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The plant economics spectrum describes the trade-off between plant resource acquisition and storage, and sheds light on plant responses to environmental changes. However, the data used to construct the plant economics spectrum comes mainly from seed plants, thereby neglecting vascular non-seed plant lineages such as the ferns. To address this omission, we evaluated whether a fern economics spectrum exists using leaf and root traits of 23 fern species living under three subtropical forest conditions differing in light intensity and nutrient gradients. The fern leaf and root traits were found to be highly correlated and formed a plant economics spectrum. Specific leaf mass and root tissue density were found to be on one side of the spectrum (conservative strategy), whereas photosynthesis rate, specific root area, and specific root length were on the other side of the spectrum (acquisitive strategy). Ferns had higher photosynthesis and respiration rates, and photosynthetic nitrogen-use efficiency under high light conditions and higher specific root area and lower root tissue density in high nutrient environments. However, environmental changes did not significantly affect their resource acquisition strategies. Thus, the plant economics spectrum can be broadened to include ferns, which expands its phylogenetic and ecological implications and utility.

Carbon absorption capability and morphological traits are crucial for plant leaf function performance. Here, we investigated the five bamboos at different elevations in Wuyi Mountain to clarify how the leaf trait responds to the elevational gradient and drives the photosynthetic capacity variations. The Standardized Major Axis Regression (SMA) analyses and the Structural Equation Model (SEM) are applied to identify how the bamboo leaf trait, including the ratio of leaf width to length (W/L), leaf mass per area (LMA), photosynthesis rates (Pn), leaf nitrogen, and phosphorus concentration (Leaf N and Leaf P) response to elevation environment, and the driving mechanism of Pn changes. Across the five bamboo species, our results revealed that leaf P scaled isometrically with respect to W/L, leaf N scaled allometrically as the 0.80-power of leaf P, and leaf N and leaf P scaled allometrically to Pn, with the exponents of 0.58 and 0.73, respectively. Besides, the SEM result showed altitude, morphological trait (W/L and LMA), and chemical trait (leaf N and leaf P) could together explain the 44% variations of Pn, with a standard total effect value of 70.0%, 38.5%, 23.6% to leaf P, leaf N, and W/L, respectively. The five bamboo species along the different elevational share an isometric scaling relationship between their leaf P and W/L, providing partial support for the general rule and operating between morphological and chemical traits. More importantly, the leaf W/L and leaf P as the main trait that affects leaf area and P utilization in growth and thus drives bamboo leaf photosynthetic capacity variations in different elevations.

Evergreen and deciduous species coexist in the subtropical forests in southeastern China. It has been suggested that phosphorus (P) is the main limiting nutrient in subtropical forests, and that evergreen and deciduous species adopt different carbon capture strategies to deal with this limitation. However, these hypotheses have not been examined empirically to a sufficient degree. In order to fill this knowledge gap, we measured leaf photosynthetic and respiration rates, and nutrient traits related to P-, nitrogen (N)- and carbon (C)-use efficiencies and resorption using 75 woody species (44 evergreen and 31 deciduous species) sampled in a subtropical forest. The photosynthetic N-use efficiency (PNUE), respiration rate per unit N and P (Rd,N and Rd,P, respectively) of the deciduous species were all significantly higher than those of evergreen species, but not in the case of photosynthetic P-use efficiency. These results indicate that, for any given leaf P, evergreen species manifest higher carbon-use efficiency (CUE) than deciduous species, a speculation that is empirically confirmed. In addition, no significant differences were observed between deciduous and evergreen species for nitrogen resorption efficiency, phosphorus resorption efficiency or N:P ratios. These results indicate that evergreen species coexist with deciduous species and maintain dominance in P-limited subtropical forests by maintaining CUE. Our results also indicate that it is important to compare the PNUE of deciduous species with evergreen species in other biomes. These observations provide insights into modeling community dynamics in subtropical forests, particularly in light of future climate change.

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.

Climate change could negatively alter plant ecosystems if rising temperatures exceed optimal conditions for obtaining carbon. The acclimation of plants to higher temperatures could mitigate this effect, but the potential of subtropical forests to acclimate still requires elucidation. We used space-for-time substitution to determine the photosynthetic and respiratory-temperature response curves, optimal temperature of photosynthesis (Topt), photosynthetic rate at Topt, temperature sensitivity (Q10), and the rate of respiration at a standard temperature of 25°C (R25) for Pinus taiwanensis at five elevations (1200, 1400, 1600, 1800, and 2000 m) in two seasons (summer and winter) in the Wuyi Mountains in China. The response of photosynthesis in P. taiwanensis leaves to temperature at the five elevations followed parabolic curves, and the response of respiration to temperature increased with temperature. Topt was higher in summer than winter at each elevation and decreased significantly with increasing elevation. Q10 decreased significantly with increasing elevation in summer but not winter. These results showed a strong thermal acclimation of foliar photosynthesis and respiration to current temperatures across elevations and seasons, and that R25 increased significantly with elevation and were higher in winter than summer at each elevation indicating that the global warming can decrease R25. These results strongly suggest that this thermal acclimation will likely occur in the coming decades under climate change, so the increase in respiration rates of P. taiwanensis in response to climatic warming may be smaller than predicted and thus may not increase atmospheric CO2 concentrations.

