The influence of seasonality in estimating mangrove leaf chlorophyll-a content from hyperspectral data

Mangrove photosynthetic activity and, consequently, physiological stress can be monitored indirectly using leaf chlorophyll-a (Chla) measurements. Recent studies have demonstrated the feasibility of mangrove leaf Chla content estimation from in situ hyperspectral vegetation indices (VI) but no such research has been conducted using data collected from contrasting seasons (i.e. dry and rainy). In this study, mangrove leaves were collected in a sub-tropical forest of the Mexican Pacific for Chla content determination and in situ hyperspectral measurements (450–1,000 nm). Specifically, we tested 35 VI to estimate Chla content based on a leaf sample of 360 collected from the same trees during both the dry and rainy seasons. The forest examined contained three species of mangrove (Rhizophora mangle, Avicennia germinans and Laguncularia racemosa) exhibiting various conditions of health (dwarf condition, tall and healthy). A principal component analysis, followed by linear regression analyses, were conducted in order to identify those VI that best predict mangrove leaf Chla content during the two seasons. The results indicate that VI derived from hyperspectral measurements collected during the dry season are better at estimating leaf Chla content than those collected during the rainy season. Among the 35 VI, the Vog1 (R₇₄₀/R₇₂₀) index was found to be the best predictor of mangrove leaf Chla content, resulting in R ² values of 0.80 and 0.68 for the dry and rainy season respectively. These results would suggest that for identifying variation in mangrove forest stress (i.e. health) in sub-tropical regions, hyperspectral measurements should be carried out during the dry season.     

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.     

Seasonal variations in leaf δ<Superscript>13</Superscript>C and nitrogen associated with foliage turnover and carbon gain for a wet subalpine fir forest in the Gongga Mountains,eastern Tibetan Plateau

It is still unclear to what extent variations in foliar δ¹³C and nitrogen can be used to detect seasonal changes in canopy productivity. We hypothesize that in a wet and cloudy fir forest, seasonally higher litterfall and lower leaf area index (LAI) are correlated with higher mass-based leaf nitrogen (N _mass) and net primary productivity (NPP), while foliar δ¹³C may change with specific leaf area (SLA), area-based leaf nitrogen (N _area), and/or starch concentration. In order to test our hypotheses, stand-level litterfall and the means of δ¹³C, N _mass, N _area, SLA, and starch concentration of canopy needles for a wet and cloudy Abies fabri forest in the Gongga Mountains were monthly measured during the growing season. Seasonal estimates of LAI were obtained from our previous work. A conceptual model was used to predict seasonal NPP of the fir forest. Seasonal mean δ¹³C and N _mass and climatic variables were used as inputs. The δ¹³C across 1–7-year-old needles increased from May to September associated with decreasing SLA and increasing N _area. There were no significant differences in seasonal starch concentration. With increasing litterfall and decreasing LAI, seasonal mean N _mass increased, while the δ¹³C varied little. The simulated NPP increased with increasing litterfall and related traits of N _mass and N _area. Our data generally supported the hypotheses. The results also suggest that in the forest with relatively moist and cloudy environment, the largest fraction of annual carbon gain may occur in the early part of the growing season when higher litterfall results in higher N _mass of canopy leaves.     

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.     

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 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.     

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     

三种温带树种叶片呼吸的时间动态及其影响因子

为认知叶片呼吸(RL)的季节变化格局及其影响因子,以东北东部山区3个主要树种(红松Pinus koraiensis、樟子松P.sylvestris var.mongolica和白桦Betula platyphylla)为对象,采用红外气体分析法在2011年生长季(常绿树4月至10月;落叶树6月至9月)测定了自然条件下叶片气体交换及其相关生理特征的季节变化,探索了RL与空气温度(Tair)和相关叶片特征之间的关系.结果表明:红松和樟子松基于叶面积的RL(RL-area)表现为生长季初期和末期较大,而白桦RL-area则随生长季进程而逐渐减小.在生长季中,RL-area与叶片总光合之比的时间动态明显.红松、樟子松RL-area与Tair关系显著,而白桦RL-area与Tair关系不显著;但3种树种基于叶质量的RL(RL-mass)与Tair均呈显著的指数函数关系.叶片特征(包括可溶性糖、淀粉、氮、比叶面积等参数)也有明显的季节变化.影响RL的叶片特征参数因树种而异,其中可溶性糖浓度对3种树种的RL均有显著影响.可见,RL的季节变化格局受树木的生长节律、温度和叶片特征的联合控制.     

