Temperature dependence of carbon metabolism in the leaves in sun and shade in a subtropical forest.

Rising temperatures pose a threat to the stability of climate regulation by carbon metabolism in subtropical forests. Although the effects of temperature on leaf carbon metabolism traits in sun-exposed leaves are well understood, there is limited knowledge about its impacts on shade leaves and the implications for ecosystem-climate feedbacks. In this study, we measured temperature response curves of photosynthesis and respiration for 62 woody species in summer (including both evergreen and deciduous species) and 20 evergreen species in winter. The aim was to uncover the temperature dependence of carbon metabolism in both sun and shade leaves in subtropical forests. Our findings reveal that shade had no significant effects on the mean optimum photosynthetic temperatures (TOpt) or temperature range (T90). However, there were decreases observed in mean stomatal conductance, mean area-based photosynthetic rates at TOpt and 25 °C, as well as mean area-based dark respiration rates at 25 °C in both evergreen and deciduous species. Moreover, the respiration-temperature sensitivity (Q10) of sun leaves was higher than that of shade leaves in winter, with the reverse being true in summer. Leaf economics spectrum traits, such as leaf mass per area, and leaf concentration of nitrogen and phosphorus across species, proved to be good predictors of TOpt, T90, mass-based photosynthetic rate at TOpt, and mass-based photosynthetic and respiration rate at 25 °C. However, Q10 was poorly predicted by these leaf economics spectrum traits except for shade leaves in winter. Our results suggest that model estimates of carbon metabolism in multilayered subtropical forest canopies do not necessitate independent parameterization of T90 and TOpt temperature responses in sun and shade leaves. Nevertheless, a deeper understanding and quantification of canopy variations in Q10 responses to temperature are necessary to confirm the generality of temperature-carbon metabolism trait responses and enhance ecosystem model estimates of carbon dynamics under future climate warming.

Prediction of photosynthetic light-response curves using traits of the leaf economics spectrum for 75 woody species: effects of leaf habit and sun-shade dichotomy.

PREMISE: Photosynthetic light-response (PLR) curves for leaves are important components of models related to carbon fixation in forest ecosystems, linking the Mitscherlich equation and Michaelis-Menten equation to traits of the leaf economics spectrum (LES). However, models do not consider changes in leaf habits (i.e., evergreen and deciduous) and within-canopy shading variation in these PLR curves. METHODS: Here, we measured the PLR curves in sun and shade leaves of 44 evergreen and 31 deciduous species to examine the relationships between variables of the Mitscherlich equation and Michaelis-Menten equation, leaf nitrogen (N) and phosphorus (P) content, and leaf mass per area (LMA). RESULTS: Small changes were caused by different leaf habits and shade variations in relationships linking variables of the two equations to leaf N and P content and LMA. Values of the scaling exponents for PLR curve parameters did not differ regardless of canopy position and leaf habit (P > 0.05). The PLR curves in species with different leaf habits (i.e., evergreen and deciduous) at different canopy positions could be predicted using the general allometric relations between leaf traits and PLR parameters in the two equations. For photosynthetic photon flux densities from 0 to 2000 µmol m-2 s-1 , approximately 71% (Mitscherlich equation) and 70% (Michaelis-Menten equation) of the net assimilation rates could be predicted. CONCLUSIONS: These findings indicate that leaf net assimilation rates can be predicted through the large available data for LES traits. Incorporation of values for these traits available in the LES databases into ecosystem models of forest productivity and carbon fixation warrants further investigation.

[Nitrogen and phosphorus contents and resorption efficiency of thirty broadleaved woody plants in Yangjifeng, Jiangxi, China.] / æ±Ÿè¥¿é˜³é™ å³°30ç§é˜”å¶æ ‘å¶ç‰‡æ°®ç£·å«é‡åŠå†å¸æ”¶æ•ˆçŽ‡.

Nutrient resorption is an important strategy of nutrient conservation, which reflecting the ability of plants to conserve and utilize nutrients and adapt to environment. To explore the relationship between nutrient content and nutrient resorption of broadleaved woody species of different life forms (i.e., evergreen vs. deciduous), we sampled 30 broadleaved woody species in subtropical region of China located in Yangjifeng National Nature Reserve, Jiangxi Province. The nitrogen (N) and phosphorus (P) concentrations in green and senescent leaves of each species were measured to calculate nutrient resorption efficiency. Furthermore, we analyzed the relationship of leaf nutrient concentration and resorption efficiency for the different life forms. The results showed that N and P concentrations in green leaves were significantly higher in deciduous trees than those in evergreen trees. The P concentrations of senescent leaves in deciduous woody species was significantly higher than that in evergreen woody species. There was no significant difference of N concentration in senescent leaves between evergreen and deciduous species. Nitrogen resorption efficiency (NRE) and phosphorus resorption efficiency (PRE) of the 30 broadleaved woody species were 49.6% and 50.9%, respectively. There were no significant differences between the NRE and PRE of evergreen and deciduous species. NRE and PRE negatively correlated with N and P concentrations in senescent leaves, respectively. Additionally, evergreen and deciduous species showed similar relationships between nutrient resorption efficiency and nutrient concentration in senescent leaves. The sca-ling exponent of allometric relationship between NRE and PRE was 1.18 across all the species. The nutrient resorption efficiency of all the species were affected by the nutrient status of the senesced leaves. Plants examined in this study generally re-absorbed P from senescing leaves than N.

