“Diminishing returns” and leaf area-biomass scaling of ferns in subtropical ecosystems

Foliage leaves are the primary photosynthetic organ of the majority of vascular plants, and their area vs. biomass scaling relationships provide valuable insights into the capacity and investment in light interception, which is critical to plant growth and performance. The “diminishing returns” hypothesis (DRH), which is based primarily on data from gymnosperms and angiosperms, posits that leaf (lamina) area scales with leaf dry mass. on average with a scaling exponent less than 1.0. However, it remains uncertain whether DRH applies to ferns or whether ecological factors affect the scaling exponents governing fern leaf morphometrics. To address this issue, 182 individuals of 28 subtropical ferns species were studied at low, medium, and high elevations (i.e., 600 m, 900 m, and 1200 m, respectively) in Mount Wuyi National Park, Jiangxi Province, China. The scaling relationships between leaf area and leaf biomass for individual and total leaf of ferns at different elevations were examined by using standardized major axis regression protocols. Analyses of the 28 fern species (using Blomberg K-value protocols) indicated no phylogenetic biases among the species compositions of the three different elevations. In addition, at the individual plant level, individual leaf area (ILA) did not differ significantly among the three different elevations (P > 0.05). However, individual leaf mass (ILM) was significantly higher at 900m than at 1200m (P < 0.05), resulting in a significantly higher leaf mass per area (LMA) at the 900m elevation than at the 600m and 1200m elevations (P < 0.05). The ILA and ILM at the 900m elevation were significantly higher than at the 600m elevation (P < 0.05). At the species level, ILA and ILM did not differ significantly among the three elevations (P > 0.05). The total leaf area per individual (TLA) did not differ significantly across the different elevations (P > 0.05). However, total leaf mass per individual (TLM) did differ significantly (P < 0.05). At the individual plant level, the scaling exponents for ILA vs. ILM and TLA vs. TLM at the three different elevations were all significantly less than 1.0 (P < 0.05), which was consistent with the DRH. At the species level, the scaling exponents for the ILA vs. ILM were significantly less than 1.0 at the middle and high elevations, but not at the low elevation. The scaling exponents of the TLA and TLM were numerically highest in the middle elevation, and all were less than 1.0 for the three elevations. These results indicate that the scaling relationships of leaf area versus mass of subtropical ferns at different elevations support the DRH hypothesis. The study further informs our understanding of the resource allocation strategies of an ancient and diverse plant lineage

Patterns and driving factors of leaf C, N, and P stoichiometry in two forest types with different stand ages in a mid-subtropical zone

Background: Carbon (C), nitrogen (N), and phosphorus (P) stoichiometry is a key indicator of nutrient utilization in plants, and C/N/P ratios are related to the life histories and adaptation strategies of tree species. However, no consensus has been reached on how leaf stoichiometric characteristics are affected by forest type and stand ages. The relationships between leaf stoichiometry and geographical, meteorological, and soil factors also remain poorly understood. Methods: Leaf and soil were sampled from forest stands of different age groups (young, middle-aged, near-mature, and mature) in two forest types (Chinese fir (Cunninghamia lanceolata) forests and evergreen broadleaved forests). The relationships between leaf C, N, and P stoichiometric parameters and geographical, meteorological, and soil factors were analysed by using redundancy analysis (RDA) and stepwise linear regression analysis. Results: Leaf C concentrations peaked in the near-mature stands with increasing age irrespective of forest type. Leaf N and P concentrations fluctuated with a rising trend in Chinese fir forests, while decreased first and increased later from young to mature phases in natural evergreen broadleaved forests. Chinese fir forests were primarily limited by N and P, while natural evergreen broadleaved forests were more susceptible to P limitation. Leaf C, N, and P stoichiometric characteristics in Chinese fir forests were mainly affected by the soil total P concentration (SP), longitude (LNG), growing season precipitation (GSP) and mean temperature in July (JUT). The leaf C concentration was mainly affected by GSP and JUT; leaf N and P concentrations were both positively correlated with LNG; and leaf P was positively correlated with SP. In evergreen broadleaved forests, however, leaf stoichiometric parameters displayed significant correlations with latitude (LAT) and mean annual precipitation (MAP). Conclusions: Leaf stoichiometry differed among forest stands of different age groups and forest types. Leaf C, N, and P stoichiometry was primarily explained by the combinations of SP, LNG, GSP and JUT in Chinese fir forests. LAT and MAP were the main controlling factors affecting the variations in the leaf C, N, and P status in natural evergreen broadleaved forests, which supports the temperature-plant physiological hypothesis. These findings improve the understanding of the distribution patterns and driving mechanisms of leaf stoichiometry linked with stand age and forest type

The morphology and nutrient content drive the leaf carbon capture and economic trait variations in subtropical bamboo forest

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

Leaf structural traits vary with plant size in even-aged stands of Sapindus mukorossi

Sapindus mukorossi Gaertn., 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 of S. 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

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

Scaling relationships of twig biomass allocation in Pinus hwangshanensis along an altitudinal gradient.

