Preston’s equation is a general model describing the egg shape of birds. The parameters of Preston’s equation are usually estimated after re-expressing it as the Todd-Smart equation and scaling the egg’s actual length to two. This method assumes that the straight line through the two points on an egg’s profile separated by the maximum distance (i.e., the longest axis of an egg’s profile) is the mid-line. It hypothesizes that the photographed egg’s profile is perfectly bilaterally symmetrical, which seldom holds true because of photographic errors and placement errors. The existing parameter estimation method for Preston’s equation considers an angle of deviation for the longest axis of an egg’s profile from the mid-line, which decreases prediction errors to a certain degree. Nevertheless, this method cannot provide an accurate estimate of the coordinates of the egg’s center, and it leads to sub-optimal parameter estimation. Thus, it is better to account for the possible asymmetry between the two sides of an egg’s profile along its mid-line when fitting egg-shape data. In this paper, we propose a method based on the optimization algorithm (optimPE) to fit egg-shape data and better estimate the parameters of Preston’s equation by automatically searching for the optimal mid-line of an egg’s profile and testing its validity using profiles of 59 bird eggs spanning a wide range of existing egg shapes. We further compared this method with the existing one based on multiple linear regression (lmPE). This study demonstrated the ability of the optimPE method to estimate numerical values of the parameters of Preston’s equation and provide the theoretical egg length (i.e., the distance between two ends of the mid-line of an egg’s profile) and the egg’s maximum breadth. This provides a valuable approach for comparing egg shapes among conspecifics or across different species, or even different classes (e.g., birds and reptiles), in future investigations.

/pdf/a-new-program-to-estimate-the-parameters-of-preston-s-3t7vww9s.pdf

A New Program to Estimate the Parameters of Preston's Equation, a General Formula for Describing the Egg Shape of Birds

COPYRIGHT © 2023 Niklas, Shi, Gielis, Schrader and Niinemets. This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms. TYPE Editorial PUBLISHED 21 March 2023 DOI 10.3389/fpls.2023.1169558

/pdf/editorial-leaf-functional-traits-ecological-and-evolutionary-22llegsq.pdf

Editorial: Leaf functional traits: Ecological and evolutionary implications

Leaf shape and size can vary between hybrids and their parents. However, this has seldom been quantitatively tested. Photinia × fraseri is an important landscaping plant in East Asia as a hybrid between evergreen shrubs P. glabra and P. serratifolia. Its leaf shape looks like that of P. serratifolia. To investigate leaf shape, we used a general equation for calculating the leaf area (A) of broad-leaved plants, which assumes a proportional relationship between A and product of lamina length (L) and width (W). The proportionality coefficient (which is referred to as the Montgomery parameter) serves as a quantitative indicator of leaf shape, because it reflects the proportion of leaf area A to the area of a rectangle with L and W as its side lengths. The ratio of L to W, and the ellipticalness index were also used to quantify the complexity of leaf shape for elliptical leaves. A total of >4000 leaves from P. × fraseri and P. serratifolia (with >2000 leaves for each taxon) collected on a monthly basis was used to examine: (i) whether there is a significant difference in leaf shape between the two taxa, and (ii) whether there is a monotonic or parabolic trend in leaf shape across leaf ages. There was a significant difference in leaf shape between the two taxa (p < 0.05). Although there were significant differences in leaf shape on a monthly basis, the variation in leaf shape over time was not large, i.e., leaf shape was relatively stable over time for both taxa. However, the leaf shape of the hybrid was significantly different from its parent P. serratifolia, which has wider and more elliptical leaves than the hybrid. This work demonstrates that variations in leaf shape resulting from hybridization can be rigorously quantified and compared among species and their hybrids. In addition, this work shows that leaf shape does not changes as a function of age either before or after the full expansion of the lamina.

