Leaves of Lianas and Self-Supporting Plants Differ in Mass per Unit Area and in Nitrogen Content

Abstract: The aim of this study was to test the hypothesis that the reduction in supporting tissues in climbers compared to self-supporting plants is also true for their leaves, and that climbers generally require higher leaf nitrogen than self-supporting plants to accomplish fast growth. This hypothesis was tested using paired samples of both growth forms with assessment of leaf area index above the sampled plants (LAI_a) in a tropical rain forest in Gabon. The sampling protocol ensured that within a highly fluctuating low canopy environment, the growth conditions were identical for each pair sampled. The results confirmed the hypothesis. Lianas had significantly lower leaf mass per unit leaf area (LMA) than their supporters. Liana leaves also contained significantly more nitrogen than host tree leaves. The differences in nitrogen concentration between liana and tree leaves reversed for the most shaded sites, when nitrogen was expressed on a leaf area base (N_area). Significant regression between leaf nitrogen and LAI_a was found for the climbers on the shaded sites but not for their supporters. This indicated better acclimation of climbers to prevailing light conditions. Better nitrogen allocation at low LMA, together with lower carbon costs for building supporting tissues, makes lianas highly competitive, especially where high nitrogen availability is assured.     

Plastic changes of leaf mass per area and leaf nitrogen content in response to canopy openings in saplings of eight deciduous broad-leaved tree species

Leaf nitrogen content per area (N_area) is a good indicator of assimilative capacity of leaves of deciduous broad-leaved trees. This study examined the degrees of increase in N_area in response to canopy openings as leaf mass per area (LMA) and leaf nitrogen content per mass (N_mass) in saplings of eight deciduous broad-leaved tree species in Hokkaido, northern Japan. Five of the species were well-branched species with a large number of small leaves (lateral-growth type), and the other three species were less-branched species with a small number of large leaves (vertical-growth type). The degrees of increase in N_area were compared between the two crown types. In closed-canopy conditions, leaves of the vertical-growth species tended to have a lower LMA and higher N_mass than those of the lateral-growth species, which resulted in similar N_area for both. LMA increased in canopy openings in the eight species, and the degrees of increase were not largely different between the lateral- and vertical-growth species. On the contrary, N_mass was unchanged in canopy openings in the eight species. As a result, N_area of each species increased in canopy openings in proportion to the increase in LMA, and the degrees of increase in N_area were similar in the lateral- and vertical-growth species. Therefore, this study showed that the degrees of increase in N_area were not correlated with the crown architecture (i.e., the lateral- and vertical-growth types).     

Leaf nitrogen distribution in relation to leaf age and photon flux density in dominant and subordinate plants in dense stands of a dicotyledonous herb

We studied the effects of photon flux density (PFD) and leaf position, a measure of developmental age, on the distribution of nitrogen content per unit leaf area (N _area) in plants of different heights, in dense stands grown at two nitrogen availabilities and in solitary plants of the erect dicotyledonous herb Xanthium canadense. Taller more dominant plants received higher PFD levels and experienced a larger difference in relative PFD between their youngest and oldest leaves than shorter subordinate plants in the stands. Differences in PFD between leaves of solitary plants were assumed to be minimal and differences in leaf traits, found for these plants, could thus be mainly attributed to an effect of leaf position. In the solitary plants, N _area decreased with leaf position while in the plants from the stands it decreased with decreasing relative PFD, indicating both factors to be important in determining the distribution of N _area. Due to the effect of leaf position on N _area, leaves of subordinate plants had a higher N _area than older leaves of dominant plants which were at the same height or slightly higher in the canopy. Consequently, the N _area distribution patterns of individual plants plotted as a function of relative PFD were steeper, and probably closer to the optimal distribution which maximizes photosynthesis, than the average distribution in the stand. Leaves of subordinate plants had a lower mass per unit area (LMA) than those of dominant plants. In the dominant plants, LMA decreased with decreasing relative PFD (and with leaf position) while in the subordinate plants it increased. This surprising result for the subordinate plants can be explained by the fact that, during the course of a growing season, these plants became increasingly shaded and newer leaves were thus formed at progressively lower light availability. This indicates that LMA was strongly determined by the relative PFD at leaf formation and to a lesser extent by the current PFD. Leaf N content per unit mass (N _mass) was strongly determined by leaf position independent of relative PFD. This indicates that N _mass is strongly ontogenetically related to the leaf-aging process while changes in N _area, in response to PFD, were regulated through changes in LMA. Received: 11 May 1997 / Accepted: 9 September 1997     

