生物量分配影响三种不同海拔起源的松树生长

松属的思茅松（Pinus kesiya var. langbianensis）、云南松（P. yunnanensis）和高山松（P. densata）是组成中国西南不同海拔针叶森林的主要树种,然而这三个树种在发育速度尤其是高生长方面表现出明显的差异。为了弄清引起这些变异的生理和形态学原因,本文将三种松树种植于同一环境下,对其光合作用、生物量分配、生长速率和叶片性状进行了研究。研究发现,与来源于高海拔的树种相比,低海拔的树种有更高的株高、以及更大的干物质重量、相对生长速率、叶质比、茎质比和比叶面积,但叶片氮含量、碳含量和根质比较低。高海拔树种的光合速率并不明显低于低海拔树种。相对生长速率和树高均与叶质比呈显著正相关,与根质比负相关,但与最大光合速率没有显著关系。这些结果表明,生物量的分配式样和长期的形态特性能够更好地预测不同海拔松树的生长表现。     

生物量分配影响三种不同海拔起源的松树生长

松属的思茅松（Pinus kesiya var．1angbianensis）、云南松（P．yunnanensis）和高山松（P．densata）是组成中国西南不同海拔针叶森林的主要树种,然而这三个树种在发育速度尤其是高生长方面表现出明显的差异。为了弄清引起这些变异的生理和形态学原因．本文将三种松树种植于同一环境下,对其光合作用、生物量分配、生长速率和叶片性状进行了研究。研究发现,与来源于高海拔的树种相比,低海拔的树种有更高的株高、以及更大的干物质重量、相对生长速率、叶质比、茎质比和比叶面积,但叶片氮含量、碳含量和根质比较低。高海拔树种的光合速率并不明显低于低海拔树种。相对生长速率和树高均与叶质比呈显著正相关,与根质比负相关,但与最大光合速率没有显著关系。这些结果表明,生物量的分配式样和长期的形态特性能够更好地预测不同海拔松树的生长表现。     

植物光合结构与非光合结构的功能平衡：来自三种亚热带乔木树种的实验证据

植物的地上部分和地下部分存在功能性平衡现已十分清楚,但植物的地上部分是否在其光合结构（叶组织）和非光合结构（枝和茎）之间也存在功能性平衡尚不明晰,本文提出两个研究假设并检验之：1）植物地上部分在其光合与非光合结构之间存在功能性平衡;2）此功能性平衡的维持依赖于对光合和非光合结构生物量分配的调节,为验证此假设,采用枝叶修剪的方式（连续两年修剪,四个修剪强度：0,20%,50%,70%）对3种亚热带乔木树种榕（Ficus microcarpa),黄桷树（Ficus virens)和樟（Cinnamomum camphora)进行了研究。结果表明,修剪使所有树种地上部分的光合与非光合结构生物量比率（P/NP）立即下降,下降程度随修剪强度的程式高而增大,但不论是首次修剪还是第二次修剪,修剪处理一年后,修剪株地上部分的光合与非光合结构生物量比率升高,且此生物量比率不低于非修剪株的光合与非光合结构生物量比率。此研究结果证实了植物地上部分光合与非光合结构间存在功能性平衡的假设,与假设一致。植株的生物量分配在修剪后发生了改变;修剪株加大了对光合结构（叶组织）的生物量分配（大量的新生产地上部分生物量被分配到光合结构）,同时却减少了对非光合结构（枝和茎）的分配,此分析格局的改变保证了光合与非光合结构功能性平衡的恢复与维持,可以认为,通过改变生物量分配格局以维持光合与非光合结构功能平衡是植物低御外来干扰和/或损伤的一种有效策略。     

Exogenous and endogenous determinants of spatial aggregation patterns in Tibetan Plateau meadow vegetation

Aims We aim to quantify the relative importance of various endogenous and exogenous processes influencing the spatial distribution of the individuals of plant species at different temporal and spatial scales in a species-rich and high-cover meadow in the eastern Tibetan Plateau.Methods We calculated Green's index of dispersion to infer the spatial distribution patterns of 73 herbaceous species at two scales (0.25 and 1.0 m 2). We constructed a series of generalized linear models to test the hypotheses that different species traits such as mean plant stem density, per capita dry biomass, maximum plant height and mean seed mass contribute to their spatial distribution. We used the first principal component of soil C, N and P to explain abundance variation across quadrats and sub-plots.Important findings The individuals of the species studied were highly spatially aggregated. At both spatial scales, biomass and stem density explained the most variation in aggregation, but there was no evidence for an effect of mean seed mass on aggregation intensity. The effects of soil carbon, nitrogen and phosphorus at different depths affected plant abundance mostly at the broader spatial scale. Our results demonstrate that self-thinning and habitat heterogeneity all contribute to determine the spatial aggregation patterns of plant individuals in alpine meadow vegetation in the eastern Tibetan Plateau.     