Application of leaf size and leafing intensity scaling across subtropical trees.

Understanding the scaling between leaf size and leafing intensity (leaf number per stem size) is crucial for comprehending theories about the leaf costs and benefits in the leaf size-twig size spectrum. However, the scaling scope of leaf size versus leafing intensity changes along the twig leaf size variation in different leaf habit species remains elusive. Here, we hypothesize that the numerical value of scaling exponent for leaf mass versus leafing intensity in twig is governed by the minimum leaf mass versus maximum leaf mass (M min versus M max) and constrained to be ≤-1.0. We tested this hypothesis by analyzing the twigs of 123 species datasets compiled in the subtropical mountain forest. The standardized major axis regression (SMA) analyses showed the M min scaled as the 1.19 power of M max and the -α (-1.19) were not statistically different from the exponents of M min versus leafing intensity in whole data. Across leaf habit groups, the M max scaled negatively and isometrically with respect to leafing intensity. The pooled data's scaling exponents ranged from -1.14 to -0.96 for M min and M max versus the leafing intensity based on stem volume (LIV). In the case of M min and M max versus the leafing intensity based on stem mass (LIM), the scaling exponents ranged from -1.24 to -1.04. Our hypothesis successfully predicts that the scaling relationship between leaf mass and leafing intensity is constrained to be ≤-1.0. More importantly, the lower limit to scaling of leaf mass and leafing intensity may be closely correlated with M min versus M max. Besides, constrained by the maximum leaf mass expansion, the broad scope range between leaf size and number may be insensitive to leaf habit groups in subtropical mountain forest.

Convergent nitrogen-phosphorus scaling relationships in different plant organs along an elevational gradient.

A general relationship between the nitrogen (N) and phosphorus (P) content of all plant organs (e.g. leaf, stem, and root) is hypothesized to exist according to whole-plant economics spectrum (PES) theory, but the evidence supporting these expected patterns remains scarce. We measured the N and P content of the leaves, twigs and fine roots of 64 species in three different forest communities along an elevational gradient (evergreen broad-leaved forest, 1319 m a.s.l., coniferous and broad-leaved mixed forest, 1697 m a.s.l., and deciduous forest, 1818 m a.s.l.) in the Wuyishan National Nature Reserve, southeastern China. The scaling relationship between the N and P content and the linear regression relationship between the N:P ratio and N and P content were analysed. The leaf N and P content was significantly higher at the high-elevation site than at the low- or middle-elevation sites (P < 0.001). The N and P content followed a power-law relationship with similar scaling slopes between organs. The N (common slope, 1.13) and P (common slope, 1.03) content isometrically covaried among leaves, twigs and roots. The scaling exponents of the N-P relationship were not significantly different from 1.0 in all organs, with a common slope of 1.08. The scaling constants of N-P decreased significantly (P < 0.05) from the highest value in fine roots (ß = 1.25), followed by leaves (ß = 1.17), to the lowest value in twigs (ß = 0.88). Standardized major axis (SMA) analyses and comparisons of 95 % confidence intervals also showed that the numerical values of the scaling slopes and the scaling constants did not differ regardless of elevation. The N content, but not the P content, accounted for a large proportion of the variation in the N:P ratio in leaves (N:P and N: r 2 = 0.31, F = 33.36, P < 0.001) and fine roots (N:P and N: r 2 = 0.15, F = 10.65, P < 0.05). In contrast, the N:P ratio was significantly related to both the N and P content in the twigs (N:P and N: r 2 = 0.20, F = 17.86, P < 0.001; N:P and P: r 2 = 0.34, F = 35.03, P < 0.001, respectively). Our results indicate that different organs of subtropical woody plants share a similar isometric scaling relationship between their N and P content, providing partial support for the PES hypothesis. Moreover, the effects of the N and P content on the N:P ratio differ between metabolic organs (leaves and fine roots) and structural organs (twigs), elucidating the stoichiometric regulatory mechanism of different organs.

