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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.

Alpine treelines are thought to be controlled by low temperature, which affects tree physiology and limits growth. Irrespective of carbon and nutrient limitations are physiological mechanisms affecting the formation of alpine treelines still needs to be defined. We measured the rates of tree growth (basal area increment, BAI), nutrient concentrations in leaves and roots, foliar concentration of nonstructural carbohydrates (NSCs), and gas exchange in Pinus taiwanensis at five elevations (1200, 1400, 1600, 1800, and 2000 m) in the Wuyi Mountains, China. Leaves and roots were sampled twice (summer and winter). The soil nitrogen (N) and phosphorus (P) concentrations and BAI were measured during the summer. We analyzed the foliar traits in summer and winter. The N:P ratio was also analyzed. BAI decreased significantly as elevation increased, accompanied by increases in foliar NSC, N, and P concentrations in both summer and winter. The root P concentration increased with elevation in summer, but the foliar N:P ratio and root N and P concentrations were not affected by elevation in winter. Foliar photosynthesis and respiration did not change in winter, but increased in summer as elevation increased. These results suggest that C and nutrients may not be limiting resources in P. taiwanensis at this alpine treeline site, which instead may be controlled by temperature. P. taiwanensis at alpine treelines accumulates C and nutrient to increase its rates of biochemical reactions at low temperatures.

Sapindus mukorossiGaertn., an important oleaginous woody plant, has garnered increasing research attention owing to its potential as a source of renewable energy (biodiesel). Leaf structural traits are closely related to plant size, and they affect the fruit yield and oil quality. However, plant size factors that predominantly contribute to leaf structural traits remain unknown. Therefore, the purpose of this study was to understand the associations between leaf structural traits and plant size factors in even-aged stands ofS. mukorossi. Results showed that leaf length (LL) and leaf area (LA) markedly increased with the increasing diameter at breast height (DBH) and tree height (TH), although other leaf structural traits did not show noticeable changes. Difference in slopes also indicated that the degree of effect of plant size factors on leaf structural traits was in the order of TH > DBH. Leaf structural traits showed no systematic variation with crown width (CW). LA was significantly positively correlated with LL, leaf width (LW), LL/LW, and leaf thickness (LT) and was significantly but negatively correlated with leaf tissue density (LTD) and leaf dry mass content (LDMC). Specific leaf area showed a significantly negative correlation with LT, LDMC, and LTD. LTD showed a significantly positive correlation with LDMC, but a negative correlation with LT. The results were critical to understand the variability of leaf structural traits with plant size, can provide a theoretical foundation for further study in the relationship between leaf structural traits and fruit yield, and regulate leaf traits through artificial management measures to promote plant growth and fruit yield.

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 Nmass, Pmass, Amass, and Rmass (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.

Premise of study Leaf area and dry mass are crucial for plant metabolic performance. The "diminishing returns" hypothesis predicts that leaf area will scale less than one with respect to leaf dry mass, indicating that the cost of light interception increases with leaf area. However, it remains unclear whether and how this scaling relationship varies among species growing in different environments. Methods More than 2000 measurements from five bamboo species adapted to high and low light and growing at different elevations in Wuyi Mountains, Southeast China, were used to explore how the leaf area vs. dry mass scaling relationship was affected by light and elevation. Key results The data indicate that (1) the normalization constants for leaf area vs. dry mass were positively but not significantly correlated with increasing leaf size and that (2) the scaling exponents remained numerically invariant among all five bamboo species, with a common slope of 0.85. Standardized major axis (SMA) analyses and comparisons of 95% confidence intervals also showed that the numerical values of the scaling exponents did not differ regardless of elevation and were similar between shaded and unshaded adapted species, whereas the numerical values of the normalization constants increased with decreasing light. Conclusions The data collected for all five bamboo species are consistent with the "diminishing returns" hypothesis, i.e., the scaling exponents governing the leaf area vs. dry mass scaling relationship are less than one within and across species and are insensitive to light conditions or elevation.

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.