Leaf chlorophyll content as a proxy for leaf photosynthetic capacity

Improving the accuracy of estimates of forest carbon exchange is a central priority for understanding ecosystem response to increased atmospheric CO₂ levels and improving carbon cycle modelling. However, the spatially continuous parameterization of photosynthetic capacity (Vcmax) at global scales and appropriate temporal intervals within terrestrial biosphere models (TBMs) remains unresolved. This research investigates the use of biochemical parameters for modelling leaf photosynthetic capacity within a deciduous forest. Particular attention is given to the impacts of seasonality on both leaf biophysical variables and physiological processes, and their interdependent relationships. Four deciduous tree species were sampled across three growing seasons (2013–2015), approximately every 10 days for leaf chlorophyll content (Chl_Leaf) and canopy structure. Leaf nitrogen (N_Area) was also measured during 2014. Leaf photosynthesis was measured during 2014–2015 using a Li‐6400 gas‐exchange system, with A‐Ci curves to model Vcmax. Results showed that seasonality and variations between species resulted in weak relationships between Vcmax normalized to 25°C () and N_Area (R² = 0.62, P < 0.001), whereas Chl_Leaf demonstrated a much stronger correlation with (R² = 0.78, P < 0.001). The relationship between Chl_Leaf and N_Area was also weak (R² = 0.47, P < 0.001), possibly due to the dynamic partitioning of nitrogen, between and within photosynthetic and nonphotosynthetic fractions. The spatial and temporal variability of was mapped using Landsat TM/ETM satellite data across the forest site, using physical models to derive Chl_Leaf. TBMs largely treat photosynthetic parameters as either fixed constants or varying according to leaf nitrogen content. This research challenges assumptions that simple N_Area– relationships can reliably be used to constrain photosynthetic capacity in TBMs, even within the same plant functional type. It is suggested that Chl_Leaf provides a more accurate, direct proxy for and is also more easily retrievable from satellite data. These results have important implications for carbon modelling within deciduous ecosystems.     

Leaf habit does not predict leaf functional traits in cerrado woody species

Plant species with a high leaf life span (LLS) commonly have a low specific leaf area (SLA), leaf nitrogen per unit mass (N), and phosphorous concentration (P), whereas species with low LLS have a high SLA, N and P. However, LLS tends to be longer in species growing in low-nutrient soils and, therefore, differences in LLS and other leaf traits may not be consistent with a plant classification according to leaf habit. Here we investigated whether leaf habit is consistent with leaf economic spectrum trade-offs in cerrado (a Neotropical savanna) woody species. We analyzed the SLA, N and P of 125 woody species with a distinct leaf habit (deciduous, semideciduous, brevideciduous or evergreen). We also gathered data on the LLS (33 species), maximum net photosynthesis per leaf area (A_area, 56 species) and per leaf mass (A_mass, 31 species), comprising the most extensive database analyzed so far for the cerrado. Differences among leaf habit groups were tested using generalized linear mixed models and ANOVA. We did not find differences in SLA and N among species with a distinct leaf habit, but deciduous species had a higher leaf P concentration than evergreens. Species did not differ in LLS and A_mass, but A_area varied among groups. Semideciduous species had higher A_area values than deciduous and brevideciduous species, but all other groups had similar A_area values. Because of the small difference in the LLS, SLA, leaf N, leaf P and maximum net photosynthesis, we argue that deciduous, brevideciduous, semideciduous and evergreen species may not constitute different functional groups in cerrado woody species.     

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.     