Allometric theory and the mechanical stability of large trees: proof and conjecture

Recent allometric theory has postulated that standing leaf mass will scale as the 3/4 power of stem mass and as the 3/4 power of root mass such that stem mass scales isometrically with respect to root mass across very large vascular plant species with self-supporting stems. We show that the isometric scaling of stem mass with respect to root mass (i.e., M(S) ∝ M(R)) can be derived directly from mechanical theory, specifically from the requirement that wind-induced bending moments acting at the base of stems must be balanced by a counter-resisting moment provided by the root system to prevent uprooting. This derivation provides indirect verification of the allometric theory. It also draws attention to the fact that leaf, stem, and root biomass partitioning patterns must accommodate the simultaneous performance of manifold functional obligations.     

Linking multiple-level tree traits with biomass accumulation in native tree species used for reforestation in Panama

To improve establishment yield and carbon accumulation during reforestation, analyses of species adaptations to local environments are needed. Here we measured, at the individual scale, links between biomass accumulation and multiple-level tree traits: biomass partitioning, crown morphology and leaf physiology. The study was carried out on one- and three-year-old individuals of five tropical tree species assigned to pioneer (P) or non-pioneer (NP) functional groups. Among the species, Cedrela odorata, Luehea seemannii and Hura crepitans showed the greatest biomass accumulation. On our seasonally dry site, species performance during the first year was dependent on a greater investment in above-ground foraging, while performance after three years was mainly related to water relations. However, large biomass accumulations were not simply associated with an efficient water use but also with contrasting water uses, based on inter-specific relationships. Generally, greater carbon isotope discrimination (Δ_leaf) was related to greater allocation to roots. Species with high Δ_leaf generally showed high leaf potential nitrogen use efficiency (PNUE), suggesting that lower water use efficiency (WUE) increases the efficiency of photosynthetically active N. Also, PNUE was negatively correlated to leaf mass per area (LMA), implying that photosynthetically active N is diluted as total leaf mass increases. Finally, no distinction in measured traits, including biomass accumulation, was observed between the two functional groups.     

Predicting leaf traits of herbaceous species from their spectral characteristics

Trait predictions from leaf spectral properties are mainly applied to tree species, while herbaceous systems received little attention in this topic. Whether similar trait–spectrum relations can be derived for herbaceous plants that differ strongly in growing strategy and environmental constraints is therefore unknown. We used partial least squares regression to relate key traits to leaf spectra (reflectance, transmittance, and absorbance) for 35 herbaceous species, sampled from a wide range of environmental conditions. Specific Leaf Area and nutrient‐related traits (N and P content) were poorly predicted from any spectrum, although N prediction improved when expressed on a per area basis (mg/m² leaf surface) instead of mass basis (mg/g dry matter). Leaf dry matter content was moderately to good correlated with spectra. We explain our results by the range of environmental constraints encountered by herbaceous species; both N and P limitations as well as a range of light and water availabilities occurred. This weakened the relation between the measured response traits and the leaf constituents that are truly responsible for leaf spectral behavior. Indeed, N predictions improve considering solely upper or under canopy species. Therefore, trait predictions in herbaceous systems should focus on traits relating to dry matter content and the true, underlying drivers of spectral properties.     