/pdf/comparison-of-leaf-shape-between-a-photinia-hybrid-and-one-3tltnx67.pdf

Comparison of Leaf Shape between a Photinia Hybrid and One of Its Parents

The inequality in leaf and fruit size distribution per plant can be quantified using the Gini index, which is linked to the Lorenz curve depicting the cumulative proportion of leaf (or fruit) size against the cumulative proportion of the number of leaves (or fruits). Prior researches have predominantly employed empirical models-specifically the original performance equation (PE-1) and its generalized counterpart (GPE-1)-to fit rotated and right-shifted Lorenz curves. Notably, another potential performance equation (PE-2), capable of generating similar curves to PE-1, has been overlooked and not systematically compared with PE-1 and GPE-1. Furthermore, PE-2 has been extended into a generalized version (GPE-2). In the present study, we conducted a comparative analysis of these four performance equations, evaluating their applicability in describing Lorenz curves related to plant organ (leaf and fruit) size. Leaf area was measured on 240 culms of dwarf bamboo (Shibataea chinensis Nakai), and fruit volume was measured on 31 field muskmelon plants (Cucumis melo L. var. agrestis Naud.). Across both datasets, the root-mean-square errors of all four performance models were consistently smaller than 0.05. Paired t-tests indicated that GPE-1 exhibited the lowest root-mean-square error and Akaike information criterion value among the four performance equations. However, PE-2 gave the best close-to-linear behavior based on relative curvature measures. This study presents a valuable tool for assessing the inequality of plant organ size distribution. 

Comparison of four performance models in quantifying the inequality of leaf and fruit size distribution

The Lamé curve is an extension of an ellipse, the latter being a special case. Dr. Johan Gielis further extended the Lamé curve in the polar coordinate system by introducing additional parameters (n1, n2, n3; m): rφ=1Acosm4φn2+1Bsinm4φn3−1/n1, which can be applied to model natural geometries. Here, r is the polar radius corresponding to the polar angle φ; A, B, n1, n2 and n3 are parameters to be estimated; m is the positive real number that determines the number of angles of the Gielis curve. Most prior studies on the Gielis equation focused mainly on its applications. However, the Gielis equation can also generate a large number of shapes that are rotationally symmetric and axisymmetric when A = B and n2 = n3, interrelated with the parameter m, with the parameters n1 and n2 determining the shapes of the curves. In this paper, we prove the relationship between m and the rotational symmetry and axial symmetry of the Gielis curve from a theoretical point of view with the condition A = B, n2 = n3. We also set n1 and n2 to take negative real numbers rather than only taking positive real numbers, then classify the curves based on extremal properties of r(φ) at φ = 0, π/m when n1 and n2 are in different intervals, and analyze how n1, n2 precisely affect the shapes of Gielis curves.

/pdf/effects-of-the-numerical-values-of-the-parameters-in-the-1zj5bb1j.pdf

Effects of the Numerical Values of the Parameters in the Gielis Equation on Its Geometries

Egg geometry can be described using Preston's equation, which has seldom been used to calculate egg volume (V) and surface area (S) to explore S versus V scaling relationships. Herein, we provide an explicit re‐expression of Preston's equation (designated as EPE) to calculate V and S, assuming that an egg is a solid of revolution. The side (longitudinal) profiles of 2221 eggs of six avian species were digitized, and the EPE was used to describe each egg profile. The volumes of 486 eggs from two avian species predicted by the EPE were compared with those obtained using water displacement in graduated cylinders. There was no significant difference in V using the two methods, which verified the utility of the EPE and the hypothesis that eggs are solids of revolution. The data also indicated that V is proportional to the product of egg length (L) and maximum width (W) squared. A 2/3‐power scaling relationship between S and V for each species was observed, that is, S is proportional to (LW2)2/3. These results can be extended to describe the shapes of the eggs of other species to study the evolution of avian (and perhaps reptilian) eggs.

A simple way to calculate the volume and surface area of avian eggs

Many geometries of plant organs can be described by the Gielis equation, a polar coordinate equation extended from the superellipse equation, r=a|cosm4φ|n2+|1ksinm4φ|n3−1/n1. Here, r is the polar radius corresponding to the polar angle φ; m is a positive integer that determines the number of angles of the Gielis curve when φ ∈ [0 to 2π); and the rest of the symbols are parameters to be estimated. The pentagonal radial symmetry of calyxes and corolla tubes in top view is a common feature in the flowers of many eudicots. However, prior studies have not tested whether the Gielis equation can depict the shapes of corolla tubes. We sampled randomly 366 flowers of Vinca major L., among which 360 had five petals and pentagonal corolla tubes, and six had four petals and quadrangular corolla tubes. We extracted the planar coordinates of the outer rims of corolla tubes (in top view) (ORCTs), and then fitted the data with two simplified versions of the Gielis equation with k = 1 and m = 5: r=acos54φn2+sin54φn3−1/n1 (Model 1), and r=acos54φn2+sin54φn2−1/n1 (Model 2). The adjusted root mean square error (RMSEadj) was used to evaluate the goodness of fit of each model. In addition, to test whether ORCTs are radially symmetrical, we correlated the estimates of n2 and n3 in Model 1 on a log-log scale. The results validated the two simplified Gielis equations. The RMSEadj values for all corolla tubes were smaller than 0.05 for both models. The numerical values of n2 and n3 were demonstrated to be statistically equal based on the regression analysis, which suggested that the ORCTs of V. major are radially symmetrical. It suggests that Model 1 can be replaced by the simpler Model 2 for fitting the ORCT in this species. This work indicates that the pentagonal or quadrangular corolla tubes (in top view) can both be modeled by the Gielis equation and demonstrates that the pentagonal or quadrangular corolla tubes of plants tend to form radial symmetrical geometries during their development and growth.