Sapindus mukorossi is an environmentally friendly plant and renewable energy source whose fruit has been widely used for biomedicine, biodiesel, and biological chemicals due to its richness in saponin and oil contents. Here, we report the first chromosome-scale genome assembly of S. mukorossi (covering ~391 Mb with a scaffold N50 of 24.66 Mb) and characterize its genetic architecture and evolution by resequencing 104 S. mukorossi accessions. Population genetic analyses showed that genetic diversity in the southwestern distribution area was relatively higher than that in the northeastern distribution area. Gene flow events indicated that southwest species may be the donor population for the distribution areas in China. Genome-wide selective sweep analysis showed that a large number of genes are involved in defense responses, growth and development, including SmRPS2, SmRPS4, SmRPS7, SmNAC2, SmNAC23, SmNAC102, SmWRKY6, SmWRKY26, and SmWRKY33. We also identified several candidate genes controlling six agronomic traits by genome-wide association studies, including SmPCBP2, SmbHLH1, SmCSLD1, SmPP2C, SmLRR-RKs, and SmAHP. Our study not only provides a rich genomic resource for further basic research on Sapindaceae woody trees but also identifies several economically significant genes for genomics-enabled improvements in molecular breeding.

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.

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 (β = 1.25), followed by leaves (β = 1.17), to the lowest value in twigs (β = 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: r2 = 0.31, F = 33.36, P < 0.001) and fine roots (N:P and N: r2 = 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: r2 = 0.20, F = 17.86, P < 0.001; N:P and P: r2 = 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., We found 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 numerical values of the scaling slopes and the scaling constants did not differ regardless of elevation. These results indicated the different organs of subtropical woody plants share a similar isometric scaling relationship between their N and P content. 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.

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 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, Mmax, or leaf area, Amax) 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 (Mmax, Amax) 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., 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. We present a model (stem-leaf growth hypothesis, SLGH) to provide a theoretical explanation for the trade-off between the maximum leaf size vs. leafing intensity. We found that the scaling exponents of maximum leaf size vs. the leafing intensity are close to −1.0 and are insensitive to forest types and different elevations. Therefore, our results successfully provide a general explanation for this trade-off as a consequence of mechanical-hydraulic constraints on stem and leaf growth per year.

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.

Background and Aims Empirical studies and allometric partitioning (AP) theory indicate that plant above-ground biomass (MA) scales, on average, one-to-one (isometrically) with below-ground biomass (MR) at the level of individual trees and at the level of entire forest communities. However, the ability of the AP theory to predict the biomass allocation patterns of understorey plants has not been established because most previous empirical tests have focused on canopy tree species or very large shrubs. Methods In order to test the AP theory further, 1586 understorey sub-tropical forest plants from 30 sites in south-east China were harvested and examined. The numerical values of the scaling exponents and normalization constants (i.e. slopes and y-intercepts, respectively) of log–log linear MA vs. MR relationships were determined for all individual plants, for each site, across the entire data set, and for data sorted into a total of 19 sub-sets of forest types and successional stages. Similar comparisons of MA/MR were also made. Key Results The data revealed that the mean MA/MR of understorey plants was 2·44 and 1·57 across all 1586 plants and for all communities, respectively, and MA scaled nearly isometrically with respect to MR, with scaling exponents of 1·01 for all individual plants and 0·99 for all communities. The scaling exponents did not differ significantly among different forest types or successional stages, but the normalization constants did, and were positively correlated with MA/MR and negatively correlated with scaling exponents across all 1586 plants. Conclusions The results support the AP theory’s prediction that MA scales nearly one-to-one with MR (i.e. MA ∝ MR ≈1·0) and that plant biomass partitioning for individual plants and at the community level share a strikingly similar pattern, at least for the understorey plants examined in this study. Furthermore, variation in environmental conditions appears to affect the numerical values of normalization constants, but not the scaling exponents of the MA vs. MR relationship. This feature of the results suggests that plant size is the primary driver of the MA vs. MR biomass allocation pattern for understorey plants in sub-tropical forests.

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.

 Premise of the study: Empirical studies and theory indicate that respiration rates ( R ) of small plants scale nearly isometrically with both leaf biomass ( M L ) and total plant biomass ( M T ). These predictions are based on angiosperm species and apply only across a small range of body mass. Whether these relationships hold true for different plants, such as conifers, remains unclear .  Methods: We tested these predictions using the whole-plant maintenance respiration rates and the biomass allocation patterns of the seedlings of two conifer tree species and two angiosperm tree species. Model Type II regression protocols were used to compare the scaling exponents ( α ) and normalization constants ( β ) across all four species and within each of the four species.  Key results: The data show that the scaling exponents varied among the four species and that all differed signifi cantly from isometry. For conifers, scaling exponents for R vs. M T , and R and M L were numerically smaller than those of the broadleaved angiosperm species. However, across the entire data set, R scaled isometrically with M L and with M T as predicted by the West, Brown, and Enquist (WBE) theory. We also observed higher respiration rates for small conifer seedlings compared to compa- rably sized angiosperm seedlings.  Conclusions: Our data add credence to the view that the R vs. M scaling relationship differs among species, and that in general, the numerical values of this interspecifi c scaling relationship will depend on the species pooled in the analysis and on the range of body sizes within the data set.