Leaf trait variation captures climate differences but differs with species irrespective of functional group

Aims To clarify whether variation in leaf traits with climate differs with scale, i.e. across species and within a species, and to detect whether plant functional group affects species-specific response.Methods Leaf dry matter content (LDMC), specific leaf area (SLA), mass- and area-based leaf N (N mass, N area) and leaf P concentrations (P mass, P area) and leaf chlorophyll concentration (SPAD) were measured for 92 woody plant species in two botanical gardens in China. The two gardens share plant species in common but differ in climate. Leaf trait variation between the two gardens was examined via mean comparison at three scales: all species together, species grouped into plant functional groups and within a species. A meta-analysis was performed to summarize the species-specific responses.Important findings At the scale of all species together, LDMC, SLA, P mass and N mass were significantly lower in the dry-cold habitat than in the wet-warm one, whereas N area and SPAD showed an inverse pattern, indicating a significant environmental effect. The meta-analysis showed that the above-mentioned patterns persisted for SLA, N area and SPAD but not for the other variables at the species-specific scale, indicating that intraspecific variation affects the overall pattern of LDMC, P mass and N mass and P area. In terms of species-specific response, positive, negative or nonsignificant patterns were observed among the 92 species. Contrary to our prediction, species-specific responses within a functional group were not statistically more similar than those among functional groups. Our results indicated that leaf trait variation captured climatic difference yet species-specific responses were quite diverse irrespective of plant functional group, providing new insights for interpreting trait variability with climate.     

Genetic effects on the biomass partitioning and growth of Pisum and Lycopersicon

We examined a series of eight pea genotypes differing in three naturally occurring allelic mutations, i.e., af (afila), st (stipules reduced), and tl (tendril-less) and three species, five cultivars, and one interspecific hybrid of tomato differing in SP (SELF-PRUNING) allele composition to determine whether different phenotypes ontogenetically express different biomass partitioning patterns compared to the isometric partitioning pattern and an interspecific 3/4 scaling "rule" governing annual growth with respect to body mass. The slopes and "elevations" (i.e., α and log β, respectively) of log-log linear regression curves of bivariate plots of leaf, stem, and root dry mass and of annual growth vs. total body mass were used to assess pattern homogeneity. The annual growth of all pea and tomato phenotypes complied with the 3/4 growth rule. The biomass partitioning patterns of all tomato phenotypes were statistically indistinguishable from the isometric pattern as were those of the pea wild type and three single-mutant genotypes. However, significant departures from the isometric (and pea wild type) biomass allocation pattern were observed for three genotypes, all of which were homozygous for the af allele. These results open the door to explore the heritability and genetics underlying the allometry of biomass partitioning patterns and growth.     

Leaf structural and photosynthetic characteristics,and biomass allocation to foliage in relation to foliar nitrogen content and tree size in three Betula species

Young trees 0.03-1.7 m high of three coexisting Betula species were investigated in four sites of varying soil fertility, but all in full daylight, to separate nutrient and plant size controls on leaf dry mass per unit area (MA), light-saturated foliar photosynthetic electron transport rate (J) and the fraction of plant biomass in foliage (F(L)). Because the site effect was generally non-significant in the analyses of variance with foliar nitrogen content per unit dry mass (N(M)) as a covariate, N(M) was used as an explaining variable of leaf structural and physiological characteristics. Average leaf area (S) and dry mass per leaf scaled positively with N(M) and total tree height (H) in all species. Leaf dry mass per unit area also increased with increasing H, but decreased with increasing N(M), whereas the effects were species-specific. Increases in plant size led to a lower and increases in N(M) to a greater FL and total plant foliar area per unit plant biomass (LAR). Thus, the self-shading probably increased with increasing N(M) and decreased with increasing H. Nevertheless, the whole-plant average M(A), as well as M(A) values of topmost fully exposed leaves, correlated with N(M) and H in a similar manner, indicating that scaling of MA with N(M) and H did not necessarily result from the modified degree of within-plant shading. The rate of photosynthetic electron transport per unit dry mass (J(M)) scaled positively with N(M), but decreased with increasing H and M(A). Thus, increases in M(A) with tree height and decreasing nitrogen content not only resulted in a lower plant foliar area (LAR = F(L)/M(A)), but also led to lower physiological activity of unit foliar biomass. The leaf parameters (J(M), N(M) and M(A)) varied threefold, but the whole-plant characteristic FL varied 20-fold and LAR 30-fold, indicating that the biomass allocation was more plastically adjusted to different plant internal nitrogen contents and to tree height than the foliar variables. Our results demonstrate that: (1) tree height and N(M) may independently control foliar structure and physiology, and have an even greater impact on biomass allocation; and (2) the modified within-plant light availabilities alone do not explain the observed patterns. Although there were interspecific differences with respect to the statistical significance of the relationships, all species generally fit common regressions. However, these differences were consistent, and suggested that more competitive species with inherently larger growth rates also more plastically respond to N and H.     

