（BD）、枝长（BL）的预估模型。对于大小相同树木的一级枝条,这些树冠变量随着DINC的增加而增大,而林木的胸径（DBH）、树高（HT）变量又很好地反映了不同大小树木的基径和枝长的变化。采用独立检验样本对构建的树冠内一级枝条基径和枝长模型进行了拟合统计量和精度检验,结果表明：模型预测效果良好,精度均达到95％以上。构建的一级枝条基径和枝长模型为进一步合理地描述樟子松人工林树冠的形状及其变化以及三维可视化经营提供依据。     

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。     

沈阳城市森林主要树种叶片形态生态特征与生长规律

利用统计和多元分析方法研究了沈阳城市森林10个主要树种叶片的形态生态特征和生长规律.结果表明,影响树木叶片生长发育节律的主要气象因子是≥5 ℃积温、累积日照时数和各物候期中旬平均温度.不同物候期树木所需要的气象因子有所变化,树木萌动期要求低温和寒冷指数,而在生长期需要一定的温暖指数和湿润指数.10个树种的叶片形态数量特征差异较大,按叶形指数依次为短序胡枝子＞华北丁香＞国槐＞银白杨＞红瑞木＞金银忍冬＞水腊＞水曲柳＞稠李＞黄菠萝;按叶面积依次为华北丁香＞银白杨＞黄菠萝＞稠李＞水曲柳＞红瑞木＞短序胡枝子＞金银忍冬＞国槐＞水腊.在10个树种叶长度与宽度、周长、叶面积之间相互关系中,基本属于异速生长类型,符合y=ax^k模型,但在叶长度与宽度之间关系中,除银白杨k＞1外,其它9个树种均k＜1;在叶长度与周长关系中,黄菠萝k＞1, 银白杨k≈1,呈线性关系,其它树种均k＜1;在叶长度与叶面积关系中,10个树种k＞1,而 银白杨k=2.1028,叶面积增长速率近似于叶长的2倍.建立了10个树种的叶面积最优回归估测模型,并对模型进行了估测检验.     

长苞铁杉天然林群落种内及种间竞争关系研究

通过各种竞争指数的比较筛选,得到较能客观反映长苞铁杉种内、种间竞争关系的竞争指数,定量地分析了长苞铁杉种内和种间竞争强度。结果表明:长苞铁杉种内竞争随胸径的增大而逐渐减少;种间与种内竞争关系顺序为:长苞铁杉-长苞铁杉>猴头杜鹃-长苞铁杉>青冈栎-长苞铁杉>其它树种-长苞铁杉。竞争强度与对象木的胸高直径服从双曲线回归关系,利用模型预测了长苞铁杉种内、种间的竞争强度。     

不同种源樟树叶片形态特征及生长差异分析

为了解不同种源樟树叶片形态特征和生长差异,该文以30个种源樟树为研究对象,对其叶长、叶宽、叶柄长、周长、叶面积、长宽比、形态因子、株高、地径等指标进行测定和差异性分析.结果表明:(1)30个种源间叶片性状的变异系数为3.88％～16.14％,显示不同种源樟树叶片形态特征存在显著差异;叶长、叶宽、叶柄长、周长、面积、叶厚...     

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。     

Petiole mechanics, leaf inclination, morphology, and investment in support in relation to light availability in the canopy of Liriodendron tulipifera

To determine the role of leaf mechanical properties in altering foliar inclination angles, and the nutrient and carbon costs of specific foliar angle variation patterns along the canopy, leaf structural and biomechanical characteristics, biomass partitioning into support, and foliar nitrogen and carbon concentrations were studied in the temperate deciduous species Liriodendron tulipifera L., which possesses large leaves on long petioles. We used beam theory to model leaf lamina as a uniform load, and estimated both the lamina and petiole flexural stiffness, which characterizes the resistance to bending of foliar elements at a common load and length. Petiole and lamina vertical inclination angles with respect to horizontal increased with increasing average daily integrated photon flux density (Q_int). Yet, the light effects on lamina inclination angle were primary determined by the petiole inclination angle. Although the petioles and laminas became longer, and the lamina loads increased with increasing Q_int, the flexural stiffness of both lamina and petiole increased to compensate for this, such that the lamina vertical displacement was only weakly related to Q_int. In addition, increases and decreases in the petiole inclination angle with respect to the horizontal effectively reduced the distance of lamina load from the axis of rotation, thereby reducing the bending moments and lamina inclination due to gravity. We demonstrate that large investments, up to 30% of total leaf biomass, in petiole and large veins are necessary to maintain the lamina at a specific position, but also that light has no direct effect on the fractional biomass investment in support. However, we provide evidence that apart from light availability, structural and chemical characteristics of the foliage may also be affected by water stress, magnitude of which scales positively with Q_int.     