Quantifying the response of photosynthesis to changes in leaf nitrogen content and leaf mass per area in plants grown under atmospheric CO2 enrichment

Previous modelling exercises and conceptual arguments have predicted that a reduction in biochemical capacity for photosynthesis (A_area) at elevated CO₂ may be compensated by an increase in mesophyll tissue growth if the total amount of photosynthetic machinery per unit leaf area is maintained (i.e. morphological upregulation). The model prediction was based on modelling photosynthesis as a function of leaf N per unit leaf area (N_area), where N_area = N_mass×LMA. Here, N_mass is percentage leaf N and is used to estimate biochemical capacity and LMA is leaf mass per unit leaf area and is an index of leaf morphology. To assess the relative importance of changes in biochemical capacity versus leaf morphology we need to control for multiple correlations that are known, or that are likely to exist between CO₂ concentration, N_area, N_mass, LMA and A_area. Although this is impractical experimentally, we can control for these correlations statistically using systems of linear multiple-regression equations. We developed a linear model to partition the response of A_area to elevated CO₂ into components representing the independent and interactive effects of changes in indexes of biochemical capacity, leaf morphology and CO₂ limitation of photosynthesis. The model was fitted to data from three pine and seven deciduous tree species grown in separate chamber-based field experiments. Photosynthetic enhancement at elevated CO₂ due to morphological upregulation was negligible for most species. The response of A_area in these species was dominated by the reduction in CO₂ limitation occurring at higher CO₂ concentration. However, some species displayed a significant reduction in potential photosynthesis at elevated CO₂ due to an increase in LMA that was independent of any changes in N_area. This morphologically based inhibition of A_area combined additively with a reduction in biochemical capacity to significantly offset the direct enhancement of A_area caused by reduced CO₂ limitation in two species. This offset was 100% for Acer rubrum, resulting in no net effect of elevated CO₂ on A_area for this species, and 44% for Betula pendula. This analysis shows that interactions between biochemical and morphological responses to elevated CO₂ can have important effects on photosynthesis.     

Variation in woody plant δ13C along a topoedaphic gradient in a subtropical savanna parkland

δ¹³C values of C₃ plants are indicators of plant carbon–water relations that integrate plant responses to environmental conditions. However, few studies have quantified spatial variation in plant δ¹³C at the landscape scale. We determined variation in leaf δ¹³C, leaf nitrogen per leaf area (N_area), and specific leaf area (SLA) in April and August 2005 for all individuals of three common woody species within a 308 × 12-m belt transect spanning an upland–lowland topoedaphic gradient in a subtropical savanna in southern Texas. Clay content, available soil moisture, and soil total N were all negatively correlated with elevation. The δ¹³C values of Prosopis glandulosa (deciduous N₂-fixing tree legume), Condalia hookeri (evergreen shrub), and Zanthoxylum fagara (evergreen shrub) leaves increased 1–4‰ with decreasing elevation, with the δ¹³C value of P. glandulosa leaves being 1–3‰ higher than those of the two shrub species. Contrary to theory and results from previous studies, δ¹³C values were highest where soil water was most available, suggesting that some other variable was overriding or interacting with water availability. Leaf N_area was positively correlated with leaf δ¹³C of all species (p < 0.01) and appeared to exert the strongest control over δ¹³C along this topoedaphic gradient. Since leaf N_area is positively related to photosynthetic capacity, plants with high leaf N_area are likely to have low p _I/p _a ratios and therefore higher δ¹³C values, assuming stomatal conductance is constant. Specific leaf area was not correlated significantly with leaf δ¹³C. Following a progressive growing season drought in July/August, leaf δ¹³C decreased. The lower δ¹³C in August may reflect the accumulation of ¹³C-depleted epicuticular leaf wax. We suggest control of leaf δ¹³C along this topoedaphic gradient is mediated by leaf N_area rather than by stomatal conductance limitations associated with water availability.     