Branching, nitrogen distribution, and carbon gain in solitary plants of an annual herb, Xanthium canadense

The theory of optimal nitrogen (N) distribution predicts that the carbon gain of plants will be maximised when leaves of higher irradiance have higher N content per area (N _area). Most previous studies have examined optimal N distribution without explicitly considering the branching status of plants. I investigated light environment, N distribution and photosynthetic traits of individual leaves of an herbaceous species, Xanthium canadense. X. canadense was grown solitary under high (HN) and low nutrients (LN). Light availability, leaf mass per unit area and N _area were measured for all leaves within plants. Daily photosynthesis of the plants of actual N distribution was compared with those of optimal and constant N distribution. Branch production was facilitated in HN but not in LN plants. N _area was correlated more with leaf order than with leaf light environment. Although N was more limited and the light environment was less heterogeneous within crowns in LN than in HN plants, leaf N distribution was closer to optimal in the latter. These results suggest that leaf N distribution was not optimised in solitary plants of X. canadense. Because this species often regenerates in a dense stand, leaf N distribution might be selected to maximise carbon gain only in such a stand. Leaf N distribution might thus be constrained by the regeneration strategy of the species.     

Leaf trait co‐variation,response and effect in a chronosequence

Question: Is there any generality in terms of leaf trait correlations and the multiple role of leaf traits (response to and/or effect on) during secondary succession? Location: A secondary successional sere was sampled at four different ages since abandonment from several years to nearly 150 years on the Loess Plateau of northwestern China. Method: Specific leaf area (SLA), leaf mass per area (LMA), leaf nitrogen (N_mass, N_area), leaf phosphorus (P_mass, P_area) and leaf dry matter content (LDMC) were measured for all species recorded in the successional sere. Above‐ground net primary productivity (ANPP) and specific rate of litter mass loss (SRLML) were measured as surrogates for ecosystem properties. Soil total carbon (C) and nitrogen (N) were measured in each stage. Leaf traits were related to ecosystem properties and soil nutrient gradients, respectively. Results: LMA is correlated with N_area and P_area' and negatively with N_mass. Correlation between N_area and P_area was higher than between N_mass and P_mass. At the community level, field age, community hierarchy and their interaction explain 64.4 ‐ 93.5% of the variation in leaf traits. At the species level, field age explains 22.4 ‐ 45.5% of the variation in leaf traits (excl. P_area) while plant functional group has a significant effect only for N_mass. LDMC is correlated with ANPP and negatively with SRLML; P_mass is correlated with SRLML. Conclusions: Mean values of LMA, N_mass and N_area are close to the worldwide means, suggesting that large‐scale climate has a profound effect on leaf mass and leaf nitrogen allocation, while environmental gradients represented by succession have little influence on leaf‐trait values. Correlations between leaf traits, such as LMA‐N_area, LMA‐P_area and LMA‐N_mass shown in previous studies, are confirmed here. Although none of the leaf traits is proved to be both a response trait and an effect trait independent of time scale and community hierarchy, mass‐based leaf N is likely a sensitive response trait to soil C and N gradients. In addition, LDMC can be a marker for ANPP and SRLML, while mass‐based leaf P can be a marker for SRLML.     

Functional correlates of leaf demographic response to gap release in saplings of a shade-tolerant tree,<Emphasis Type="Italic"> Elateriospermum tapos</Emphasis>