Temporal patterns of growth and nutrient accumulation of plant species in a Mediterranean mountainous grassland

The temporal patterns of growth and nutrient accumulation into shoots of coexisting species were studied at two neighbouring areas contrasting with respect to long-term water availability, in an upland herbaceous grassland. Plant growth limiting nutrients were nitrogen (N) in the wet area, and N and phosphorus (P) in the dry area. A series of seven harvests allowed assessment of temporal patterns of peaks for dry matter, N, P and potassium (K) accumulation into shoots and their respective rates of accumulation. Additionally, “pairwise species’ proportion similarities” and “pairwise species’ overlaps” were estimated from shoot biomass data. The peaks of N and P accumulation preceded those of K and dry matter accumulation. Similar trends were evident for the respective rates of accumulation. Compared to the dry area, in the wet area there was a prolonged growing period that increased the inter-species temporal variability in the rates of nutrient accumulation. Abundance of species was correlated (negatively) to species N and P concentrations only in the dry area. It is argued that the significant negative correlation between abundance and either N or P concentrations in the dry area was indicative of intensified inter-species competition resulting from declined temporal complementarity.     

降水变化和种间竞争对红松和蒙古栎幼苗生长的影响

针对全球变暖导致的降水格局的变化,选择长白山红松针阔叶混交林主要树种蒙古栎和红松幼苗为研究对象,在野外自然条件下人工模拟增、减水30%对单种和混种的针阔叶树种形态、生长和生物量分配的影响. 结果表明：对于蒙古栎幼苗,与单种处理相比,混种显著增加了其冠幅和主根长,与对照处理相比,减水处理显著提高了其茎质比、降低了其主根长; 对于红松幼苗,与单种处理相比,混种显著减少了其基径、树高、叶片数和根、茎、叶质量及总干质量,与对照处理相比,减水处理显著降低了其主根长、叶片数、叶质量和总干质量以及叶质比,同时显著提高了茎质比. 增水处理对二者的影响均不显著. 在树木生长初期,种间竞争和降水格局变化对蒙古栎和红松幼苗形态和生长均产生显著影响,对红松幼苗的影响更大.     

The Altitudinal Patterns of Leaf C∶N∶P Stoichiometry Are Regulated by Plant Growth Form,Climate and Soil on Changbai Mountain,China

Understanding the geographic patterns and potential drivers of leaf stoichiometry is critical for modelling the nutrient fluxes of ecosystems and to predict the responses of ecosystems to global changes. This study aimed to explore the altitudinal patterns and potential drivers of leaf C∶N∶P stoichiometry. We measured the concentrations of leaf C, N and P in 175 plant species as well as soil nutrient concentrations along an altitudinal transect (500–2300 m) on the northern slope of Changbai Mountain, China to explore the response of leaf C∶N∶P stoichiometry to plant growth form (PGF), climate and soil. Leaf C, N, P and C∶N∶P ratios showed significant altitudinal trends. In general, leaf C and C∶N∶P ratios increased while leaf N and P decreased with elevation. Woody and herbaceous species showed different responses to altitudinal gradients. Trees had the largest variation in leaf C, C∶N and C∶P ratios, while herbs showed the largest variation in leaf N, P and N∶P ratio. PGF, climate and soil jointly regulated leaf stoichiometry, explaining 17.6% to 52.1% of the variation in the six leaf stoichiometric traits. PGF was more important in explaining leaf stoichiometry variation than soil and climate. Our findings will help to elucidate the altitudinal patterns of leaf stoichiometry and to model ecosystem nutrient cycling.     

N, P, and C stoichiometry of Eranthis hyemalis (Ranunculaceae) and the allometry of plant growth

We report the nitrogen (N), phosphorus (P), and carbon (C) stoichiometry for each of the five organ-types (leaves, aerial stems, reproductive organs, roots, and tubers) of 17 actively growing Eranthis hyemalis plants differing in size (as measured in g C). We also report the N, P, and C stoichiometry of 20 winterized tubers, which are the only perennial organs of this species. Comparisons between whole-plant and winterized N/C and P/C levels indicate that N was resorbed from aerial organs and stored in tubers by the end of the growing season. Leaves were substantial reservoirs for N and P. With few exceptions, N scaled isometrically with respect to C for each organ-type, whereas P scaled as the 3/4 power of C. Thus, N is proportional to P(3/4), which is proportional to C regardless of organ-type. Additionally, annual growth rate G of shoots (leaves and aerial stems) scaled as the -3 power of leaf N/P quotients such that G was proportional to the 3/4 power of leaf P. We suggest that these scaling relationships (together with previously reported allometric trends across herbaceous species) show that growth is constrained by organ-specific N and P allocation patterns (presumably to proteins and ribosomes, respectively).     

