不同光强下生长的两种榕树叶片光合能力与比叶重、氮含量及分配的关系

观测了不同光强下生长的两种榕树幼苗叶片的光合能力、比叶重(LMA)、氮含量及在光合机构中的分配.结果表明,两种榕树幼苗的最大净光合速率(Pmax)均随生长光强的升高而升高,这与LMA、单位面积氮含量(NA)和光合氮利用效率随生长光强的升高而升高有关.除36%光强下外,相同光强下生长的喜光的斜叶榕(Ficus tinctoria)的Pmax均显著高于耐荫的假斜叶榕(Ficus subulata),这与其叶片中氮在羧化组份和生物力能学组份中的分配系数、LMA和NA较高有关.     

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.     

Evolutionarily stable leaf area production in plant populations

Using an analytical model, it was shown that for a given amount of nitrogen in the canopy of a stand (N(T)), there exists an evolutionarily stable leaf area index (ES-LAI), and therefore an evolutionarily stable average leaf nitrogen content (n(ES)(av);n(ES)(av) =N(T)/ES-LAI), at which no individual plant in the stand can increase its photosynthesis by changing its leaf area. It was also shown that this ES-LAI is always greater than the optimal LAI that maximizes photosynthesis per unit N(T) of the stand. This illustrates that the canopy structure that maximizes photosynthesis of a population is not the same as the canopy structure that maximizes photosynthesis of individuals within a population. It was further derived that the ES-LAI at given N(T) increases with the ratio between the light-saturated photosynthesis and the N content per unit leaf area (leaf-PPNUE) and that it decreases with the canopy extinction coefficient for light (K(L)), the light availability and the apparent quantum yield (phi). These hypotheses were tested by comparing calculated ES-LAI and n(ES)(av) values to actual LAIs and leaf N contents measured for stands of a large variety of herbaceous plants. There was a close correspondence between the calculated and measured values. As predicted by the model, plants with high leaf-PPNUEs produced more leaf area per unit nitrogen than those with low leaf-PPNUEs while plants with horizontal leaves, forming stands with higher K(L) values, produced less leaf area than those with more vertically inclined leaves. These results suggest that maximization of individual plant photosynthesis per unit of nitrogen plays an important role in determining leaf area production of plants and the resulting canopy structure of stands of vegetation. They further suggest this optimization to be a mechanism by which leaf traits such as leaf-PPNUE and leaf inclination angle are causally related to structural characteristics of the population, i.e. the leaf area index of the stand.     

Photosynthetic acclimation to light in woody and herbaceous species: a comparison of leaf structure, pigment content and chlorophyll fluorescence characteristics measured in the field

Acclimation of foliage photosynthetic properties occurs with varying time kinetics, but structural, chemical and physiological factors controlling the kinetics of acclimation are poorly understood, especially in field environments. We measured chlorophyll fluorescence characteristics, leaf total carotenoid (Car), chlorophyll (Chl) and nitrogen (N) content and leaf dry mass per area (LMA) along vertical light gradients in natural canopies of the herb species, Inula salicina and Centaurea jacea, and tree species, Populus tremula and Tilia cordata, in the middle of the growing season. Presence of stress was assessed on the basis of night measurements of chlorophyll fluorescence. Our aim was to compare the light acclimation of leaf traits, which respond to light availability at long (LMA and N), medium (Chl a/b ratio, Car/Chl ratio) and short time scales (fluorescence characteristics). We found that light acclimation of nitrogen content per unit leaf area (N(area)), chlorophyll content per unit dry mass (Chl(mass)) and Chl/N ratio were related to modifications in LMA. The maximum PSII quantum yield (F(v) /F(m)) increased with increasing growth irradiance in I. salicina and P. tremula but decreased in T. cordata. Leaf growth irradiance, N content and plant species explained the majority of variability in chlorophyll fluorescence characteristics, up to 90% for steady-state fluorescence yield, while the contribution of leaf total carotenoid content was generally not significant. Chlorophyll fluorescence characteristics did not differ strongly between growth forms, but differed among species within a given growth form. These data highlight that foliage acclimation to light is driven by interactions between traits with varying time kinetics.     