For a shade-tolerant SE Asian tropical tree, Elateriospermum tapos (Euphorbiaceae), we studied field-established saplings in gaps and the shaded understory to test the hypothesis that differences in leaf demography and leaf life span under contrasting light regimes should be functionally correlated with architecture, self-shading and nitrogen distribution within the sapling crown. Rates of leaf production and net leaf gain were greater for saplings in gaps than those in the understory. Median leaf life span was approximately 26 months in the gap saplings, while it was estimated to be greater than 38 months in the understory saplings. Consequently, gap saplings had a greater standing leaf number and experienced greater degrees of self-shading than understory saplings. Light availability at individual leaves, estimated by a combination of canopy photos and a three-dimensional architecture model, were negatively correlated with leaf age in gap saplings but not so in understory saplings. Leaf nitrogen content per unit area (N_area) was influenced more by light availability than by leaf age in the gap saplings. In contrast, in understory saplings, N_area was neither correlated with light availability nor with leaf age, and did not decrease significantly before 38 months in leaf age. We conclude that saplings of this shade-tolerant species apparently prolong their leaf life span in the shaded understory through slower rates of leaf production, lower standing number of leaves and lower degrees of self shading than in gap, and that the rate of decline of N_area with leaf age depends on architecture and self-shading regimes that respond to changes in light regimes.     

Photosynthesis, leaf morphology and chemistry of Pinus koraiensis and Quercus mongolica in broadleaved Korean pine mixed forest

Leaf traits and physiology are species-specific and various with canopy position and leaf age. Leaf photosynthesis, morphology and chemistry in the upper and lower canopy positions of Pinus koraiensis Sieb. et Zucc and Quercus mongolica Fisch. ex Turoz in broadleaved Korean pine forest were determined in September 2009. Canopy position did not significantly affect light-saturated photosynthetic rate based on unit area (P _area) and unit dry mass (P _mass), apparent quantum yield (α), light compensation point (LCP), light saturation point (LSP); total nitrogen (N_m), phosphorus (P_m), carbon (C_m), and chlorophyll content (Chl_m) per unit dry mass; leaf dry mass per unit area (LMA) and photosynthetic nitrogen-use efficiency (PNUE) for P. koraiensis current-year needles and Q. mongolica leaves. While in P. koraiensis one-year-old needles, P _area, P _mass, α and LCP in the upper canopy were lower than those in the lower canopy. The needles of P. koraiensis had higher Cm and LMA than leaves of Q. mongolica, but P _mass, Chl_m and PNUE showed opposite trend. There were no differences in P _area, LSP, N_m, and P_m between the two species. Needle age significantly influenced photosynthetic parameters, chemistry and LMA of P. koraiensis needles except LCP, LSP and C_m. In contrast to LMA, P _area, P _mass, N_m, P_m, Chl_m, and PNUE of one-year-old needles were significantly lower than those of current-year needles for P. koraiensis. The negative correlations between LMA and P _mass, N_m, P_m, Chl_m, and positive correlations between P _mass and N_m, P_m, Chl_m were found for P. koraiensis current-year needles and Q. mongolica leaves. Our results indicate that leaf nitrogen and phosphorus contents and nutrient absorption from soil are similar for mature P. koraiensis and Q. mongolica growing in the same environment, while difference in carbon content between P. koraiensis and Q. mongolica may be attributed to inherent growth characteristics.     

荆条叶性状对野外不同光环境的表型可塑性

光照是影响植物生长和分布的重要环境因子。对生长在野外5种不同光环境下(林外、阔叶林林缘、阔叶林林下、针叶林林窗和针叶林林下)的荆条的叶片进行取样研究,通过对光合作用光响应曲线、叶绿素荧光、叶绿素含量、叶片氮磷含量以及叶片形态的测量,来反映荆条对不同光环境的表型可塑性。研究结果表明,荆条叶片对于野外不同的光环境具有很好的适应机制,叶片功能性状受到结构性状的调节。低光下通过高的比叶面积(SLA)、单位质量叶绿素含量、光系统II最大量子产量,低的暗呼吸速率、光饱和点、光补偿点、叶绿素a,b的比值来提高对光能的利用效率,维持生长;高光下则通过与SLA有关的叶片结构的变化对光合作用进行调节。大多数的叶性状只受到日光照总量的影响,SLA的大小与日最高光强有关,可以对不同日变化模式的光照做出迅速的响应,是适应不同光照的敏感指标。尽管光照是不同光环境下影响荆条叶性状的主要环境因子,土壤养分含量同样会对叶性状产生影响,高土壤养分下的高叶长与叶柄长的比值体现了植物对资源获取和支撑结构之间分配的权衡。     

Leaf nitrogen distribution in relation to crown architecture in the tall canopy species,Fagus crenata