峨眉冷杉幼苗叶片功能特征及其N、P化学计量比对模拟大气氮沉降的响应

通过人工施氮模拟大气氮沉降,研究了施氮对峨眉冷杉(Abies fabiri)幼苗叶片功能特征、氮和磷含量及其化学计量比的影响,以及幼苗对氮素的积累效应。结果表明:经过2个生长季节的施氮处理(2009年和2010年,N2)幼苗的总生物量、叶干重、叶重比、叶片氮和磷含量及其N:P分别高于对照处理11.29%、46.70%、41.40%、37.30%、22.33%和6.43%,而比叶面积则降低了6.61%,其中叶干重、叶重比和叶片氮含量与对照处理差异显著(P<0.05),N:P差异极显著(P<0.01),并且叶干重与叶片氮含量具有较强的线性相关;与经1个生长季节施氮处理(2009年,N1)的幼苗总生物量、叶干重、叶重比、比叶面积、叶片氮和磷含量及其N:P的比较分析表明,除叶重比和比叶面积外,其他指标N2均高于N1;人工施氮显著促进了幼苗叶片的生长,提高了叶片氮、磷含量及其N:P,但也反映幼苗生长仍受氮素限制,同时,峨眉冷杉幼苗具有氮素积累效应。     

Leaf size modifies support biomass distribution among stems, petioles and mid-ribs in temperate plants

The implications of extensive variation in leaf size for biomass distribution between physiological and support tissues and for overall leaf physiological activity are poorly understood. Here, we tested the hypotheses that increases in leaf size result in enhanced whole-plant support investments, especially in compound-leaved species, and that accumulation of support tissues reduces average leaf nitrogen (N) content per unit dry mass (N(M)), a proxy for photosynthetic capacity. Leaf biomass partitioning among the lamina, mid-rib and petiole, and whole-plant investments in leaf support (within-leaf and stem) were studied in 33 simple-leaved and 11 compound-leaved species. Support investments in mid-ribs and petioles increased with leaf size similarly in simple leaves and leaflets of compound leaves, but the overall support mass fraction within leaves was larger in compound-leaved species as a result of prominent rachises. Within-leaf and within-plant support mass investments were negatively correlated. Therefore, the total plant support fraction was independent of leaf size and lamina dissection. Because of the lower N(M) of support biomass, the difference in N(M) between the entire leaf and the photosynthetic lamina increased with leaf size. We conclude that whole-plant support costs are weakly size-dependent, but accumulation of support structures within the leaf decreases whole-leaf average N(M), potentially reducing the integrated photosynthetic activity of larger leaves.     

Leaf size versus leaf numbertrade-offs in dioecious angiosperms

Aims We explore the possible role of leaf size/number trade-offs for the interpretation of leaf size dimorphism in dioecious plant species.Methods Total above-ground biomass (both male and female) for three herbaceous dioecious species and individual shoots (from both male and female plants) for three woody dioecious species were sampled to record individual leaf dry mass, number of leaves, dry mass of residual above-ground tissue (all remaining non-leaf biomass), number of flowers/inflorescences (for herbaceous species) and number of branches.Important findings For two out of three woody species and two out of three herbaceous species examined, male plants produced smaller leaves but with higher leafing intensity—i.e. more leaves per unit of supporting (residual) shoot tissue or plant body mass—compared with females. Male and female plants, however, did not differ in shoot or plant body mass or branching intensity. We interpret these results as possible evidence for a dimorphic leaf deployment strategy that promotes both male and female function, respectively. In male plants, capacity as a pollen donor may be favored by selection for a broadly spaced floral display, hence favoring relatively high leafing intensity because this provides more numerous axillary meristems that can be deployed for flowering, thus requiring a relatively small leaf as a trade-off. In one herbaceous species, higher leafing intensity in males was associated with greater flower production than in females. In contrast, in female plants, selection favors a relatively large leaf, we propose, because this promotes greater capacity for localized photosynthate production, thus supporting the locally high energetic cost of axillary fruit and seed development, which in turn requires a relatively low leafing intensity as a trade-off.     