Research Highlights: Short-term nitrogen (N) addition did not significantly alter the effects of seasonal drought on the leaf functional traits in Machilus pauhoi Kanehira seedlings in N-rich subtropical China. Background and Objectives: Seasonal drought and N deposition are major drivers of global environmental change that affect plant growth and ecosystem function in subtropical China. However, no consensus has been reached on the interactive effects of these two drivers. Materials and Methods: We conducted a full-factorial experiment to analyze the single and combined effects of seasonal drought and short-term N addition on chemical, morphological and physiological traits of M. pauhoi seedlings. Results: Seasonal drought (40% of soil field capacity) had significant negative effects on the leaf N concentrations (LNC), phosphorus (P) concentrations (LPC), leaf thickness (LT), net photosynthetic rate (A), transpiration rate (E), stomatal conductance (Gs), and predawn leaf water potential (&psi, PD), and significant positive effects on the carbon:N (C:N) ratio and specific leaf area (SLA). Short-term N addition (50 kg N&middot, hm&minus, 2&middot, year&minus, 1 and 100 kg N&middot, 1) tended to decrease the C:N ratio and enhance leaf nutrient, growth, and photosynthetic performance because of increased LNC, LPC, LT, leaf area (LA), SLA, A, E, and &psi, PD, however, it only had significant effects on LT and Gs. No significant interactive effects on leaf traits were detected. Seasonal drought, short-term N addition, and their interactions had significant effects on soil properties. The soil total C (STC), nitrate N (NO3&minus, N) and soil total N (STN) concentrations were the main factors that affected the leaf traits. Conclusions: Seasonal drought had a stronger effect on M. pauhoi seedling leaf traits than short-term N deposition, indicating that the interaction between seasonal drought and short-term N deposition may have an additive effecton M. pauhoi seedling growth in N-rich subtropical China.

Background and aims: The &ldquo, diminishing returns&rdquo, hypothesis postulates that the scaling exponent governing the lamina area versus lamina mass scaling relationships has, on average, a numerical value less than one. Theoretically, a similar scaling relationship may exist at the twig level. However, this possibility has not been explored empirically. Methods: We tested both hypotheses by measuring the lamina area and mass, petiole mass of individual leaves, and the total foliage area and stem mass of individual current-year shoots (twigs) of 64 woody species growing in three characteristic forest community types:(1) Evergreen broad-leaved, (2) mixed coniferous and broad-leaved, and (3) deciduous. Key results: The results demonstrate that lamina area vs. mass and lamina area vs. petiole mass differ significantly among the three forest types at both the individual leaf and twig levels. Nevertheless, the scaling exponents of lamina area vs. mass were &lt, 1.0 in each of the three community types, as were the corresponding exponents for lamina area vs. petiole mass, both within and across the three community types. Similar trends were observed at the individual twig level. The numerical values of the scaling exponent for lamina area vs. petiole mass and total foliage area vs. stem mass per twig decreased with increased elevation. Conclusions: These data support the &ldquo, hypothesis at both the individual leaf level and at the individual twig level, phenomena that can inform future inquiries into the mechanistic basis of biomass allocation patterns to physiological (leaf) and mechanical (stem) plant organs.

Empirical studies indicate that the exponents governing the scaling of plant respiration rates (R) with respect to biomass (M) numerically vary between three‐fourth for adult plants and 1.0 for seedlings and saplings and are affected by nitrogen (N) and phosphorus (P) content. However, whether the scaling of R with respect to M (or N and P) varies among different phylogenetic groups (e.g., gymnosperms vs. angiosperms) or during the growing and dormant seasons remains unclear. We measured the whole‐plant R and M, and N and P content of the seedlings of four woody species during the growing season (early October) and the dormant season (January). The data show that (i) the scaling exponents of R versus M, R versus N, and R versus P differed significantly among the four species, but (ii), not between the growing and dormant seasons for each of the four species, although (iii) the normalization constants governing the scaling relationships were numerically greater for the growing season compared to the dormant season. In addition, (iv) the scaling exponents of R versus M, R versus N, and R versus P were numerically larger for the two angiosperm species compared to those of the two gymnosperm species, (v) the interspecific scaling exponents for the four species were greater during the growing season than in the dormant season, and (vi), interspecifically, P scaled nearly isometric with N content. Those findings indicate that the metabolic scaling relationships among R, M, N, and P manifest seasonal variation and differ between angiosperm and gymnosperm species, that is, there is no single, canonical scaling exponent for the seedlings of woody species.