Complex adjustments of photosynthetic potentials and internal diffusion conductance to current and previous light availabilities and leaf age in Mediterranean evergreen species Quercus ilex

Mature non-senescent leaves of evergreen species become gradually shaded as new foliage develops and canopy expands, but the interactive effects of integrated light during leaf formation (Q(int)G), current light (Q(int)C) and leaf age on foliage photosynthetic competence are poorly understood. In Quercus ilex L., we measured the responses of leaf structural and physiological variables to Q(int)C and Q(int)G for four leaf age classes. Leaf aging resulted in increases in leaf dry mass per unit area (M(A)), and leaf dry to fresh mass ratio (D(F)) and decreases in N content per dry mass (N(M)). N content per area (N(A)) was independent of age, indicating that decreases in N(M) reflected dilution of leaf N because of accumulation of dry mass (NA = N(M) M(A)). M(A), D(F) and N(A) scaled positively with irradiance, whereas these age-specific correlations were stronger with leaf growth light than with current leaf light. Area-based maximum ribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco) carboxylase activity (V(cmax)A), capacity for photosynthetic electron transport (J(max)A) and the rate of non-photorespiratory respiration in light (R(d)A) were also positively associated with irradiance. Differently from leaf structural characteristics, for all data pooled, these relationships were stronger with current light with little differences among leaves of different age. Acclimation to current leaf light environment was achieved by light-dependent partitioning of N in rate-limiting proteins. Mass-based physiological activities decreased with increasing leaf age, reflecting dilution of leaf N and a larger fraction of non-photosynthetic N in older leaves. This resulted in age-dependent modification of leaf photosynthetic potentials versus N relationships. Internal diffusion conductance (g(m)) per unit area (g(m)A) increased curvilinearly with increasing irradiance for two youngest leaf age classes and was independent of light for older leaves. In contrast, g(m) per dry mass (g(m)M) was negatively associated with light in current-year leaves. Greater photosynthetic potentials and moderate changes in diffusion conductance resulted in greater internal diffusion limitations of photosynthesis in higher light. Both area- and mass-based g(m) decreased with increasing leaf age. The decrease in diffusion conductance was larger than changes in photosynthetic potentials, leading to larger CO2 drawdown from leaf internal air space to chloroplasts (delta(c)) in older leaves. The increases in diffusion limitations in older leaves and at higher light scaled with age- and light-dependent increases in MA and D(F). Overall, our study demonstrates a large potential of foliage photosynthetic acclimation to changes in leaf light environment, but also highlights enhanced structural diffusion limitations in older leaves that result from leaf structural acclimation to previous rather than to current light environment and accumulation of structural compounds with leaf age.     

Leaf age and season influence the relationships between leaf nitrogen, leaf mass per area and photosynthesis in maple and oak trees

Abstract. Seasonal changes in photosynthesis, leaf nitrogen (N) contents and leaf mass per area (LMA) were observed over three growing seasons in open-grown sun-lit leaves of red maple ( Acer rubrum ), sugar maple ( A. sacchamm ) and northern pin oak ( Quereus ellipsoidalis ) trees in southern Wisconsin. Net photosynthesis and leaf N were highly linearly correlated on both mass and area bases within all species from late spring until leaf senescence in fall. Very early in the growing season leaves had high N concentrations, but low photosynthetic rates per unit leaf N, suggesting that leaves were not fully functionally developed at that time. Leaf N per unit area and LMA had nonparallel seasonal patterns, resulting in differing relationships between leaf N/area and LMA in the "early versus late growing season. As a result of differences in seasonal patterns between leaf N/area and LMA, net photosynthesis/area was higher for a given LMA in the spring than fall, and the overall relationships between these two parameters were poor.     

Do Variations in Local Leaf Irradiance Explain Changes to Leaf Nitrogen within Row Maize Canopies?

