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.     

How is light interception efficiency related to shoot structure in tall canopy species?

Coexistence of multiple species is a fundamental aspect of plant and forest ecology. Although spatial arrangement of leaves within crowns is an important determinant of light interception and productivity, shoot structure varies considerably among coexisting canopy species. We investigated the relative importance of structural traits in determining the light availability of leaves (I) and light interception efficiency at the current-year shoot level (LIE_CS; the total light interception of leaves divided by shoot biomass) at the top of crowns of 11 canopy species in a cool-temperate forest in Japan. In accordance with Corner’s rules, the total mass, stem mass, total mass of leaf laminae, individual leaf area, and stem cross-sectional area of current-year shoot were positively correlated with each other, and branching intensity (the number of current-year shoots per branch unit of 1-m length) was inversely correlated with these traits across species. In contrast, I was correlated not with these traits, but with leaf elevation angle (a _L). Moreover, variation in LIE_CS across species was caused by variation in I (thus in a _L). Thus, a _L is a key parameter for the leaf light interception of canopy shoots in this cool-temperate forest. Differences in a _L across species might be related to different physiological strategies that developed in the high light and water-limited environment of forest canopies. Small variation in the length of current-year shoots among species implies that variations in I and LIE_CS would be important for the coexistence of these canopy species.     

Changes in crown development patterns and current-year shoot structure with light environment and tree height in Fagus crenata (Fagaceae)

The relative effects of light and tree height on the architecture of leader crowns (i.e., the leading section of the main trunk, 100 cm in length) and current-year shoots for a canopy species, Fagus crenata, occupying both the ridge top and the valley bottom in a cool-temperate forest in Japan were investigated. For leader crowns, the number of current-year shoots and leaves increased with increasing tree height, whereas the mean length of current-year shoots increased with increasing relative photon flux density (PFD). The leader crown area decreased, and the depth and leaf area index of leader crowns increased, with increasing relative PFD. The mass of current-year shoots increased with relative PFD. However, this total mass was allocated differently between stems and leaves depending on tree height, such that the relative allocation to stems increased with increasing tree height. Furthermore, stem structures within current-year shoots also changed with height, such that taller trees produced thicker and shorter stems of the same volume. In contrast, leaf structure and leaf biomass allocations changed with relative PFD. Specific leaf area decreased with increasing relative PFD. In addition, leaf number increased more rapidly with increasing individual leaf mass for trees exposed to greater relative PFD. Consequently, the total leaf area supported by a stem of a given diameter decreased with increasing tree height and relative PFD. Thus, the architecture of leader crowns and current-year shoots were related differently to light and tree height, which are considered important for efficient light capture and the growth of small and tall trees in different environments.     

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.     

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     

Needle-age related variability in nitrogen, mobile carbohydrates, and δ13C within Pinus koraiensis tree crowns

For both ecologists and physiologists, foliar physioecology as a function of spatially and temporally variable environmental factors such as sunlight exposure within a tree crown is important for understanding whole tree physiology and for predicting ecosystem carbon balance and productivity. Hence, we studied concentrations of nitrogen (N), non-structural carbohydrates (NSC = soluble sugars + starch), and δ¹³C in different-aged needles within Pinus koraiensis tree crowns, to understand the needle age- and crown position-related physiology, in order to test the hypothesis that concentrations of N, NSC, and δ¹³C are needle-age and crown position dependent (more light, more photosynthesis affecting N, NSC, and δ¹³C), and to develop an accurate sampling strategy. The present study indicated that the 1-yr-old needles had significantly higher concentration levels of mobile carbohydrates (both on a mass and an area basis) and N_area (on an area basis), as well as NSC-N ratios, but significantly lower levels of N_mass (on a mass basis) concentration and specific leaf area (SLA), compared to the current-year needles. Azimuthal (south-facing vs. north-facing crown side) effects were found to be significant on starch [both on a mass (ST_mass) and an area basis (ST_area)], δ¹³C values, and N_area, with higher levels in needles on the S-facing crown side than the N-facing crown side. Needle N_mass concentrations significantly decreased but needle ST_mass, ST_area, and δ¹³C values significantly increased with increasing vertical crown levels. Our results suggest that the sun-exposed crown position related to photosynthetic activity and water availability affects starch accumulation and carbon isotope discrimination. Needle age associated with physiological activity plays an important role in determining carbon and nitrogen physiology. The present study indicates that across-scale sampling needs to carefully select tissue samples with equal age from a comparable crown position.     