Understanding the response of biomass allocation in twigs (the terminal branches of current-year shoots) to environmental change is crucial for elucidating forest ecosystem carbon storage, carbon cycling, and plant life history strategies under a changing climate. On the basis of interspecies investigations of broad-leaved plants, previous studies have demonstrated that plants respond to environmental factors by allocating biomass in an allometric manner between support tissues (i.e., stems) and the leaf biomass of twigs, where the scaling exponent (i.e., slope of a log—log linear relationship, α) is constant, and the scaling constant (i.e., intercept of a log—log linear relationship, log β) varies with respect to environmental factors. However, little is known about whether the isometric scaling exponents of such biomass allocations remain invariant for single species, particularly conifers, at different altitudes and in different growing periods. In this study, we investigated how twig biomass allocation varies with elevation and period among Pinus hwangshanensis Hsia trees growing in the mountains of Southeast China. Specifically, we explored how twig stem mass, needle mass, and needle area varied throughout the growing period (early, mid-, late) and at three elevations in the Wuyi Mountains. Standardized major axis analysis was used to compare the scaling exponents and scaling constants between the biomass allocations of within-twig components. Scaling relationships between these traits differed with growing period and altitude gradient. During the different growing periods, there was an isometric scaling relationship, with a common slope of 1.0 (i.e., α ≈ 1.0), between needle mass and twig mass (the sum of the total needle mass and the stem mass), whereas there were allometric scaling relationships between the stem mass and twig mass and between the needle mass and stem mass of P. hwangshanensis. The scaling constants (log β) for needle mass vs. twig mass and for needle mass vs. stem mass increased progressively across the growing stages, whereas the scaling constants of stem mass vs. twig mass showed the opposite pattern. The scaling exponents (α) of needle area with respect to needle biomass increased significantly with growing period, changing from an allometric relationship (i.e., α < 1.0) during the early growing period to a nearly isometric relationship (i.e., α ≈ 1.0) during the late growing period. This change possibly reflects the functional adaptation of twigs in different growing periods to meet their specific reproductive or survival needs. At different points along the altitudinal gradient, the relationships among needle mass, twig mass, and stem mass were all isometric (i.e., α ≈ 1.0). Moreover, significant differences were found in scaling constants (log β) along the altitudinal gradient, such that species had a smaller stem biomass but a relatively larger needle mass at low altitude. In addition, the scaling exponents remained numerically invariant among all three altitudes, with a common slope of 0.8, suggesting that needle area failed to keep pace with the increasing needle mass at different altitudes. Our results indicated that the twig biomass allocation pattern was significantly influenced by altitude and growing period, which reflects the functional adaptation of twigs to meet their specific survival needs under different climatic conditions.

Metabolic scaling theory (MST) posits that the scaling exponents among plant height H, diameter D, and biomass M will covary across phyletically diverse species. However, the relationships between scaling exponents and normalization constants remain unclear. Therefore, we developed a predictive model for the covariation of H, D and stem volume V scaling relationships and used data from Chinese fir (Cunninghamia lanceolata) in Jiangxi province, China to test it. As predicted by the model and supported by the data, normalization constants are positively correlated with their associated scaling exponents for D vs. V and H vs. V, whereas normalization constants are negatively correlated with the scaling exponents of H vs. D. The prediction model also yielded reliable estimations of V (mean absolute percentage error = 10.5 ± 0.32 SE across 12 model calibrated sites). These results (1) support a totally new covariation scaling model, (2) indicate that differences in stem volume scaling relationships at the intra-specific level are driven by anatomical or ecophysiological responses to site quality and/or management practices and (3) provide an accurate non-destructive method for predicting Chinese fir stem volume.