The theory of optimal leaf N distribution predicts that the C gain of plants is maximized when the N content per unit area (N _area) scales with light availability, but most previous studies have demonstrated that the N distribution is not proportional to light availability. In tall trees, the leaves are often clustered on twigs (leaf cluster) and not evenly distributed within the crowns. Thus, we hypothesized that the suboptimal N distribution is partly caused by the limited capacity to translocate N between leaf clusters, and consequently, the relationship between light and N _area differs for leaves in different clusters. We investigated the light availability and N content of all individual leaves within several leaf clusters on tall trees of a deciduous canopy species Fagus crenata in Japan. We observed that the within-cluster leaf N distribution patterns differed from the between-cluster patterns and the slopes of the relationships between light and N _area were lower within clusters than between clusters. According to the detailed analysis of the N distribution within leaf clusters, N _area was greater for current-year shoots with greater light availability or a larger total leaf area. The latter pattern was probably caused by the greater sink strength of the current-year shoots with a larger leaf area. These N distribution patterns suggest that leaf clusters are fairly independent with respect to their N use, and the productivity of real F. crenata crowns may be less than optimal.     

Converging patterns of vertical variability in leaf morphology and nitrogen across seven Eucalyptus plantations in Brazil and Hawaii,USA

Key message

Across sites in Brazil and Hawaii, LMA and N _mass were strongly correlated with height and shade index, respectively, which may help simplify canopy function modeling of Eucalyptus plantations.

Abstract

Within tree canopies, leaf mass per area (LMA) and leaf nitrogen per unit area (N _area) commonly increase with height. Previous research has suggested that these patterns occur as a strategy to optimize carbon gain by allocating available resources to upper canopy leaves that are exposed to greater light availability. We tested three hypotheses about the influences of height, shade index (a proxy for light), and stand age on LMA and leaf nitrogen for even-aged Eucalyptus saligna and Eucalyptus grandis × urophylla plantations in Brazil and Hawaii, USA, spanning most of the environmental conditions found across 19.6 million ha of Eucalyptus spp. plantations around the world. Shade index was developed by incorporating canopy depth (inner-crown shading) and a tree height ratio relative to neighbor trees (shading from other trees). Across all sites and ages, leaf height accounted for 45 % of the variation in LMA, whereas shade index accounted for only 6 %. A combination of both factors was slightly better in accounting for LMA variation than height alone. LMA–height relationships among sites were strongest under greater light availability and in older stands. Leaf nitrogen per unit mass (N _mass) consistently decreased with shade index, whereas N _area showed no consistent pattern with height or shade index. These relationships indicate that N _mass is primarily driven by light, while height is the primary driver for LMA. The general relationships between LMA and leaf N _mass across all sites may simplify canopy function modeling of E. saligna and E. grandis × urophylla plantations.     

Photosynthetic nitrogen and water use efficiency of acacia and eucalypt seedlings as afforestation species

The ecophysiological traits of acacia and eucalypt are important in assessing their suitability for afforestation. We measured the gas-exchange rate, the leaf dry mass per area (LMA) and the leaf nitrogen content of two acacia and four eucalypt species. Relative to the eucalypts, the acacias had lower leaf net photosynthetic rate (P _N), lower photosynthetic nitrogen-use efficiency (PNUE), higher water-use efficiency (WUE), higher LMA and higher leaf nitrogen per unit area (N _area). No clear differences were observed within or between genera in the maximum rate of carboxylation (V _cmax) or the maximum rate of electron transport (J _max), although these parameters tended to be higher in eucalypts. PNUE and LMA were negatively correlated. We conclude that acacias with higher LMA do not allocate nitrogen efficiently to photosynthetic system, explaining why their P _N and PNUE were lower than in eucalypts.     

Diaheliotropic leaf movement enhances leaf photosynthetic capacity and photosynthetic light and nitrogen use efficiency via optimising nitrogen partitioning among photosynthetic components in cotton (Gossypium hirsutum L.)