Variation in shoot mortality within crowns of severely defoliated <Emphasis Type="Italic">Betula maximowicziana</Emphasis> trees in Hokkaido,northern Japan

To clarify mortality patterns of current-year shoots within the crown of Betula maximowicziana Regel after severe insect herbivory in central Hokkaido, northern Japan, we investigated the degree of defoliation, pattern of shoot development, shoot mortality, and leaf tissue-water relations. One hundred current-year long shoots growing in a B. maximowicziana plantation were observed for defoliation and mortality in June 2002. An outbreak of herbivorous insects (Caligula japonica and Lymantria dispar praeterea) occurred in the stand in mid-to-late June, and the monitored shoots were defoliated to various degrees. Within 1 month of defoliation, some of the severely defoliated shoots had produced new leaves on short shoots that had emerged from axillary buds. Stepwise logistic regression revealed that the probability that current-year long shoots would put out axillary short shoots with leaves is closely related to the degree of defoliation. To evaluate the water relations of the leaves, we determined pressure–volume curves for the leaves that survived the herbivorous insect outbreak and the new leaves that emerged after defoliation. The water potential at turgor loss (Ψ_l,tlp) and the osmotic potential at full turgidity (Ψ_π,sat) were higher for the new leaves than for the surviving leaves, indicating a lower ability to maintain leaf cell turgor against leaf dehydration in the new leaves. Of the 100 shoots, 13 died after the emergence of new leaves. Stepwise logistic regression revealed that the probability that the long shoots would die generally increased with the emergence of new leaves, with increasing shoot height. This result suggests that the combined effect of the vulnerability of newly emerged leaves and low water availability, associated with higher shoot positions within the crown, caused shoot mortality. Based on our results, some possible mechanisms for mortality in severely defoliated B. maximowicziana are discussed.     

Variation in crown light utilization characteristics among tropical canopy trees

BACKGROUND AND AIMS: Light extinction through crowns of canopy trees determines light availability at lower levels within forests. The goal of this paper is the exploration of foliage distribution and light extinction in crowns of five canopy tree species in relation to their shoot architecture, leaf traits (mean leaf angle, life span, photosynthetic characteristics) and successional status (from pioneers to persistent). METHODS: Light extinction was examined at three hierarchical levels of foliage organization, the whole crown, the outermost canopy and the individual shoots, in a tropical moist forest with direct canopy access with a tower crane. Photon flux density and cumulative leaf area index (LAI) were measured at intervals of 0.25-1 m along multiple vertical transects through three to five mature tree crowns of each species to estimate light extinction coefficients (K). RESULTS: Cecropia longipes, a pioneer species with the shortest leaf life span, had crown LAI <0.5. Among the remaining four species, crown LAI ranged from 2 to 8, and species with orthotropic terminal shoots exhibited lower light extinction coefficients (0.35) than those with plagiotropic shoots (0.53-0.80). Within each type, later successional species exhibited greater maximum LAI and total light extinction. A dense layer of leaves at the outermost crown of a late successional species resulted in an average light extinction of 61% within 0.5 m from the surface. In late successional species, leaf position within individual shoots does not predict the light availability at the individual leaf surface, which may explain their slow decline of photosynthetic capacity with leaf age and weak differentiation of sun and shade leaves. CONCLUSION: Later-successional tree crowns, especially those with orthotropic branches, exhibit lower light extinction coefficients, but greater total LAI and total light extinction, which contribute to their efficient use of light and competitive dominance.     

Nutritional differences and leaf acclimation of climbing plants and the associated vegetation in different types of an Andean montane rainforest

Climbing plants are known to play an important role in tropical forest systems, but key features for their distribution are only partly understood. Investigation was carried out to find if climbers differ from self-supporting vegetation in their adjustment of leaf parameters over a wide variety of light regimes in different forest types along an altitudinal gradient. Relative photon flux density (PFD_rel) was assessed above 75 pairs of strictly linked climbers and supporting vegetation on seven plots between 2,020 and 2,700 m a.s.l. along a mountain range in South-Ecuador up to the Páramo vegetation. Leaf samples from both growth forms were analyzed for leaf area (LA), specific leaf mass (LMA), mass and area-based carbon and nitrogen concentration (C, C_area, N, and N_area) and concentrations of P, K, Ca, Mg, Mn and Al. Leaf size of climbers was independent of general light condition, whereas the leaf size of the self-supporting vegetation increased in shade. LMA increased as expected with altitude and irradiance for both growth forms, but climbers generally built smaller leaves with lower LMA. N, P, and K concentrations were higher in the leaves of climbers than in their supporters. Relationships of LMA and N_area to the light conditions were more pronounced within the climbers than within their supporters. Slope for the regression between climber’s N_area and LMA was twice as steep as for the supporter leaves. Al accumulators were only found within the self-supporting vegetation. The investigated traits indicate improved adjustment towards light supply within climbers compared to self-supporting vegetation. Thus climbing plants seem to have a higher potential trade off in resource-use efficiency regarding irradiance and nutrients.     