Numerous studies have dealt with the relationship between leafnitrogen content and leaf irradiance. However, most of themrefer to dense stands presenting reduced horizontal heterogeneityof foliage distribution. Both gradients of leaf nitrogen andleaf irradiance related to canopy depth are significant undersuch conditions, and modelling radiative exchange using a turbid-mediumanalogy and dividing the canopy into vegetation layers is sufficient.Conversely, row crops such as maize are characterized by stronghorizontal heterogeneity of foliage distribution and the one-dimensional(1D) approach may be unsuitable. We thus modelled the three-dimensional(3D) geometry of maize canopies with varying densities and atdifferent developmental stages using plant digitizing underfield conditions. The nitrogen content per unit area of eachleaf part was obtained subsequently by nitrogen analysis. Wenext calculated radiative exchange using a 3D volume-based approachwithin the canopies in order to estimate local leaf irradianceon a daily integration scale. Vertical gradients in leaf nitrogencontent per unit area observed in dense stands during the vegetativephase corresponded largely to those reported in the literature.We also identified significant gradients in nitrogen contentalong the leaves, which had not before been clearly demonstrated.Our study shows that local light climate during plant developmentplays a major role in leaf nitrogen distribution and remobilization.Moreover, brutal plant thinning involves rapid changes in leafnitrogen partitioning. It is concluded that taking account ofthe 3D heterogeneity of nitrogen and irradiance distributionmay have implications for modelling crop photosynthesis andproduction. Copyright 1999 Annals of Botany Company 3D plant architecture, horizontal gradients in leaf nitrogen, leaf irradiance, leaf nitrogen content per unit area, maize, nitrogen partitioning, nitrogen remobilization, virtual plant, Zea mays L.     

Photosynthetic performance along a light gradient as related to leaf characteristics of a naturally occurring <Emphasis Type="Italic">Cypripedium flavum</Emphasis>

Photosynthesis, leaf structure, nitrogen content and nitrogen allocation in photosynthetic functions of Cypripedium flavum were studied in a naturally varying light regime. Light-saturated leaf net photosynthetic rate (A (max)) was strongly correlated with leaf dry mass per area (LMA), mesophyll conductance (g (m)) and area-based leaf nitrogen content (N(area)), with all variables increasing with increasing irradiance. Such coordinate variation of all these parameters illustrates the plastic response of leaf structure to high light (HL). Leaf N(area) was greater under HL than in low light (LL). The fractions of leaf nitrogen partitioning in carboxylation (P (R)) and bioenergetics (P (B)) were positively related to LMA. In contrast, P (R) and P (B) decreased with increasing mass-based leaf nitrogen content (N(mass)). However, no correlation was found between leaf nitrogen investment in light harvesting (P (L)) and either LMA or N(mass). Like maximum rate of carboxylation (V (cmax)) and electron transport (J (max)), the J (max)/V (cmax) ratio, which was strongly correlated to LMA, also increased significantly with irradiance. Under HL, leaf maximum photosynthetic nitrogen efficiency (ANUE) and intrinsic water use efficiency (WUE) were greater than in LL conditions, despite a small difference in WUE. This suggests that a functional balance in the photosynthetic machinery favors leaf photosynthetic plasticity of C. flavum in response to different light conditions. Given an ample soil nitrogen supply, C. flavum may offset its susceptibility to HL by efficient nitrogen use and higher stomatal and mesophyll conductance against photoinhibition so as to keep leaf photosynthesis positive.     

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.     

Nitrogen in cell walls of sclerophyllous leaves accounts for little of the variation in photosynthetic nitrogen-use efficiency

Photosynthetic rate per unit nitrogen generally declines as leaf mass per unit area (LMA) increases. To determine how much of this decline was associated with allocating a greater proportion of leaf nitrogen into cell wall material, we compared two groups of plants. The first group consisted of two species from each of eight genera, all of which were perennial evergreens growing in the Australian National Botanic Gardens (ANBG). The second group consisted of seven Eucalyptus species growing in a greenhouse. The percentage of leaf biomass in cell walls was independent of variation in LMA within any genus, but varied from 25 to 65% between genera. The nitrogen concentration of cell wall material was 0.4 times leaf nitrogen concentration for all species apart from Eucalyptus , which was 0.6 times leaf nitrogen concentration. Between 10 and 30% of leaf nitrogen was recovered in the cell wall fraction, but this was independent of LMA. No trade-off was observed between nitrogen associated with cell walls and the nitrogen allocated to ribulose 1·5-bisphosphate carboxylase/oxygenase (Rubisco). Variation in photosynthetic rate per unit nitrogen could not be explained by variation in cell wall nitrogen.     