/pdf/quantifying-the-variation-in-the-geometries-of-the-outer-399y9fjv.pdf

Quantifying the Variation in the Geometries of the Outer Rims of Corolla Tubes of Vinca major L.

Foliage leaves are essential for plant survival and growth, and how plants allocate biomass to their leaves reveals their economic and ecological strategies. Prior studies have shown that leaf-age significantly influences leaf biomass allocation patterns. However, unravelling the effects of ontogeny on partitioning biomass remains a challenge because it is confounded by the effects of environmental factors. Here, we aim to elucidate whether leaf-age affects the allocation to the lamina and petiole by examining leaves of known age growing in the same general environmental context. We sampled 2698 Photinia serratifolia leaves developing in the same environment from April to November 2021, representing eight leaf-ages (n > 300 for each leaf-age). Petiole and lamina biomass, and lamina area were measured to evaluate the scaling relationships using reduced major axis regression protocols. The bootstrap percentile method was used to determine the differences in scaling exponents among the different leaf-ages. ANOVA with Tukey’s HSD was used to compare the ratios of petiole and lamina biomass to lamina area across the leaf-ages. Correlation tests were used to determine if exponents, intercepts, and ratios differed significantly across the different leaf-ages. The data indicated that (i) the ratio of petiole and lamina biomass to lamina area and the scaling exponent of lamina biomass versus lamina area correlate positively with leaf-age, and (ii) the scaling exponent of petiole biomass versus lamina area correlates negatively with leaf-age. Leaf maturation process involves an inverse proportional allocation between lamina and petiole biomass for expanding photosynthetic area. This phenomenon underscores the effect of leaf-age on biomass allocation and the importance of adopting an ontogenetic perspective when entertaining plant scaling theories and unravelling the principles governing shifts in biomass allocation throughout the leaf lifespan.

Leaf-age and petiole biomass play significant roles in leaf scaling theory

The proportions in the size of the avian egg albumen, yolk, and shell are crucial for understanding bird survival and reproductive success, because their relationships with volume and surface area can affect ecological and life history strategies. Prior studies have focused on the relationship between the albumen and the yolk, but little is known about the scaling relationship between eggshell mass and shape, and the mass of the albumen and the yolk. Toward this end, 691 eggs of six precocial species were examined, and their 2-D egg profiles were photographed and digitized. The explicit Preston equation, which assumes bilateral symmetrical geometry, was used to fit the 2-D egg profiles and to calculate surface areas and volumes based on the hypothesis that eggs can be treated as solids of profile revolution. The scaling relationships of eggshell mass (Ms), albumen mass (Ma), and yolk mass (My), as well as the surface area (S), volume (V), and total mass (Mt) were determined. The explicit Preston equation was validated in describing the 2-D egg profiles. The scaling exponents of Ma vs. Ms, My vs. Ms, and My vs. Ma were smaller than unity, indicating that increases in Ma and My fail to keep pace with increases in Ms, and that increases in My fail to keep pace with increases in Ma. Therefore, increases in unit nutrient contents (i.e., the yolk) involve disproportionately larger increases in eggshell mass and disproportionately larger increases in albumen mass. The data also revealed a 2/3-power scaling relationship between S and V for each species, i.e., simple Euclidean geometry is obeyed. These findings help to inform our understanding of avian egg construction and reveal evolutionary interspecific trends in the scaling of egg shape, volume, mass, and mass allocation.

Scaling relationships among the mass of eggshell, albumen, and yolk in six precocial birds.

Abstract 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 ∝ 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.

Scaling relationships of leaf vein and areole traits versus leaf size for nine Magnoliaceae species differing in venation density

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