Optimal Leaf-to-Root Ratio and Leaf Nitrogen Content Determined by Light and Nitrogen Availabilities

Plants exhibit higher leaf-to-root ratios (L/R) and lower leaf nitrogen content (N _area) in low-light than in high-light environments, but an ecological significance of this trait has not been explained from a whole-plant perspective. This study aimed to theoretically and experimentally demonstrate whether these observed L/R and N _area are explained as optimal biomass allocation that maximize whole-plant relative growth rate (RGR). We developed a model which predicts optimal L/R and N _area in response to nitrogen and light availability. In the model, net assimilation rate (NAR) was determined by light-photosynthesis curve, light availability measured during experiments, and leaf temperature affecting the photosynthesis and leaf dark respiration rate in high and low-light environments. Two pioneer trees, Morus bombycis and Acer buergerianum, were grown in various light and nitrogen availabilities in an experimental garden and used for parameterizing and testing the model predictions. They were grouped into four treatment groups (relative photosynthetic photon flux density, RPPFD 100% or 10%×nitrogen-rich or nitrogen-poor conditions) and grown in an experimental garden for 60 to 100 days. The model predicted that optimal L/R is higher and N _area is lower in low-light than high-light environments when compared in the same soil nitrogen availability. Observed L/R and N _area of the two pioneer trees were close to the predicted optimums. From the model predictions and pot experiments, we conclude that the pioneer trees, M. bombycis and A. buergerianum, regulated L/R and N _area to maximize RGR in response to nitrogen and light availability.     

Temporal and spatial variations in leaf herbivory within a canopy of<Emphasis Type="Italic"> Fagus crenata</Emphasis>

This study investigated spatio-temporal variation in the leaf area consumed by insect herbivores within a canopy of Fagus crenata, with reference to the light conditions of leaf clusters. There was no clear relationship between photosynthetic photon flux density (PPFD) and consumed leaf area (CLA) in May, immediately after leaf flush, but CLA decreased with an increase in PPFD after June. Leaf mass per area, carbon concentration, C/N ratio, concentration of total phenolics, and condensed tannin concentration were higher in leaves under high light intensity than those of leaves under low light. On the other hand, the nitrogen concentration of leaves decreased as light availability increased. Consequently, within-tree variation in light availability affects the consumption of leaves by insect herbivores through temporal changes in leaf characteristics.     

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.     

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     

Herbivory alters plant carbon assimilation,patterns of biomass allocation and nitrogen use efficiency

Herbivory can trigger physiological processes resulting in leaf and whole plant functional changes. The effects of chronic infestation by an insect on leaf traits related to carbon and nitrogen economy in three Prunus avium cultivars were assessed. Leaves from non-infested trees (control) and damaged leaves from infested trees were selected. The insect larvae produce skeletonization of the leaves leaving relatively intact the vein network of the eaten leaves and the abaxial epidermal tissue. At the leaf level, nitrogen content per mass (N_mass) and per area (N_area), net photosynthesis per mass (A_mass) and per area (A_area), photosynthetic nitrogen-use efficiency (PNUE), leaf mass per area (LMA) and total leaf phenols content were measured in the three cultivars. All cultivars responded to herbivory in a similar fashion. The N_mass, A_mass, and PNUE decreased, while LMA and total content of phenols increased in partially damaged leaves. Increases in herbivore pressure resulted in lower leaf size and total leaf area per plant across cultivars. Despite this, stem cumulative growth tended to increase in infected plants suggesting a change in the patterns of biomass allocation and in resources sequestration elicited by herbivory. A larger N investment in defenses instead of photosynthetic structures may explain the lower PNUE and A_mass observed in damaged leaves. Some physiological changes due to herbivory partially compensate for the cost of leaf removal buffering the carbon economy at the whole plant level.     