• Phototropic leaf movement of plants is an effective mechanism for adapting to light conditions. Light is the major driver of plant photosynthesis. Leaf N is also an important limiting factor on leaf photosynthetic potential. Cotton (Gossypium hirsutum L.) exhibits diaheliotropic leaf movement. Here, we compared the long‐term photosynthetic acclimation of fixed leaves (restrained) and free leaves (allowed free movement) in cotton.
• The fixed leaves and free leaves were used for determination of PAR, leaf chlorophyll concentration, leaf N content and leaf gas exchange. The measurements were conducted under clear sky conditions at 0, 7, 15 and 30 days after treatment (DAT).
• The results showed that leaf N allocation and partitioning among different components of the photosynthetic apparatus were significantly affected by diaheliotropic leaf movement. Diaheliotropic leaf movement significantly increased light interception per unit leaf area, which in turn affected leaf mass per area (LMA), leaf N content (N_A) and leaf N allocation to photosynthesis (N_P). In addition, cotton leaves optimised leaf N allocation to the photosynthetic apparatus by adjusting leaf mass per area and N_A in response to optimal light interception.
• In the presence of diaheliotropic leaf movement, cotton leaves optimised their structural tissue and photosynthetic characteristics, such as LMA, N_A and leaf N allocation to photosynthesis, so that leaf photosynthetic capacity was maximised to improve the photosynthetic use efficiency of light and N under high light conditions.

     

南亚热带不同演替阶段森林优势树种叶片构建成本与机械抗性的协同关系

为探究不同演替阶段森林优势种叶片资源获取策略的差异以及叶片构建成本与机械抗性的关系,对我国南亚热带不同演替阶段森林14优势种的叶片构建成本、机械抗性、角质层厚度和比叶重等结构性状进行测定。结果表明,与演替早期相比,演替晚期优势种具有更高的单位面积叶片构建成本、叶片撕裂力以及穿透力,但其叶片最大光合速率较低;同时,单位面积叶片构建成本与机械抗性呈显著正相关关系,而叶片角质层厚度、比叶重等结构性状也与叶片构建成本、机械抗性均呈显著正相关。因此,从叶片能量投资策略上反映了南亚热带森林演替进程中叶片构建成本与机械抗性的协同关系。     

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.     

Acclimation to irradiance in seedlings of three tropical rain forest Garcinia species after simulated gap formation

We investigated the acclimation of seedlings of three tropical rain forest sub-canopy Garcinia species (G. xanthochymus, G. cowa, and G. bracteata) after transfer from 4.5 (LI) to 40 % (HI) sunlight and 12.5 (MI) sunlight to HI (LH₁ and LH₂ denoting transfer from LI to HI and MI to HI transfer, respectively). The changes of chlorophyll (Chl) fluorescence, net photosynthetic rate (P _N), dark respiration rate (R _D), Chl content per unit area (Chl_area), leaf mass per unit area (LMA), and seedling mortality were monitored over two months after transfer. These parameters together with leaf anatomy of transferred and control seedlings (kept in LI, MI, and HI) were also examined after two months. No seedlings died during the two months. F_v/F_m, P _N, and Chl_area of the transferred seedlings decreased in the first 3 to 12 d. LH₁ leaves showed larger reduction in F_v/F_m (>23 % vs. <16 %) and slower recovery of F_v/F_m than LH₂ leaves. P _N started to recover after about one week of I transfer and approached higher values in all G. cowa seedlings and G. xanthochymus LH₁ seedlings than those before the transfer. However, P _N of G. bracteata seedlings approached the values before transfer. The final P _N values in leaves of transferred G. xanthochymus and G. cowa seedlings approached that of leaves kept in HI, while the final P _N values of transferred leaves of G. bracteata were significantly lower than that of leaves grown under HI (p<0.05). R _D of G. xanthochymus LH₁ seedlings and all G. cowa seedlings increased and approached the value of the seedlings in HI. The final Chl_area of both G. xanthochymus and G. cowa approached the values before transfer, but that of G. bracteata did not recover to the level before transfer. The final Chl_area of all transferred seedlings was not significantly different from that of seedlings in HI except that G. cowa LH₁ seedlings had higher Chl_area than that in HI. LMA decreased within 2 d and then increased continuously until about 30 d and approached the value under HI. Spongy/palisade mesophyll ratio decreased after transfer because of the increase in palisade thickness. Leaf thickness did not change, so LMA increase of transferred seedlings was mainly due to the increase of leaf density. Thus the mature leaves under LI and MI of G. xanthochymus and G. cowa are able to acclimate to HI by leaf physiological and anatomical adjustment, while G. bracteata had limited ability to acclimate to HI.     