The Leaf Economics Spectrum Constrains Phenotypic Plasticity Across a Light Gradient.

The leaf economics spectrum (LES) characterizes multivariate correlations that confine the global diversity of leaf functional traits onto a single axis of variation. Although LES is well established for traits of sun leaves, it is unclear how well LES characterizes the diversity of traits for shade leaves. Here, we evaluate LES using the sun and shade leaves of 75 woody species sampled at the extremes of a within-canopy light gradient in a subtropical forest. Shading significantly decreased the mean values of LMA and the rates of photosynthesis and dark respiration, but had no discernable effect on nitrogen and phosphorus content. Sun and shade leaves manifested the same relationships among N mass, P mass, A mass, and R mass (i.e., the slopes of log-log scaling relations of LES traits did not differ between sun and shade leaves). However, the difference between the normalization constants of shade and sun leaves was correlated with functional trait plasticity. Although the generality of this finding should be evaluated further using larger datasets comprising more phylogenetically diverse taxa and biomes, these findings support a unified LES across shade as well as sun leaves.

[Effects of the current-year shoot stem configuration on leaf biomass in different canopy heights of woody plants in evergreen broad-leaved forest in Jiangxi Province, China.] / æ±Ÿè¥¿å¸¸ç»¿é˜”å¶æž—æœ¨æœ¬æ¤ç‰©ä¸åŒå† å±‚é«˜åº¦å½“å¹´ç”Ÿå°æžèŒŽæž„åž‹å¯¹å¶ç”Ÿç‰©é‡çš„å½±å“.

To investigate the effects of stem configuration on leaf biomass allocation in different organs of the current-year shoots at different canopy heights, relationships of biomass in different organs (i.e., leaves, stems, and twigs) and stem configuration (i.e., stem diameter, length, width/length, stem volume and stem density) were analyzed using the data of 69 woody species from the Yangjifeng Natural Reserve, Jiangxi Provence. Standardized major axis (SMA) was used to explore the regression between biomass and stem configuration. The results showed that there was no significant difference in leaf biomass, stem biomass, twig biomass, stem diameter, stem length, stem width/length and stem volume of current year shoots from upper and lower canopy heights and life forms (i.e., evergreen and deciduous woody plants). Stem density differed significantly in the current year shoots at different heights for both evergreen and deciduous woody species. There were isometric relationships among leaf, stem and total biomass of shoots in different canopy heights and in different life forms. Leaf biomass scaled allometrically with stem diameter and volume, with the scaling exponents being not different significantly among different canopy heights. With respect to the stem configuration of the twigs, stem length, stem width/length and stem density contributed less than 24% to the leaf biomass variation in the current-year shoots. On the contrary, stem diameter and volume had greater effects on leaf biomass of the current-year shoots than stem length, stem width/length and stem density. Canopy heights did not significantly affect the allometric scaling relationships between the stem configuration and leaf biomass of the current-year shoots.

Stem and leaf growth rates define the leaf size vs. number trade-off.

The trade-off between leaf number and individual leaf size on current-year shoots (twigs) is crucial to light interception and thus net carbon gain. However, a theoretical basis for understanding this trade-off remains elusive. Here, we argue that this trade-off emerges directly from the relationship between annual growth in leaf and stem mass, a hypothesis that predicts that maximum individual leaf size (i.e. leaf mass, M max, or leaf area, A max) will scale negatively and isometrically with leafing intensity (i.e. leaf number per unit stem mass, per unit stem volume or per stem cross-sectional area). We tested this hypothesis by analysing the twigs of 64 species inhabiting three different forest communities along an elevation gradient using standardized major axis (SMA) analyses. Across species, maximum individual leaf size (M max, A max) scaled isometrically with respect to leafing intensity; the scaling constants between maximum leaf size and leafing intensity (based on stem cross-sectional area) differed significantly among the three forests. Therefore, our hypothesis successfully predicts a scaling relationship between maximum individual leaf size and leafing intensity, and provides a general explanation for the leaf size-number trade-off as a consequence of mechanical-hydraulic constraints on stem and leaf growth per year.

Stem Diameter (and Not Length) Limits Twig Leaf Biomass.