Seasonal changes in photosynthesis and nitrogen allocation in leaves of different ages in evergreen understory shrub <Emphasis Type="Italic">Daphniphyllum humile</Emphasis>

Seasonal changes in photosynthetic capacity, leaf nitrogen (N) content, leaf chlorophyll (Chl) content and leaf N allocation patterns in leaves of different ages in the evergreen understory shrub, Daphniphyllum humile Maxim, growing at a forest border and an understory site were studied. In current-year leaves at the understory site, the N and Rubisco contents increased from spring to autumn although their light-saturated photosynthetic rate at 22°C (P _max²²) remained stable, indicating that their mesophyll conductance rates declined as they completed their development and/or that they invested increasing amounts of their resources in photosynthetic enzymes during this period. In contrast, seasonal changes in P _max²² in current-year leaves at the forest border site were correlated with changes in Rubisco content. In 1-year old leaves at the understory site, P _max²² and contents of Chl, leaf N, and Rubisco remained stable from spring to autumn, while these parameters decreased in 1-year-old forest border leaves, indicating that N may have been remobilized from shaded 1-year-old leaves to sunlit current-year leaves. When leaves senesced at the forest border site the Rubisco content decreased more rapidly than that of light-harvesting proteins such as LHCII, suggesting that N remobilization from Rubisco may be more efficient, possibly because Rubisco has greater N costs and is soluble, whereas the light-harvesting proteins are membrane components.     

Development of models for estimating leaf chlorophyll and nitrogen contents in tree species with respect to seasonal changes

Models were developed to estimate nondestructively chlorophyll (Chl) content per unit of leaf area (Chl_area) and nitrogen content per unit of leaf area (N_area) using readings of two optical meters for five warm-temperate, evergreen, broadleaved tree species (Castanopsis sieboldii, Cinnamomum tenuifolium, Eurya japonica, Machilus thunbergii, and Neolitsea sericea). It was determined whether models should be adjusted seasonally. Readings (were obtained six times during a year period and Chl_area and N_area were determined using destructive methods. Bayesian inference was used to estimate parameters of models that related optical meter readings to Chl_area or N_area for each species. Deviance information criterion values were used to select the best among models, including the models with seasonal adjustment. The selected models were species-specific and predicted Chl_area accurately (R ² = 0.93–0.96). The best model included parameters with seasonal adjustments for one out of five species. Model-based estimates of N_area were not as accurate as those for Chl_area, but they were still adequate (R ² = 0.64–0.82). For all species studied, the best models did not include parameters with seasonal adjustments. The estimation methods used in this study were rapid and nondestructive; thus, they could be used to assess a function of many leaves and/or repeatedly on individual leaves in the field.     

Photosynthesis-nitrogen relations in Amazonian tree species

The relationships between leaf nitrogen (N), specific leaf area (SLA) (an inverse index of leaf thickness or density), and photosynthetic capacity (A_max) were studied in 23 Amazonian tree species to characterize scaling in these properties among natural populations of leaves of different ages and light microenvironments, and to examine how variation within species in N and SLA can influence the expression of the A_max-to-N relationship on mass versus area bases. The slope of the A_max-N relationship, change in A per change in N (mol CO₂ gN^-1 s^-1), was consistently greater, by as much as 300%, when both measures were expressed on mass rather than area bases. The x-intercept of this relationship (N-compensation point) was generally positive on a mass but not an area basis. In this paper we address the causes and implications of such differences. Significant linear relationships (p<0.05) between mass-based leaf N (N_mass) and SLA were observed in 12 species and all 23 regressions had positive slopes. In 13 species, mass-based A_max (A_mass) was positively related (p<0.05) with SLA. These patterns reflect the concurrent decline in N_mass and SLA with increasing leaf age. Significant (p<0.05) relationships between area-based leaf N (N_area) and SLA were observed in 18 species. In this case, all relationships had negative slopes. Taken collectively, and consistent in all species, as SLA decreased (leaves become thicker) across increasing leaf age and light gradients, N_mass also decreased, but proportionally more slowly, such that N_area increased. Due to the linear dependence of A_mass on N_mass and a negative 4-intercept, thicker leaves (low SLA) therefore tend, on average, to have lower N_mass and A_mass but higher N_area than thinner leaves. This tendency towards decreasing A_mass with increasing N_area, resulting in a lower slope of the A_max-N relationship on an area than mass basis in 16 of 17 species where both were significant. For the sole species exception (higher area than mass-based slope) variation in N_area was related to variation in N_mass and not in SLA, and thus, these data are also consistent with this explanation. The relations between N, SLA and A_max explain how the rate of change in A_max per change in N can vary three-fold depending on whether a mass or area mode of expression is used.     

