Abstract
Leaves, as the most important photosynthetic organ of plants, are intimately associated with plant function and adaptation to environmental changes. The scaling relationship of the leaf dry mass (or the fresh mass) vs. leaf surface area has been referred to as “diminishing returns”, suggesting that the leaf area fails to increase in proportion to leaf dry mass (or fresh mass). However, previous studies used materials across different families, and there is lack of studies testing whether leaf fresh mass is proportional to the leaf dry mass for the species in the same family, and examining the influence of the scaling of leaf dry mass vs. fresh mass on two kinds of diminishing returns based on leaf dry mass and fresh mass. Bamboo plants (Poaceae: Bambusoideae) are good materials for doing such a study, which have astonishingly similar leaf shapes across species. Bamboo leaves have a typical parallel venation pattern. In general, a parallel venation pattern tends to produce a more stable symmetrical leaf shape than the pinnate and palmate venation patterns. The symmetrical parallel veins enable leaves to more regularly hold water, which is more likely to result in a proportional relationship between the leaf dry mass and absolute water content, which consequently determines whether the scaling exponent of the leaf dry mass vs. area is significantly different from (or the same as) that of the leaf fresh mass vs. area. In the present study, we used the data of 101 bamboo species, cultivars, forms and varieties (referred to as 101 (bamboo) taxa below for convenience) to analyze the scaling relationships between the leaf dry mass and area, and between leaf fresh mass and area. We found that the confidence intervals of the scaling exponents of the leaf fresh mass vs. dry mass of 68 out of the 101 taxa included unity, which indicates that for most bamboo species (67.3%), the increase in leaf water mass keeps pace with that of leaf dry mass. There was a significant scaling relationship between either leaf dry mass or fresh mass, and the leaf surface area for each studied species. We found that there was no significant difference between the scaling exponent of the leaf dry mass vs. leaf area and that of the leaf fresh mass vs. leaf area when the leaf dry mass was proportional to the leaf fresh mass. The goodness of fit to the linearized scaling relationship of the leaf fresh mass vs. area was better than that of the leaf dry mass vs. area for each of the 101 bamboo taxa. In addition, there were significant differences in the normalized constants of the leaf dry mass vs. fresh mass among the taxa (i.e., the differences in leaf water content), which implies the difference in the adaptabilities to different environments across the taxa.
Highlights
The growth of plants cannot be separated from the contributions of leaves
We examined 10,045 leaves from 101 bamboo taxa from 17 genera, aiming to determine (i) whether the leaf fresh mass was proportional to the leaf dry mass, or if there was an allometric relationship, and what proportion of species showed a proportional or allometric relationship; (ii) what proportion of species showed “diminishing returns”, referring to the scaling relationship between the leaf fresh/dry mass and leaf surface area; (iii) and whether the majority of the studied bamboo species exhibited a better scaling relationship between the leaf fresh mass vs. leaf surface area rather than the leaf dry mass vs. leaf surface area
Symmetry 2020, 1345of leaf dry mass to fresh mass was significantly different among the 101 bamboo taxa4 of 10 (Figure 1 and Table S2 in the online Supplementary Materials)
Summary
The fundamental leaf functional traits (e.g., leaf biomass, water content, leaf shape, leaf surface area, etc.) profoundly affect a variety of biological processes, including plant growth, survival, reproduction and ecosystem function [1,2,3]. Our previous studies have found this same allometric relationship between the leaf dry mass and leaf surface area for five Lauraceae, five Oleaceae, eight Rosaceae, ten climbing species and seventeen bamboo species [5,6,9,10]. This “diminishing returns” phenomenon might occur because leaf dry matter is unevenly distributed in the leaf. Plants normally invest more dry mass to the side facing the sunlight, and invest less mass in the shaded side [2,11]
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