The leaf development process and its significance for reducing self-shading of a tropical pioneer tree species

On a monoaxial erect stem of trees with continuous leafing, the older leaves would be quickly shaded by newer (upper) leaves if the trees did not have any compensating mechanisms to avoid self-shading. We hypothesized that the dynamic adjustment of leaf deployment, by regulating the patterns of leaf growth and by changing leaf orientation as leaves age, is a compensating mechanism. To verify this hypothesis, we analyzed leaf development and crown structure of a Far Eastern tropical pioneer tree species, Macaranga gigantea (Rub. f. et Toll.) M.A., which unfolds huge leaves directly on a monoaxial stem with a short leafing interval. Petioles required more than 90 days for full elongation and the petiole angle (the angle between the petiole axis and the vertical) increased over time. Thus, a series of leaves on a stem progressively increased in petiole length and petiole angle from the youngest to the oldest leaves. This is beneficial because it decreases the degree of self-shading within a crown. A simulation suggested that an average crown for the M. gigantea seedlings, which was constructed using empirically determined morphometric data cannot entirely eliminate self-shading within the crown. But an average crown had a lower degree of self-shading, with less dry mass allocation to the petiole than simulated crowns that were identical to the average crown in all but one respect: they had constant petiole lengths or petiole angles. We conclude that M. gigantea seedlings reduce self-shading by regulating elongation of the petiole and changes in the petiole angle with increasing leaf age.     

The functional morphology of light capture and carbon gain in the Redwood forest understorey plant Adenocaulon bicolor Hook

1. A three-dimensional geometric simulation model of crown architecture was utilized to investigate the efficiency of light capture and its relationship to whole-plant CO₂ assimilation of Adenocaulon bicolor .
2. Positioning of the leaves by the combined effects of ontogenetic variations in petiole length and angle and leaf size, and the leaf divergence angles were shown to be effective in minimizing self shading. The efficiency of light absorption varied from 0·64 to 0·70 among individual plants that were sampled.
3. Plant to plant variation in simulated daily carbon gain was strongly influenced by variations in the direct and diffuse PFD received by the individual plants. When simulations were run for all plants under a single common light environment, the carbon gain was strongly dependent on the efficiencies of light absorption of the different plants.
4. Simulations in which petiole length was varied showed a non-linear dependence of light absorption efficiency on petiole length. When both petiole length and leaf size were varied in a way that maintained a constant biomass then an optimal petiole length that corresponded to the observed petiole length was apparent. The observed divergence angle between successive leaves also maximized light absorption efficiency as compared to greater or lesser angles, but increases in internode length had no significant effect.
5. The results of this study provide evidence for selection for an 'optimal design' of crown architecture in Adenocaulon bicolor that maximizes light capture.     

银杏叶片形态研究

通过对银杏不同枝类、不同叶序的叶片形态进行调查,结果表明:银杏不同枝类的叶片形态差异较大,叶宽、叶柄长、叶基角:多年生鳞枝＞一年生鳞枝＞一年生长枝;叶长、叶形指数:一年生长枝＞一年生鳞枝＞多年生鳞枝;叶面积:一年生和多年生鳞枝＞一年生长枝;有缺刻叶比例:一年生长枝＞多年生鳞枝＞一年生鳞枝.一年生长枝叶片的叶宽、叶长、叶面积、叶柄长、叶基角均随叶序的增加逐渐减小,叶形指数和有缺刻叶的比例则增加.一年生和多年生鳞枝1～5叶的叶长、叶宽和叶面积随叶序增加而逐渐增加,第5～6叶达最大,以后随叶序增加而逐渐减小,叶形指数和叶柄长度随叶序增加而增加,叶基角随叶序增加而减小.一年生长枝的第2叶、一年和多年生鳞枝的第4叶可作为品种描述的标准叶.     

Allometric relationships ofmalva parviflora growing in two different bioclimatic regions

A total of 660 individual plants ofMalva parviflora, a medicinal plant in many countries, growing in two bioclimatic regions were randomly collected with the aim of examining the differences in the allometry of this herbaceous plant growing in two bioclimatic regions. Allometric relationships were found in plant height, stem width, leaf area, leaf length, leaf width, petiole length, and leaf dry weight whereas no relationship was found between plant height or petiole length with specific leaf area. Plants growing in the cool bioclimatic region showed that plant height increases more than the increase in stem width, leaf length, leaf width, and petiole length while plants growing in the warm bioclimatic region showed that plant height increase was lower than that of stem width, leaf length, leaf width, and petiole length. Plant height relationship with root length indicated that in the cool region the plant height increase was less than the increase in the root length while the opposite occurred in the warm region. These differences can be explained by the effects of the different environmental conditions present in the two bioclimatic regions such as water scarcity and availability on the growth ofM. parviflora.     