Contrasting correlation networks between leaf structure,nitrogen and chlorophyll in herbaceous and woody canopies

We studied acclimation patterns in leaf dry mass per area (M_A), nitrogen (N_A) and chlorophyll (ζ_A) content per area, and chlorophyll to nitrogen ratio (ζ/N) along vertical light gradients in natural temperate mixed herbaceous canopy and deciduous tree canopy. In the deciduous tree canopy, all leaves are formed at approximately the same time, and the light gradient during the rest of the growing season reflects the differences in light availability during leaf development, whereas in the herbaceous canopy, leaf production continues during most of the growing season and major changes in light conditions occur after leaf maturation. M_A and N_A increased strongly with increasing current light availability (I_D) in the tree canopy. In the herbaceous canopy, M_A and N_A were generally unrelated to I_D. Depending on species, the correlation between chlorophyll content per leaf area (ζ_A) and I_D was positive, negative, or non-significant. Path analyses revealed two opposite effects of I_D on the amount of leaf chlorophyll. In the tree canopy, increasing I_D enhanced ζ_A through changes in M_A and N_A, whereas the direct effect of light was negative in both canopies. The overall correlation network between foliage structural and chemical traits and the relationships with I_D were significantly stronger in the tree canopy, suggesting limited re-acclimation potential in the mixed herbaceous canopy. Within-species acclimation patterns reflected the patterns within the main functional types. These data demonstrate that the relationships of current light availability vs. leaf dry mass per area, leaf nitrogen and chlorophyll contents, and chlorophyll to nitrogen ratio differ among multi-species herbaceous canopies and deciduous tree canopies due to contrasting canopy development.     

Differences between height- and light-dependent changes in shoot traits in five deciduous tree species

The effects of tree height on shoot traits may in some cases differ in magnitude and direction from the effects of light. Nevertheless, general patterns of change in shoot traits in relation to variations in height and light have not so far been revealed. A comprehensive analysis of the differences between the effects of height and light on a range of leaf and shoot traits is important for the scaling of these traits to individual trees. We investigated the biomass allocation and structure of current-year shoots at the top of the crowns of five deciduous tree species in Japan. Height effect was investigated by comparing shoot traits among trees of different heights growing under a high light environment. The effects of light were examined by comparing saplings growing in high- and low-light environments. The effects of light were significant for most traits, while those of height were not significant for some traits. The magnitudes of the effects of light were larger than those of height for most traits related to biomass allocation. There was an extreme difference between the effects of height and light in the direction of change in the length of current-year shoots and in the number of standing leaves. The measures of both parameters increased with the increase in light, but decreased with the increase in tree height. Thus, the effects of height and light on diverse traits at the level of current-year shoots were not always similar. These results suggest that great care must be taken when scaling shoot traits from small trees to tall trees because the effects of height and light can be complex.     

唐古特白刺叶功能性状沿气候梯度的变异特征

叶功能性状与植物的生长对策及资源利用效率密切相关,研究叶功能性状沿气候梯度的变异特征能为理解植物对气候变化的响应机制提供一种简便可行的测定指标。以我国西北荒漠地区广泛分布的唐古特白刺（Nitraria tangutorum）为研究对象,对其比叶面积（SLA）、单位质量和单位面积叶氮含量（N_mass、N_area）、单位质量和单位面积叶建成成本（CC_mass、CC_area）进行测定,分析这些叶功能性状及性状相关关系沿气候梯度的变异特征。结果表明,唐古特白刺叶功能性状（CC_area除外）在气候梯度下存在显著差异,其中,温度是决定唐古特白刺SLA变化的主要因子,SLA随着温度的增加而增加;降水和温度对唐古特白刺N_mass、N_area和CC_mass均有显著影响,N_mass和N_area随着降水和温度的增加而降低,而CC_mass呈增加趋势。沿气候梯度,唐古特白刺SLA-N_mass、CC_mass-N_mass和CC_area-N_area的线性正相关关系发生平移,导致在相同SLA、CC_mass和CC_area下,降水和温度较低的地区具有更高的N_mass和N_area。这一结果表明唐古特白刺能通过调节叶功能性状之间的关系来适应气候的变化,并形成性状间的最佳功能组合。