Leaf lamina mass and area are closely correlated with the photosynthetic capacity and competitive ability of plants, whereas leaf age has been demonstrated to affect physiological processes such as photosynthesis. However, it remains unknown whether the lamina mass vs. area scaling relationship is influenced by leaf age, which is important for understanding plant adaptive strategies and, more broadly, resource utilization and growth. We measured the leaf functional traits of five leaf-age groups of Photinia × fraseri for a total of 1,736 leaves. ANOVA followed by Tukey’s honestly significant difference test was used to compare the functional traits among the five leaf-age groups. Reduced major axis regression protocols were used to fit the scaling relationship between lamina mass and area, and the bootstrap percentile method was used to compare the lamina mass vs. area scaling relationships among the leaf-age groups. Lamina area, and the ratio of lamina dry mass to lamina fresh mass increased with increasing leaf age. Lamina fresh mass per unit area, and lamina dry mass per unit area both exhibited a parabolic-like trend as leaf age increased, i.e., at the leaf maturation stage, it showed a slight but significant decline. The phenomenon called diminishing returns were confirmed by each of the five leaf-age groups, i.e., all of the numerical values of the scaling exponents of lamina mass vs. area were significantly greater than 1. There were significant differences in the scaling exponents of lamina mass vs. area for the leaves across different sampling times. The scaling exponents were lower at the early rapid growth stage, indicating a lower cost for increasing leaf area compared to the leaf maturation stage. These data are consistent with leaves undergoing a transition from resource acquisition to resource conservation in the process of their development and growth.
Leaf shape is closely related to economics of leaf support and leaf functions, including light interception, water use, and CO2 uptake, so correct quantification of leaf shape is helpful for studies of leaf structure/function relationships. There are some extant indices for quantifying leaf shape, including the leaf width/length ratio (W/L), leaf shape fractal dimension (FD), leaf dissection index, leaf roundness index, standardized bilateral symmetrical index, etc. W/L ratio is the simplest to calculate, and recent studies have shown the importance of the W/L ratio in explaining the scaling exponent of leaf dry mass vs. leaf surface area and that of leaf surface area vs. leaf length. Nevertheless, whether the W/L ratio could reflect sufficient geometrical information of leaf shape has been not tested. The FD might be the most accurate measure for the complexity of leaf shape because it can characterize the extent of the self-similarity and other planar geometrical features of leaf shape. However, it is unknown how strongly different indices of leaf shape complexity correlate with each other, especially whether W/L ratio and FD are highly correlated. In this study, the leaves of nine Magnoliaceae species (>140 leaves for each species) were chosen for the study. We calculated the FD value for each leaf using the box-counting approach, and measured leaf fresh mass, surface area, perimeter, length, and width. We found that FD is significantly correlated to the W/L ratio and leaf length. However, the correlation between FD and the W/L ratio was far stronger than that between FD and leaf length for each of the nine species. There were no strong correlations between FD and other leaf characteristics, including leaf area, ratio of leaf perimeter to area, fresh mass, ratio of leaf fresh mass to area, and leaf roundness index. Given the strong correlation between FD and W/L, we suggest that the simpler index, W/L ratio, can provide sufficient information of leaf shape for similarly-shaped leaves. Future studies are needed to characterize the relationships among FD and W/L in leaves with strongly varying shape, e.g., in highly dissected leaves.
Premise The phenomenon called “diminishing returns” refers to a scaling relationship between lamina mass (M) vs. lamina area (A) in many species, i.e., M ∝ Aα>1, where α is the scaling exponent exceeding unity. Prior studies have focused on the scaling relationships between lamina dry mass (DM) and A, or between fresh mass (FM) and A. However, the scaling between petiole mass and M and A has seldom been investigated. Here, we examine the scaling relationships among FM, DM, A, and petiole fresh mass (PFM). Methods For each of 3268 leaves from nine Lauraceae species, FM, DM, A, and PFM were measured, and their scaling relationships were fitted using reduced major axis regression protocols. The bootstrap percentile method was used to test the significance of the difference in α‐values between any two species. Results The phenomenon of diminishing returns was verified between FM vs. A and DM vs. A. The FM vs. A scaling relationship was statistically more robust than the DM vs. A scaling relationship based on bivariate regression r2‐values. Diminishing returns were also observed for the PFM vs. FM and PFM vs. A scaling relationships. The PFM vs. FM scaling relationship was statistically more robust than the PFM vs. A scaling relationship. Conclusions “Diminishing returns” was confirmed among the FM, DM, A, and PFM scaling relationships. The data collectively indicate that the petiole scales mechanically more strongly with lamina mass than with area, suggesting that static (self) loading takes precedence over dynamic (wind) loading.
Premise Across species, main leaf vein density scales inversely with leaf area (A). Yet, minor vein density manifests no clear relationship with respect to A, despite having the potential to provide important insights into the trade‐off among the investments in leaf mechanical support, hydraulics, and light interception. Methods To examine this phenomenon, the leaves of nine Magnoliaceae leaves were sampled, and the scaling relationships among A and midrib length (ML), total vein length (TVL), total vein area (TVA), total areole area (TAA), and mean areole area (MAA) were determined. The scaling relationships between MAA and areole density (the number of areoles per unit leaf area) and between MAA and A were also analyzed. Results For five of the nine species, A was proportional to ML2. For eight of the nine species, TVL and TVA were both proportional to A. The numerical values of the scaling exponents for TAA vs. A were between 1.0 and 1.07 for eight species; i.e., as expected, TAA was isometrically proportional to A. There was no correlation between MAA and A, but MAA scaled inversely with respect to areole density for each species. Conclusions The correlation between midrib “density” (i.e., ML/A) and A, and the lack of correlation between total leaf vein density and A result from the A∝ $\propto $ ML2 scaling relationship and the proportional relationship between TVL and A, respectively. Leaves with the same size can have widely varying MAA. Thus, leaf size itself does not directly constrain leaf hydraulic efficiency and redundancy.
Many plant species produce ovate leaves, but there is no general parametric model for describing this shape. Here, we used two empirical nonlinear equations, the beta and Lobry–Rosso–Flandrois (LRF) equations, and their modified forms (referred to as the Mbeta and MLRF equations for convenience), to generate bilaterally symmetrical curves along the x-axis to form ovate leaf shapes. In order to evaluate which of these four equations best describes the ovate leaf shape, we used 14 leaves from 7 Neocinnamomum species (Lauraceae) and 72 leaves from Chimonanthus praecox (Calycanthaceae). Using the AIC and adjusted root mean square error to compare the fitted results, the modified equations fitted the leaf shapes better than the unmodified equations. However, the MLRF equation provided the best overall fit. As the parameters of the MLRF equation represent leaf length, maximum leaf width, and the distance from leaf apex to the point associated with the maximum leaf width along the leaf length axis, these findings are potentially valuable for studying the influence of environmental factors on leaf shape, differences in leaf shape among closely related plant species with ovate leaf shapes, and the extent to which leaves are bilaterally symmetrical. This is the first work in which temperature-dependent developmental equations to describe the ovate leaf shape have been employed, as previous studies lacked similar leaf shape models. In addition, prior work seldom attempted to describe real ovate leaf shapes. Our work bridges the gap between theoretical leaf shape models and empirical leaf shape indices that cannot predict leaf shape profiles.
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