The relationship between leaf and stem biomass as well as the relationship between leaf biomass and stem length and diameter are important to our understanding of a broad range of important plant scaling relationship because of their relationship to photosynthesis and thus growth. To understand how twig architecture (i.e., current year leaves, and stem diameter and length) affects stem diameter and length, and leaf number and biomass, we examined the twigs of 64 woody species collected from three forest types along an elevational gradient in the Wuyi Mountains, Jiangxi Province, China. We also compared the scaling relationships we observed with biomass allocation patterns reported at the whole tree level. Our results revealed isometric relationship between leaf and stem biomass on twigs despite differences in forest communities and despite changes in environmental factors along an elevational gradient. Across the 64 species, from twigs to individual trees, leaf biomass scaled approximately as the 2.0-power of stem diameter (but not for stem length or leaf number). These results help to identify a general rule that operates at two different levels of biological organization (twigs and whole trees). The scaling relationship between leaf biomass and stem diameter in twigs is insensitive to differences in species composition, elevation, or forest type. We speculate that this rule emerges because stem diameter serves as a proxy for the amount of resources supplied per unit cross section to developing leaves and for the flow of photosynthates from mature leaves to the rest of the plant body.

Effects of light on the growth and clonal reproduction of Ligularia virgaurea.

Ligularia virgaurea is a perennial herb that is widely distributed in the alpine meadow on the eastern Qinghai-Tibet plateau. We investigated the patterns of growth and reproduction of L. virgaurea under two contrasting levels of light conditions for two continuous growing seasons. Our results showed that the light effects on the maximum relative growth rate, the shoot weight ratio and the root weight ratio differed between the two growing seasons. L. virgaurea reproduced initially through rhizome in the second growing season, rather than sexual reproduction. The proportion of genets with clonal reproduction decreased under shaded conditions. A minimum genet size should be attained for clonal reproduction to begin under the shaded conditions. There was a positive linear relationship between clonal reproduction and genet size. Light level affected the allocation of total biomass to clonal structures, with less allocation under the full natural irradiance than under the shaded conditions. There seemed to be a trade-off between vegetative growth and clonal reproduction under the full natural irradiance, in terms of smaller relative growth rates of genets with clonal reproduction than those without clonal reproduction. L. virgaurea emphasized clonal reproduction under the full natural irradiance, while the plant emphasized vegetative growth under the shaded conditions.

Calculation of Thallium's toxicity coefficient in the evaluation of potential ecological risk index: A case study.

As a common pollutive metal element, Tl is very biotoxic. The potential ecological risk index (RI) proposed by Håkanson is one of the commonest methods for evaluation of ecological risk of a metal in sedimentary environment of a water body. According to the calculation principle proposed by Håkanson, the toxicity coefficient of Tl was calculated, and determined as 10 in this paper. In addition, the environmental risk of Tl in the surface sediment of the Beijing-Hangzhou Grand Canal (Zaozhuang Section) was evaluated by RI method, enrichment factor method, etc. The South-to-North Water Diversion Project which benefits four billion people is the largest inter-basin water transfer project in China. The Zaozhuang Section is a significant water conveyance line of this project. We found that the Tl concentrations were 0.46-0.70 µg g-1 with the mean value of 0.61 µg g-1 and were higher than the local background value. The highest contents of Tl occurred in the middle of Zaozhuang section and Tai'erzhuang District, but the enrichment degree of it was much higher in the entrance of Nansihu Lake. The grain size and Al oxides/hydroxides were main factors which controlled the distribution of Tl. Analysis of enrichment factors indicated that Tl in sediments possessed obvious source of human activities which were mainly from combustion of fossil fuels such as coal and mining of mineral resources. As a whole, however, the research region has low Tl content, so Tl has a small probability of environmental pollution.

Characterization of surface sediments from the Beijing-Hangzhou Grand Canal (Zaozhuang section), China: assessment of beryllium enrichment, biological effect, and mobility.

The South-to-North Water Diversion Project is one of the world's largest water diversion projects, benefiting seven million people in China. The Zaozhuang section of the Beijing-Hangzhou Grand Canal is an important part of this project. This paper investigated the enrichment, biological effect, and mobility of beryllium (Be) in surface sediments of the Zaozhuang section. Results showed that high values were found in Tai'erzhuang District, Zaozhuang city, and the areas near the inlet of the Nansihu Lake, which might have been influenced by local human activities including metallurgy, burning of fossil fuels, and transportation. Four geochemical fractions of Be were obtained: acid-soluble fraction, reducible fraction, oxidizable fraction, and residual fraction. The non-residual fractions (the sum of the first three) accounted for 72.5 ∼ 96.1 % of the total amount of Be. Acid-soluble fraction might be mainly influenced by human activities, with the strongest mobility and bio-availability, accounting for 4.1 ∼ 44.7 % of the total amount, with an average of 20.2 %. Enrichment factor (EF) showed minor to moderate enrichment in some regions; adverse effect index (AEI) also showed that there were high levels of Be in some regions, which might have negative impacts on organisms. Generally, mobility, EF, and AEI of elements are carried out separately. But the results of this study indicated that a comprehensive assessment on the enrichment, mobility, and biological effects of Be caused by human activities is necessary in understanding the environmental risks of Be.