Response of plant functional traits to grazing for three dominant species in alpine steppe habitat of the Qinghai–Tibet Plateau,China

Above-ground biomass (AGB) is an important indicator of grassland ecosystem performance. Easily measured plant functional traits (PFTs) may provide useful predictors of the response of plants to grazing. Understanding the response of PFTs to grazing and the relationship between PFTs and AGB is very important for effectively predicting the response of ecosystems to grazing and rangeland management. A grazing experiment was conducted in Gangcha County, Qinghai Province, in the northeastern part of the Qinghai–Tibet Plateau in 2012 and 2013. We investigated the response of PFTs in three dominant species (Elymus nutans, Kobresia humilis, and Stipa purpurea) to grazing, using six stocking rates. Plant height (PH), plant weight, leaf area, and leaf dry biomass of these three dominant species had significantly negative relationships with stocking rate. Leaf thickness (LT) of these three species usually showed a unimodal response to grazing. Specific leaf area generally showed a quadratic relationship with grazing intensity. No consistent effects of grazing were observed on nitrogen content per unit mass (N _mass) and nitrogen content per unit area (N _area). PH, leaf area, and leaf dry mass (LDM) were positively associated with AGB, but LT, N _mass, and N _area had no statistically significant association with AGB. We thus conclude that PH, leaf area, and LDM best predict the effects of grazing on AGB. Finally, 2.87 sheep/ha is recommended as the optimal stocking rate in this region to maintain the health of this grassland ecosystem and to allow for sustainable development.     

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.     

Adaptive characteristics of grassland community structure and leaf traits along an altitudinal gradient on a subtropical mountain in Chongqing,China

Community structure and leaf traits are important elements of terrestrial ecosystems. Changes of community structure and leaf traits are of particular use in the study of the influence of climate change on terrestrial ecosystems. Patterns of community structure (including species richness, above- and below-ground biomass) and leaf traits (including leaf mass per area (LMA), nitrogen content both on mass and area bases (N _mass and N _area), and foliar δ¹³C) from 19 grassland plots along an altitudinal transect at Hongchiba in Chongqing, China, were analyzed. Species richness along the altitudinal transect had a hump-shaped pattern. Above-ground biomass had a quadratic decrease along the altitudinal gradient whereas below-ground biomass had the opposite pattern. Change of above-ground biomass of various taxonomic groups with altitude was also studied. Poaceae showed strong negative relationships and Asteraceae showed a hump-shaped relationship with increase of altitude. Five common species of the grassland, Trifolium pratense, Geranium wilfordii, Aster tataricus, Leontopodium leontopodioides, and Spiraea prunifolia, were particularly studied for variation of leaf traits along the altitudinal gradient. Averaged for all species, LMA, N _area and foliar δ¹³C had positive correlations with altitude. N _mass did not change significantly as altitude increased. LMA and N _area showed significant positive relationships with foliar δ¹³C. The adaptive features of leaf traits among different species were not consistent. The study highlights specific adaptation patterns in relation to altitude for different plant species, provides further insights into adaptive trends of community structure and leaf traits in a specific ecological region filling a gap in the definition of global patterns, and adds to the understanding of how adaptive patterns of plants may respond to global climate change.     

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 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.     

Seasonal changes in sugar beet photosynthesis as affected by exogenous cytokininN 6-(m- hydroxybenzyl)adenosine

Foliar sprays withN ⁶-(m-hydroxybenzytyadenosine, (mOH)-[9R]BAP, one of the synthetic cytokinins, were applied to field-grown sugar beet twice during the vegetation period: before full canopy closing(I), 6 weeks before harvest(II) or both. Application of (mOH)[9R]BAP retained high cytokinin content that usually declines prior to harvest. The total content of isoprenoid cytokinins at harvest was 2.6-fold higher in (mOH)[9R]BAP-treated plants as compared to the controls. TreatmentI had no significant effect on contents of chlorophylls (Chl)a andb and carotenoids, nor on the rates of net photosynthesis (P_N) or photorespiration (R_l), or on CO₂ compensation concentration (Γ) measured during the whole vegetation period on detached leaves under optimum environmental conditions. Increased values in P_N, R_L and Chla andb contents were found in variantsII and I+II linked with a delay in leaf senescence before harvest. Transpiration rate and stomatal conductances of adaxial and abaxial epidermes were not significantly affected by any treatment.     

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。