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

Short-Term Nitrogen Addition Does Not Significantly Alter the Effects of Seasonal Drought on Leaf Functional Traits in <i>Machilus pauhoi</i> Kanehira Seedlings

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 (&#968;PD), and significant positive effects on the carbon:N (C:N) ratio and specific leaf area (SLA). Short-term N addition (50 kg N&#183;hm&#8722;2&#183;year&#8722;1 and 100 kg N&#183;hm&#8722;2&#183;year&#8722;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 &#968;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&#8722;-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

Coupling Coordination Analysis and Prediction of Landscape Ecological Risks and Ecosystem Services in the Min River Basin

Watershed landscape ecological security and ecosystem service functions are the material basis and environmental guarantee for promoting socioeconomic development. Analyzing the spatiotemporal characteristics of landscape ecological risks (LERs) and ecosystem services (ESs) and exploring the coupling coordination relationship between the two are of great significance for promoting the construction of ecological civilization and achieving sustainable development in the watershed. With the Min River Basin as the study area, the landscape ecological risk assessment, Integrated Valuation of Ecosystem Services and Tradeoffs (InVEST), and Carnegie Ames&ndash;Stanford Approach (CASA) models were used to evaluate the LERs and ESs based on the shared socioeconomic pathways (SSPs), and the patch-generating land use simulation (PLUS) model was used to predict the land use distribution of the Min River Basin in 2030. On this basis, the coupling coordination degree model was used to explore the coupling coordination relationship between the LERs and ESs. The results show that, from 2000 to 2020, the LER of the Min River Basin gradually decreased, and the overall spatial distribution pattern was &ldquo;high in the north and low in the south&rdquo;. The ES of the Min River Basin initially decreased and then increased, showing a spatial distribution pattern of &ldquo;low in the south and high in the north&rdquo;. Among the SSPs in 2030, the LER is the largest under the SSP3 scenario and the smallest under the SSP4 scenario. The ES improvement is the most significant under the SSP1 scenario and the lowest under the SSP3 scenario. From 2000 to 2030, the coupling coordination degree of the Min River Basin first decreased and then increased, showing a spatial distribution pattern of &ldquo;high in the south and low in the north&rdquo;. Among the five SSPs, the coupling coordination degree was the highest under SSP1. The spatial distribution of urban area is the main driving factor affecting the coupling coordination relationship between the LER and ES, and the development of social and economy is the beginning of landscape pattern optimization

Coupling Coordination Analysis and Prediction of Landscape Ecological Risks and Ecosystem Services in the Min River Basin

Watershed landscape ecological security and ecosystem service functions are the material basis and environmental guarantee for promoting socioeconomic development. Analyzing the spatiotemporal characteristics of landscape ecological risks (LERs) and ecosystem services (ESs) and exploring the coupling coordination relationship between the two are of great significance for promoting the construction of ecological civilization and achieving sustainable development in the watershed. With the Min River Basin as the study area, the landscape ecological risk assessment, Integrated Valuation of Ecosystem Services and Tradeoffs (InVEST), and Carnegie Ames–Stanford Approach (CASA) models were used to evaluate the LERs and ESs based on the shared socioeconomic pathways (SSPs), and the patch-generating land use simulation (PLUS) model was used to predict the land use distribution of the Min River Basin in 2030. On this basis, the coupling coordination degree model was used to explore the coupling coordination relationship between the LERs and ESs. The results show that, from 2000 to 2020, the LER of the Min River Basin gradually decreased, and the overall spatial distribution pattern was “high in the north and low in the south”. The ES of the Min River Basin initially decreased and then increased, showing a spatial distribution pattern of “low in the south and high in the north”. Among the SSPs in 2030, the LER is the largest under the SSP3 scenario and the smallest under the SSP4 scenario. The ES improvement is the most significant under the SSP1 scenario and the lowest under the SSP3 scenario. From 2000 to 2030, the coupling coordination degree of the Min River Basin first decreased and then increased, showing a spatial distribution pattern of “high in the south and low in the north”. Among the five SSPs, the coupling coordination degree was the highest under SSP1. The spatial distribution of urban area is the main driving factor affecting the coupling coordination relationship between the LER and ES, and the development of social and economy is the beginning of landscape pattern optimization

Description of <i>Pinus hwangshanensis</i> plots at different altitudes (mean ± <i>SE</i>).

<p>Description of <i>Pinus hwangshanensis</i> plots at different altitudes (mean ± <i>SE</i>).</p