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Carbon limitation, stem growth rate and the biomechanical cause of Corner's rules.

ABSTRACT: Background and aims:Corner's rules describe a global spectrum from large-leaved plants with thick, sparingly branched twigs with low-density stem tissues and thick piths to plants with thin, highly branched stems with high-density stem tissues and thin piths. The hypothesis was tested that, if similar crown areas fix similar amounts of carbon regardless of leaf size, then large-leaved species, with their distantly spaced leaves, require higher stem growth rates, lower stem tissue densities and stiffnesses, and therefore thicker twigs. Methods:Structural equation models were used to test the compatibility of this hypothesis with a dataset on leaf size, shoot tip spacing, stem growth rate and dimensions, and tissue density and mechanics, sampling 55 species drawn from across the angiosperm phylogeny from a morphologically diverse dry tropical community. Key results:Very good fit of structural equation models showed that the causal model is highly congruent with the data. Conclusions:Given similar amounts of carbon to allocate to stem growth, larger-leaved species require greater leaf spacing and therefore greater stem extension rates and longer stems, in turn requiring lower-density, more flexible, stem tissues than small-leaved species. A given stem can have high resistance to bending because it is thick (has high second moment of area I) or because its tissues are stiff (high Young's modulus E), the so-called E-I trade-off. Because of the E-I trade-off, large-leaved species have fast stem growth rates, low stem tissue density and tissue stiffness, and thick twigs with wide piths and thick bark. The agreement between hypothesis and data in structural equation analyses strongly suggests that Corner's rules emerge as the result of selection favouring the avoidance of self-shading in the context of broadly similar rates of carbon fixation per unit crown area across species.

SUBMITTER: Olson ME

PROVIDER: S-EPMC6153482 | biostudies-literature | 2018 Sep

REPOSITORIES: biostudies-literature

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Carbon limitation, stem growth rate and the biomechanical cause of Corner's rules.

Olson Mark E ME Rosell Julieta A JA Zamora Muñoz Salvador S Castorena Matiss M

Annals of botany 20180901 4

<h4>Background and aims</h4>Corner's rules describe a global spectrum from large-leaved plants with thick, sparingly branched twigs with low-density stem tissues and thick piths to plants with thin, highly branched stems with high-density stem tissues and thin piths. The hypothesis was tested that, if similar crown areas fix similar amounts of carbon regardless of leaf size, then large-leaved species, with their distantly spaced leaves, require higher stem growth rates, lower stem tissue densiti ...[more]

PMID: 29889257

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Project description:The orientation of vascular canals in primary bone may reflect differences in growth rate and/or adaptation to biomechanical loads. Previous studies link specific canal orientations to bone growth rates, but results between different taxa are contradictory. Circumferential vascular canals (forming laminar bone) have been hypothesized to reflect either (or both) rapid growth rate or locomotion-induced torsional loading. Previous work on the hindlimb biomechanics in the emu shows that the femur and tibiotarsus experience large shear strains, likely resulting from torsional loads that increase through ontogeny. Here, we test how growth rate and biomechanical loading affect bone laminarity in wing and hindlimb elements from growing emu (2-60 wks). If laminar bone is an adaptation to torsion-induced shear strains, it should increase from juveniles to adults. Alternatively, if bone laminarity reflects rapid growth, as has been shown previously in emu, it should be abundant in fast-growing juveniles and decrease with age. Transverse mid-shaft histological sections from the limb bones (femur, tibiotarsus, humerus, ulna, and radius) were prepared and imaged. Growth rates were measured using fluorescent bone labels. Vascular canal orientation was quantified using laminarity index (proportion of circumferential canals). Principal components analysis was performed to convert highly correlated variables (i.e., mass, age, growth rate, and shear strain) into principal components. Random-intercept beta regression modeling determined which principal components best explained laminarity. The fastest growth rates were found in young individuals for all five skeletal elements. Maximum growth rate did not coincide with peak laminarity. Instead, in the femur and tibiotarsus, elevated laminarity is strongly correlated with adult features such as large size, old age, and modest growth rate. This result is contrary to predictions made based on a previous study of emu but is consistent with results observed in some other avian species (penguin, chicken). Shear strain in the caudal octant of the femur and tibiotarsus is positively correlated with laminarity but has a weaker effect on laminarity relative to mass, age, and growth rate. Laminarity in the wing elements is variable and does not correlate with ontogenetic factors (including mass, age, and growth rate). Its presence may relate to relaxed developmental canalization or a retained ancestral feature. In conclusion, ontogeny (including growth rate) is the dominant influence on vascular canal orientation at least in the hindlimb of the emu.

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Carbon limitation, stem growth rate and the biomechanical cause of Corner's rules.

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Carbon limitation, stem growth rate and the biomechanical cause of Corner's rules.

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