翅果油树叶性特征的动态变化

以山西翼城翅果油树(Elaeagnus mollis)自然保护区的翅果油树为研究对象,利用LI-3000A叶面积测定仪测量其单叶面积,用Li-6400便携式光合作用测定系统测定单位叶面积饱和光合速率(Aaraa),用H2SO4-H2O2消煮法测定叶氮含量,计算叶性参数比叶面积(SLA)、单位叶重量饱和光合速率(Amass)、光合氮利用率(PNUE)和单位重量叶氮含量(Nmass)的值,并研究它们与径级之间的关系.结果表明:随着翅果油树径级的增加,SLA、Nmass、Aarea、Amass和PNUE值先下降后上升,当翅果油树径级为7.5 cm左右时,SLA、Nmass、Aarea、Amass和PNUE值均降到最小值,表明径级为7.5 cm左右的翅果油树其光合能力最弱、光合氮利用率较低且生存压力较大.翅果油树叶性参数是研究翅果油树种群动态变化的有效指标,可为研究翅果油树种群动态提供更为便捷的方法.     

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     

Do we Underestimate the Importance of Leaf Size in Plant Economics? Disproportional Scaling of Support Costs Within the Spectrum of Leaf Physiognomy

BACKGROUND: Broad scaling relationships between leaf size and function do not take into account that leaves of different size may contain different fractions of support in petiole and mid-rib. METHODS: The fractions of leaf biomass in petiole, mid-rib and lamina, and the differences in chemistry and structure among mid-ribs, petioles and laminas were investigated in 122 species of contrasting leaf size, life form and climatic distribution to determine the extent to which differences in support modify whole-lamina and whole-leaf structural and chemical characteristics, and the extent to which size-dependent support investments are affected by plant life form and site climate. KEY RESULTS: For the entire data set, leaf fresh mass varied over five orders of magnitude. The percentage of dry mass in mid-rib increased strongly with lamina size, reaching more than 40 % in the largest laminas. The whole-leaf percentage of mid-rib and petiole increased with leaf size, and the overall support investment was more than 60 % in the largest leaves. Fractional support investments were generally larger in herbaceous than in woody species and tended to be lower in Mediterranean than in cool temperate and tropical plants. Mid-ribs and petioles had lower N and C percentages, and lower dry to fresh mass ratio, but greater density (mass per unit volume) than laminas. N percentage of lamina without mid-rib was up to 40 % higher in the largest leaves than the total-lamina (lamina and mid-rib) N percentage, and up to 60 % higher than whole-leaf N percentage, while lamina density calculated without mid-rib was up to 80 % less than that with the mid-rib. For all leaf compartments, N percentage was negatively associated with density and dry to fresh mass ratio, while C percentage was positively linked to these characteristics, reflecting the overall inverse scaling between structural and physiological characteristics. However, the correlations between N and C percentages and structural characteristics differed among mid-ribs, petioles and laminas, implying that the mass-weighted average leaf N and C percentage, density, and dry to fresh mass ratio can have different functional values depending on the importance of within-leaf support investments. CONCLUSIONS: These data demonstrate that variation in leaf size is associated with major changes in within-leaf support investments and in large modifications in integrated leaf chemical and structural characteristics. These size-dependent alterations can importantly affect general leaf structure vs. function scaling relationships. These data further demonstrate important life-form effects on and climatic differentiation in foliage support costs.     

