“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

Trait divergence and opposite above- and below-ground strategies facilitate moso bamboo invasion into subtropical evergreen broadleaf forest

Understanding the invasion of moso bamboo (Phyllostachys edulis) into adjacent evergreen broadleaf forest based on functional traits is crucial due to its significant influence on ecosystem processes. However, existing research has primarily focused on above- or below-ground traits in isolation, lacking a comprehensive integration of both. In this study, we conducted a trait-based analysis including 23 leaf traits and 11 root traits in three forest types - bamboo forest, mixed bamboo and broadleaf forest, and evergreen broadleaf forest - to investigate trait differences, phenotypic integration, and above- and below-ground resource strategies in bamboo and broadleaf species. Our findings demonstrated significant differences in leaf and root key traits between bamboo and broadleaf species, strongly supporting the "phenotypic divergence hypothesis". Bamboo exhibited stronger trait correlations compared to broadleaf species, indicating higher phenotypic integration. Above- and below-ground strategies were characterized by trade-offs rather than coordination, resulting in a multi-dimensional trait syndrome. Specifically, a unidimensional leaf economics spectrum revealed that bamboo with higher leaf N concentrations (LNC), P concentrations (LPC), and specific leaf area (SLA) adopted a "fast acquisitive" above-ground strategy, while broadleaf species with thicker leaves employed a "slow conservative" above-ground strategy. A two-dimensional root trait syndrome indicated a "conservation" gradient with bamboo adopting a "slow conservative" below-ground strategy associated with higher root tissue density (RTD), and broadleaf species exhibiting a "fast acquisitive" below-ground strategy linked to higher root N concentrations (RNC) and P concentrations (RPC), and a "collaboration" gradient probably ranging from broadleaf species with a "do-it-yourself" strategy characterized by high specific root length (SRL), to bamboo adopting an "outsourcing" strategy with thicker roots. In conclusion, key trait divergence from coexisting broadleaf species, higher phenotypic integration, and multi-dimensional opposite above- and below-ground resource strategies confer competitive advantages to moso bamboo, shedding light on the mechanistic understanding of its invasion into subtropical evergreen broadleaf forest and providing theoretical guidance for maintaining the stability of subtropical forest ecosystem

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

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

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

Carbon, Nitrogen, and Phosphorus Stoichiometry between Leaf and Soil Exhibit the Different Expansion Stages of Moso Bamboo (Phyllostachys edulis (Carriere) J. Houzeau) into Chinese Fir (Cunninghamia lanceolata (Lamb.) Hook.) Forest

The expansion of Moso bamboo (Phyllostachys edulis (Carriere) J. Houzeau) has triggered native forest retreat and a range of ecological issues, especially for the Chinese fir (Cunninghamia lanceolata (Lamb.) Hook.) forests with similar growing conditions. In order to reveal the stoichiometric characteristics of Moso bamboo succession and scientifically control the forest retreat caused by the expansion of Moso bamboo into Chinese fir, mixed forests including 0%, 30%, 50%, 60%, and 80% of Moso bamboo expanded into Chinese fir forests were used to simulate the expansion stages I, II, III, IV, and V, respectively. In addition, by measuring the C, N, and P contents in Moso bamboo leaves and soils and calculating the correlation stoichiometric ratios, the correlation and coupling of which were explored and combined with an ecological homeostasis model at different stages of Moso bamboo expansion. The results demonstrated that P was a key element for the high utilization of Moso bamboo growth, and the expansion principle was influenced by N limitation. The conclusion was that the anthropogenic regulation of C content in soil could achieve the purpose of expansion control and exploit the carbon sequestration capacity in the mixed forest with half Moso bamboo and half Chinese fir, which should discourage the expansion

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

Carbon Density Change Characteristics and Driving Factors During the Natural Succession of Forests on Xinglong Mountain in the Transition Zone Between the Qinghai&ndash;Tibet and Loess Plateaus

The transition zone between the Qinghai&ndash;Tibet and Loess Plateaus is an important ecological functional area and carbon (C) reservoir in China. Studying the main drivers of C density changes in forest ecosystems is crucial to enhance the C sink potential of those ecosystems in ecologically fragile regions. In this study, four stand types at different succession stages in the transition zone of Xinglong Mountain were selected as the study objective. The C densities of the ecosystem, vegetation, plant debris, and soil of each stand type were estimated, and the related driving factors were quantified. The results showed that the forest ecosystem C density continuously increased significantly with natural succession (381.23 Mg/hm2 to 466.88 Mg/hm2), indicating that the ecosystem has a high potential for C sequestration with progressive forest succession. The increase in ecosystem C density was mainly contributed to by the vegetation C density, which was jointly affected by the vegetation characteristics (C sink, mean diameter at breast height, mean tree height), litter C/N (nitrogen), and surface soil C/N, with factors explaining 95.1% of the variation in vegetation C density, while the net effect of vegetation characteristics was the strongest (13.9%). Overall, this study provides a new insight for understanding the C cycle mechanism in ecologically fragile areas and further improves the theoretical framework for understanding the C sink function of forest ecosystems