Relationships between branch spacing, growth rate and light in tropical forest saplings

1. The spacing of branches along central stems was related to growth rate and light level in forest saplings and trees in tropical moist forest on Barro Colorado Island (BCI), Panama. The study included 14 species with tiers of plagiotropic branches (having planar leaf arrangements) and four species with continuous distributions of plagiotropic branches.
2. All species showed increases in branch spacing with increasing light and growth rate of diameter, similar to the patterns in leaf spacing noted previously in species which initially bear large leaves on unbranched stems.
3. Non-tiered species had shorter internodes than tiered species but because the latter bear more branches per node, both groups had similar numbers of branches per unit stem length, when compared at similar growth rates.
4. Differences in the relationship between internode length and growth rate among tiered species were related to demographic characteristics, suggesting that tree architecture may influence forest composition.
5. The strong correlation observed between branch spacing and growth rate suggests that branch spacing may be used to estimate past growth histories of forest tree species with plagiotropic branches.     

Implications of opposite phyllotaxis for light interception efficiency of Mediterranean woody plants

Opposite leaves lead to a greater leaf overlapping than leaves spirally arranged along a shoot, decreasing light interception efficiency (Ea, fraction of the light reaching the plant actually intercepted by the leaves) of the crown. However, Ea results from a whole suite of morphological traits. The interplay between phyllotaxis, crown architecture, leaf morphology and Ea was explored in 12 woody species from Mediterranean-type ecosystems, where the abundance of woody species with opposite phyllotaxis is unusually high. The three-dimensional model Y-plant was used to estimate Ea in unbranched, vertical shoots of each species encompassing the natural morphological variation found from moderate shade to open light environments. Ea exhibited significant interspecific differences, ranging from 0.25 in Daphne gnidium to 0.75 in Cistus ladanifer, Olea europaea and Salvia officinalis, decreasing with leaf inclination angle and leaf area ratio (LAR), and increasing with internode-to-leaf-length ratio and supporting biomass. Species with spiral vs. opposite phyllotaxis did not differ in their mean Ea. However, the former had higher Ea than the latter at short internode lengths. The natural range of variation in internode length had a larger effect on Ea than the natural range of leaf elevation angle. Principal component analysis segregated species with opposite phyllotaxis from those with spiral leaves because of their greater self-shading for high sun elevation angles (>45°); they were in turn distributed in two groups, one with high Ea, large investment in supporting biomass and long internodes, and another with low Ea and large LAR. Species with spiral phyllotaxis all had intermediate or low Ea and steep leaf elevation angles. Species with opposite phyllotaxis can compensate their less efficient leaf arrangement by decreasing leaf elevation angle and increasing internode length, but they may experience a real phylogenetic constraint for light interception when biomass allocation to supporting tissues (internodes and petioles) becomes very costly. This constraint could be involved in the shade intolerance of woody Mediterranean species exhibiting opposite phyllotaxis.     

优良单株家系辣木叶的表型性状分析

为挖掘辣木(Moranga oleifera)优良种质资源,对30个优良单株家系的叶片表型性状进行研究。结果表明,除叶形外,辣木不同家系间的叶柄和叶片颜色、复叶数、复叶柄长度和直径、复叶间距、叶长、叶宽均存在不同程度的差异。复叶数与复叶柄长度和直径、复叶间距、叶长、叶宽呈极显著正相关;主成分分析表明,叶长、叶宽、复叶柄长度和直径、复叶间距、叶柄和叶片颜色是区分辣木不同家系最主要的叶片性状指标。聚类分析结果表明,30个辣木家系可分为3大类,叶片表型性状存在显著差异的家系的遗传距离较远。因此,叶柄和叶片颜色、复叶数、复叶柄长度和直径、复叶间距、叶长、叶宽将为直观区分辣木家系提供参考。     

Molecular genetics of growth and development in Populus (Salicaceae). v. mapping quantitative trait loci affecting leaf variation

We examined the genetic variation of leaf morphology and development in the 2-yr-old replicated plantation of an interspecific hybrid pedigree of Populus trichocarpa T. & G. and P. deltoides Marsh. via both molecular and quantitative genetic methods. Leaf traits chosen were those that show pronounced differences between the original parents, including leaf size, shape, orientation, color, structure, petiole size, and petiole cross section. Leaves were sampled from the current terminal, proleptic, and sylleptic branches. In the F2 generation, leaf traits were all significantly different among genotypes, but with significant effects due to genotype X crown-position interaction. Variation in leaf pigmentation, petiole length. And petiole length proportion appeared to be under the control of few quantitative trait loci (QTLs). More QTLs were associated with single leaf area, leaf shape, lamina angle, abaxial color, and petiole flatness, and in these traits the number of QTLs varied among crown positions. In general, the estimates of QTL numbers from Wright's biometric method were close to those derived from molecular markers. For those traits with few underlying QTLs, a single marker interval could explain from 30 to 60% of the observed phenotypic variance. For multigenic traits, certain markers contributed more substantially to the observed variation than others. Genetic cluster analysis showed developmentally related traits to be more strongly associated with each other than with unrelated traits. This finding was also supported by the QTL mapping. For example, the same chromosomal segment of linkage group L seemed to account for 20% of the phenotypic variation of all dimension-related traits, leaf size, petiole length. and midrib angle. In both traits. the P. deltoides alleles had positive effects and were dominant to the P. trichocarpa alleles. Similar relationships were also found for lamina angle. abaxial greenness, and petiole.     

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).