The vertical distribution of nitrogen and photosynthetic activity at different plant densities in Carex acutiformis

Shoots of the monocotyledonous perennial Carex acutiformis were grown in open (28 shoots m⁻²) and dense stands (280 shoots m⁻²). For fully grown stands the distribution of relative PPFD and leaf nitrogen per unit leaf area over canopy depth was determined. Light response of photosynthesis was measured on leaf segments sampled at various heights in the stands. Relations between parameters of these curves and leaf nitrogen were investigated. Simulations showed that in the open stand daily canopy photosynthesis was not affected by nitrogen redistribution in the canopy. For the dense stand however, a uniform nitrogen distribution would lead to only 73% of the maximum net carbon gain by the stand under optimal nitrogen distribution. The actual canopy photosynthesis was only 7% less than this theoretical maximum; the actual nitrogen distribution of the dense stand clearly tended to the optimal distribution. The vertical pattern of the nitrogen distribution was to a large extent determined by the minimum leaf nitrogen content. The relatively high minimum leaf nitrogen content found for Carex leaves may perhaps be necessary to maintain the physiological function of the basal parts of the leaves.     

Optimal co-allocation of carbon and nitrogen in a forest stand at steady state

Nitrogen (N) is essential for plant production, but N uptake imposes carbon (C) costs through maintenance respiration and fine-root construction, suggesting that an optimal C:N balance can be found. Previous studies have elaborated this optimum under exponential growth; work on closed canopies has focused on foliage only. Here, the optimal co-allocation of C and N to foliage, fine roots and live wood is examined in a closed forest stand. Optimal co-allocation maximizes net primary productivity (NPP) as constrained by stand-level C and N balances and the pipe model. Photosynthesis and maintenance respiration increase with foliar nitrogen concentration ([N]), and stand-level photosynthesis and N uptake saturate at high foliage and fine-root density. Optimal NPP increases almost linearly from low to moderate N availability, saturating at high N. Where N availability is very low or very high, the system resembles a functional balance with a steady foliage [N]; in between, [N] increases with N availability. Carbon allocation to fine roots decreases, allocation to wood increases, and allocation to foliage remains stable with increasing N availability. The predicted relationships between biomass density and foliage [N] are in reasonable agreement with data from coniferous stands across Finland. All predictions agree with our qualitative understanding of N effects on growth.     

干旱和湿润生境中主要优势树种叶片功能性状的比较

以云南南部的沟谷雨林(湿润生境)和干热河谷萨瓦纳(savanna)稀树灌丛(干旱生境)中共31种主要优势木本植物为材料,研究了叶片主要功能性状单位面积叶干重(LMA)、单位干重氮含量(Nmass)和单位干重最大净光合速率(Amass)在2个生境中的差异及各性状间的关系。结果表明:在物种水平上,干旱和湿润生境中各树种的LMA值分别为46.88～178.63和45.35～93.16g·m-2;Amass值分别为48.12～176.65和71.38～265.76nmol·g-1·s-1;Nmass值分别为11.8～36.71和12.98～31.78mg·g-1;在群落水平上,干旱生境中LMA显著高于湿润生境,而Amass显著低于湿润生境,Nmass在2个生境中差异不显著;在群落尺度上,干旱生境中LMA与Nmass和Amass均呈显著负相关;在2生境中Nmass与Amass均呈显著正相关。同一生境下不同植物对环境的适应策略有差异,但各性状间的相关关系具有趋同性;从湿润到干旱生境,植物由快速生长转向以提高资源利用效率为主的适应策略。     

Variation in the components of relative growth rate in 10 Acacia species from contrasting environments