Spatial patterns of photosynthetic characteristics and leaf physical traits of plants in the Loess Plateau of China

The spatial patterns of photosynthetic characteristics and leaf physical traits of 171 plants belonging to nine life-forms or functional groups (trees, shrubs, herbs, evergreen trees, deciduous trees, C₃ and C₄ herbaceous plants, leguminous and non-leguminous species) and their relationships with environmental factors in seven sites, Yangling, Yongshou, Tongchuan, Fuxian, Ansai, Mizhi and Shenmu, ranging from south to north in the Loess Plateau of China were studied. The results showed that the leaf light-saturated photosynthetic rate (P_max), photosynthetic nitrogen use efficiency (PNUE), chlorophyll content (Chl), and leaf mass per area (LMA) of all the plants in the Loess Plateau varied significantly among three life-form groups, i.e., trees, shrubs and herbs, and two groups, i.e., evergreen trees and deciduous trees, but leaf nitrogen content differed little among different life-form groups. For the 171 plants in the Loess Plateau, leaf P_max was positively correlated with PNUE. The leaf nitrogen content per unit area (N_area) was positively correlated but Chl was negatively correlated with the LMA. When controlling the LMA, the N_area was positively correlated with the Chl (partial r = 0.20, P < 0.05). With regard to relationships between photosynthetic characteristics and leaf physical traits, the P_max was positively correlated with N _area, while the PNUE was positively correlated with the Chl and negatively correlated with the N_area and LMA. For all the species in the Loess Plateau, the PNUE was negatively correlated with the latitude and annual solar radiation (ASR), but positively correlated with the mean annual rainfall (MAR) and mean annual temperature (MAT). With regard to the leaf physical traits, the leaf Chl was negatively correlated with the latitude and ASR, but positively correlated with the MAR and MAT. However, the N_area and LMA were positively correlated with the latitude and ASR, but negatively correlated with the MAR and MAT. In general, leaf N_area and LMA increased, while PNUE and Chl decreased with increases in the latitude and ASR and decreases in MAR and MAT. Electronic supplementary material The online version of this article () contains supplementary material, which is available to authorized users.     

Optimal photosynthetic use of light by tropical tree crowns achieved by adjustment of individual leaf angles and nitrogen content

Background and Aims

Theory for optimal allocation of foliar nitrogen (ONA) predicts that both nitrogen concentration and photosynthetic capacity will scale linearly with gradients of insolation within plant canopies. ONA is expected to allow plants to efficiently use both light and nitrogen. However, empirical data generally do not exhibit perfect ONA, and light-use optimization per se is little explored. The aim was to examine to what degree partitioning of nitrogen or light is optimized in the crowns of three tropical canopy tree species.

Methods

Instantaneous photosynthetic photon flux density (PPFD) incident on the adaxial surface of individual leaves was measured along vertical PPFD gradients in tree canopies at a frequency of 0·5 Hz over 9–17 d, and summed to obtain the average daily integral of PPFD for each leaf to characterize its insolation regime. Also measured were leaf N per area (N_area), leaf mass per area (LMA), the cosine of leaf inclination and the parameters of the photosynthetic light response curve [photosynthetic capacity (A_max), dark respiration (R_d), apparent quantum yield (ϕ) and curvature (θ)]. The instantaneous PPFD measurements and light response curves were used to estimate leaf daily photosynthesis (A_daily) for each leaf.

Key Results

Leaf N_area and A_max changed as a hyperbolic asymptotic function of the PPFD regime, not the linear relationship predicted by ONA. Despite this suboptimal nitrogen partitioning among leaves, A_daily did increase linearly with PPFD regime through co-ordinated adjustments in both leaf angle and physiology along canopy gradients in insolation, exhibiting a strong convergence among the three species.