Deciduous and evergreen trees differ in juvenile biomass allometries because of differences in allocation to root storage

Background and Aims

Biomass partitioning for resource conservation might affect plant allometry, accounting for a substantial amount of unexplained variation in existing plant allometry models. One means of resource conservation is through direct allocation to storage in particular organs. In this study, storage allocation and biomass allometry of deciduous and evergreen tree species from seasonal environments were considered. It was expected that deciduous species would have greater allocation to storage in roots to support leaf regrowth in subsequent growing seasons, and consequently have lower scaling exponents for leaf to root and stem to root partitioning, than evergreen species. It was further expected that changes to root carbohydrate storage and biomass allometry under different soil nutrient supply conditions would be greater for deciduous species than for evergreen species.

Methods

Root carbohydrate storage and organ biomass allometries were compared for juveniles of 20 savanna tree species of different leaf habit (nine evergreen, 11 deciduous) grown in two nutrient treatments for periods of 5 and 20 weeks (total dry mass of individual plants ranged from 0·003 to 258·724 g).

Key Results

Deciduous species had greater root non-structural carbohydrate than evergreen species, and lower scaling exponents for leaf to root and stem to root partitioning than evergreen species. Across species, leaf to stem scaling was positively related, and stem to root scaling was negatively related to root carbohydrate concentration. Under lower nutrient supply, trees displayed increased partitioning to non-structural carbohydrate, and to roots and leaves over stems with increasing plant size, but this change did not differ between leaf habits.

Conclusions

Substantial unexplained variation in biomass allometry of woody species may be related to selection for resource conservation against environmental stresses, such as resource seasonality. Further differences in plant allometry could arise due to selection for different types of biomass allocation in response to different environmental stressors (e.g. fire vs. herbivory).     

Greater than the sum of the parts: how the species composition in different forest strata influence ecosystem function

The mechanisms underpinning forest biodiversity‐ecosystem function relationships remain unresolved. Yet, in heterogeneous forests, ecosystem function of different strata could be associated with traits or evolutionary relationships differently. Here, we integrate phylogenies and traits to evaluate the effects of elevational diversity on above‐ground biomass across forest strata and spatial scales. Community‐weighted means of height and leaf phosphorous concentration and functional diversity in specific leaf area exhibited positive correlations with tree biomass, suggesting that both positive selection effects and complementarity occur. However, high shrub biomass is associated with greater dissimilarity in seed mass and multidimensional trait space, while species richness or phylogenetic diversity is the most important predictor for herbaceous biomass, indicating that species complementarity is especially important for understory function. The strength of diversity‐biomass relationships increases at larger spatial scales. We conclude that strata‐ and scale‐ dependent assessments of community structure and function are needed to fully understand how biodiversity influences ecosystem function.     

Inherent allocation patterns and potential growth rates of herbaceous climbing plants

We tested the hypothesis that herbaceous climbing plants, unlike non-climbing herbs, maximize height growth and leaf area, with minimal expenditure in support structures. The enhanced investment in leaf area was expected to result in high relative growth rates in terms of biomass increment. Four leguminous herbaceous climbers from nutrient-poor sites and four non-leguminous herbaceous climbers from nutrient-rich sites, were compared with non-climbing, self-supporting leguminous and non-leguminous herbaceous species from similar habitats. Plants were grown in hydroponic cultures in controlled environment chambers. All climbers had inherently taller shoots than self-supporting plants when compared at an equal amount of total plant dry weight, due to longer stems per unit of support biomass. In contrast to the hypothesis, the relative growth rates of all climbers were relatively low compared to the range found for self-supporting species. The biomass allocation patterns of the non-leguminous climbers were similar to those of the self-supporting species. Leguminous climbers allocated more biomass to support tissue and less biomass to leaves than non-climbers. As a result, height growth was even more emphasized in leguminous climbers than in non-leguminous climbers. Climbing legumes had high rates of net carbon gain, which partly compensated the lower relative leaf weight. We conclude that leguminous herbaceous climbers maximize height growth by a large investment in support biomass, enabling them to keep a large proportion of their leaves in the better illuminated environment at the top of the vegetation canopy.