In this study we assessed the inherent relative growth rate (RGR) under controlled environment conditions of 10 contrasting Acacia species from semi-arid and mesic environments. For several of the species, compound pinnate leaves produced early in the seedling stage, were gradually replaced by phyllodes (expanded petioles that form simple lamina). Other species either did not form phyllodes, or only did so to a minor degree by the end of the study. Phyllode production was dominant in the four slow-growing Acacia species from semi-arid environments (A. aneura, A. colei, A. coriacea and A. tetragonophylla), with leaf production being exclusive or dominant in five (A. dealbata, A. implexa, A. mearnsii, A. melanoxylon and A. irrorata) of the six faster-growing species from mesic environments. The exception was A. saligna which was fast growing but did produce phyllodes. From a carbon economy perspective, slow growth in the semi-arid species was not associated with lower net assimilation rates or less plant mass allocated to foliage. Rather, the primary factor associated with their slow growth was a smaller foliage area per unit foliage mass. This was true for comparisons based on the mean over all harvests or at set plant masses. The production of phyllodes by the semi-arid species substantially reduced foliage area per unit foliage mass, as this was lower for phyllodes than leaves in all species. To assess the impact that phyllode production had on ontogenetic changes in RGR, we modelled the situation where only leaves were formed. This analysis showed that changing from leaves to phyllodes substantially reduced the RGR. There was little difference in plant nitrogen concentration or the ratio of foliage nitrogen to plant nitrogen between the species. This resulted in foliage nitrogen productivity (dry mass gain per unit foliage nitrogen and time) being directly proportional to foliage area per unit foliage mass between species. We concluded that a smaller foliage area per unit foliage mass and phyllode production are the primary factors associated with lower RGR in contrasting Acacia species.     

不同功能型植物叶氮含量与光合特性的关系研究

在山西南部的霍山七里峪林场,确定乔木、灌木和草本物种共26个,用Li-3000A叶面积测定仪测量了叶面积的大小、用Li-6400便携式光合作用测定系统测定了叶光饱和速率（A_area）,计算了比叶重（LMA）、单位重量的光饱和光合速率（A_mass）、单位面积叶氮含量(Narea)、单位重量叶氮含量(N_mass)及光合氮利用效率(PNUE),研究了它们之间的不同和相互作用关系。结果表明：不同功能型植物的N_mass、A_area、A_mass、N_area和PNUE差异显著(p<0.05),植物叶片氮含量与植物光合生理特性具有显著相关关系,N_mass与A_area、A_mass和PNUE呈线性显著的正相关(p<0.05);N_area与A_area、A_mass、PNUE之间呈极显著的负相关(p<0.01)。     

Photosynthetic stimulation under long-term CO2 enrichment and fertilization is sustained across a closed Populus canopy profile (EUROFACE)

The long-term response of leaf photosynthesis to rising CO2 concentrations [CO2] depends on biochemical and morphological feedbacks. Additionally, responses to elevated [CO2] might depend on the nutrient availability and the light environment, affecting the net carbon uptake of a forest stand. After 6 yr of exposure to free-air CO2 enrichment (EUROFACE) during two rotation cycles (with fertilization during the second cycle), profiles of light, leaf characteristics and photosynthetic parameters were measured in the closed canopy of a poplar (Populus) short-rotation coppice. Net photosynthetic rate (A(growth)) was 49% higher in poplars grown in elevated [CO2], independently of the canopy position. Jmax significantly increased (15%), whereas leaf carboxylation capacity (Vcmax), leaf nitrogen (N(a)) and chlorophyll (Chl(a)) were unaffected in elevated [CO2]. Leaf mass per unit area (LMA) increased in the upper canopy. Fertilization created more leaves in the top of the crown. These results suggest that the photosynthetic stimulation by elevated [CO2] in a closed-canopy poplar coppice might be sustained in the long term. The absence of any down-regulation, given a sufficient sink capacity and nutrient availability, provides more carbon for growth and storage in this bioenergy plantation.     

Canopy structure and leaf nitrogen distribution in a stand of Lysimachia vulgaris L. as influenced by stand density

Summary A hypothesis that a dense stand should develop a less uniform distribution of leaf nitrogen through the canopy than an open stand to increase total canopy photosynthesis was tested with experimentally established stands of Lysimachia vulgaris L. The effect of stand density on spatial variation of photon flux density, leaf nitrogen and specific leaf weight within the canopy was examined. Stand density had little effect on the value of the light extinction coefficient, but strongly affected the distribution of leaf nitrogen per unit area within a canopy. The open stand had more uniform distribution of leaf nitrogen than the dense stand. However, different light climates between stands explained only part of the variation of leaf nitrogen in the canopy. The specific leaf weight in the canopy increased with increasing relative photon flux density and with decreasing nitrogen concentration.