Conclusions

The results suggest that canopy tree leaves in this tropical forest optimize photosynthetic use of PPFD rather than N per se. Tropical tree canopies then can be considered simple ‘big-leaves’ in which all constituent ‘small leaves’ use PPFD with the same photosynthetic efficiency.Key words: Optimal resource allocation, nitrogen, photosynthetic capacity, leaf mass per area, tropical trees, radiation use efficiency, scaling, leaf angle, canopy architecture, big leaf model     

云南文山石漠化区车桑子叶脉密度与叶氮含量关系对生境的响应

植物叶脉特征和叶氮含量的变化影响着叶片经济谱的形成,为验证叶片结构中叶脉网络构建提供了理论依据。该文以单位质量叶氮含量(N_(mass))和单位面积叶氮含量(N_(area))分别表示叶氮含量,采取主成分分析、线性回归分析的方法,研究了云南文山石漠化区旷地(Ⅰ)、林缘(Ⅱ)和林下(Ⅲ)3种自然生境下车桑子的叶脉密度(Vein density,VD)与N_(mass)和N_(area)的异速关系。结果表明:从乔灌群落的旷地到林下,车桑子的比叶面积、叶绿素总含量、光能利用率和N_(mass)逐渐增大,光饱和点、光补偿点、水分利用效率、VD、N_(area)逐渐减小,净光合速率、蒸腾速率、气孔导度呈先增大后减小的趋势。VD与叶氮含量呈不同程度的相关性,在生境I和III,VD与N_(mass)和N_(area)分别具有显著的负相关(P0.05)和正相关(P0.05);在生境Ⅱ,VD与N_(mass)和N_(area)分别呈不显著负相关(P0.05)和正相关(P0.05)。车桑子在旷地强光生境,高VD的叶片含有低N_(mass)高N_(area),而林下荫蔽生境偏向于相反的配置模式,反映了石漠化区植物较强的叶脉可塑性及其与氮利用性状的权衡机制。     

Leaf traits of natural populations of <Emphasis Type="Italic">Adiantum reniforme</Emphasis> var. <Emphasis Type="Italic">sinensis</Emphasis>, endemic to the Three Gorges region in China

Leaf mass per unit area (LMA), carbon and nitrogen contents, leaf construction cost, and photosynthetic capacity (P _max) of Adiantum reniforme var. sinensis, an endangered fern endemic to the Three Gorges region in southwest China, were compared in five populations differing in habitat such as soil moisture and irradiance. The low soil moisture and high irradiance habitat population exhibited significantly higher LMA, area-based leaf construction (CC_A), and carbon content (C_A), but lower leaf nitrogen content per unit dry mass (N_M) than the other habitat populations. The high soil moisture and low irradiance habitat populations had the lowest CC_A, but their cost/benefic ratios of CCA/P _max were similar to the medium soil moisture and irradiance habitat population due to their lower leaf P _max. Hence A. reniforme var. sinensis prefers partially shaded, moist but well-drained, slope habitats. Due to human activities, however, its main habitats now are cliffs or steeply sloped bare rocks with poor and thin soil. The relatively high energy requirements and low photosynthetic capacity in these habitats could limit the capability of the species in extending population or interspecific competition and hence increase its endangerment.     

Phenotypic correlates of the lianescent growth form: a review

Background

As proposed by Darwin, climbers have been assumed to allocate a smaller fraction of biomass to support organs in comparison with self-supporting plants. They have also been hypothesized to possess a set of traits associated with fast growth, resource uptake and high productivity.

Scope

In this review, these hypotheses are evaluated by assembling and synthesizing published and unpublished data sets from across the globe concerning resource allocation, growth rates and traits of leaves, stems and roots of climbers and self-supporting species.

Conclusions

The majority of studies offer little support for the smaller allocation of biomass to stems or greater relative growth rates in climbers; however, these results are based on small sized (<1 kg) plants. Simulations based on allometric biomass equations demonstrate, however, that larger lianas allocate a greater fraction of above-ground biomass to leaves (and therefore less biomass to stems) compared with similar sized trees. A survey of leaf traits of lianas revealed their lower average leaf mass per area (LMA), higher N and P concentration and a slightly higher mass-based photosynthetic rate, as well as a lower concentration of phenolic-based compounds than in woody self-supporting species, consistent with the specialization of lianas towards the fast metabolism/rapid turnover end of the global trait spectra. Liana stems have an efficient hydraulic design and unique mechanical features, while roots appear to penetrate deeper soil levels than in trees and are often able to generate hydraulic pressure. Much remains to be learned, however, about these and other functional specializations of their axial organs and the associated trade-offs. Developmental switches between self-supporting, searcher and climbing shoots within the same individual are a promising field of comparative studies on trait association in lianas. Finally, some of the vast trait variability within lianas may be reduced when species with different climbing mechanisms are considered separately, and when phylogenetic conservatism is accounted for.