Metabolic Scaling Theory Research Articles

Tree growth as affected by stem and crown structure

Key messagePrediction of tree growth based on size or mass as proposed by the Metabolic Scaling Theory is an over-simplification and can be significantly improved by consideration of stem and crown morphology.Tree growth and metabolic scaling theory, as well as corresponding growth equations, use tree volume or mass as predictors for growth. However, this may be an over-simplification, as the future growth of a tree may, in addition to volume or mass, also depend on its past development and aspects of the current inner structure and outer morphology. The objective of this evaluation was to analyse the effect of selected structural and morphological tree characteristics on the growth of common tree species in Europe. Here, we used eight long-term experiments with a total of 24 plots and extensive individual measurements of 1596 trees in monospecific stands of European beech (Fagus sylvatica L.), Norway spruce (Picea abies (L.) Karst.), Scots pine (Pinus sylvestris L.) and sessile oak (Quercus petraea (Matt.) Liebl.). Some of the experiments have been systematically surveyed since 1870. The selected plots represent a broad range of stand density, from fully to thinly stocked stands. We applied linear mixed models with random effects for analysing and modelling how tree growth and productivity are affected by stem and crown structure. We used the species-overarching relationship mathrm{iv}={{a}_{0}times v} between stem volume growth, mathrm{iv}and stem volume, v, as the baseline model. In this model {a}_{0} represents the allometric factor and α the allometric exponent. Then we included tree age, mean stem volume of the stand and structural and morphological tree variables in the model. This significantly reduced the AIC; RMSE was reduced by up to 43%. Interestingly, the full model estimating mathrm{iv} as a function of v and mean tree volume, crown projection area, crown ratio and mean tree ring width, revealed a alpha cong 3/4 scaling for the relationship between mathrm{iv}propto {v}^{alpha }. This scaling corresponded with Kleiber’s rule and the West-Brown-Enquist model of the metabolic scaling theory. Simplified approaches based on stem diameter or tree mass as predictors may be useful for a rough estimation of stem growth in uniform stands and in cases where more detailed predictors are not available. However, they neglect other stem and crown characteristics that can have a strong additional effect on the growth behaviour. This becomes of considerable importance in the heterogeneous mixed-species stands that in many countries of the world are designed for forest restoration. Heterogeneous stand structures increase the structural variability of the individual trees and thereby cause a stronger variation of growth compared with monocultures. Stem and crown characteristics, which may improve the analysis and projection of tree and stand dynamics in the future forest, are becoming more easily accessible by Terrestrial laser scanning.

The Influence of Ecosystem and Phylogeny on Tropical Tree Crown Size and Shape

The sizes and shapes of tree crowns are of fundamental importance in ecology, yet understanding the forces that determine them remains elusive. A cardinal question facing ecologists is the degree to which general and non-specific versus ecological and context-dependent processes are responsible for shaping tree crowns. Here, we test this question for the first time across diverse tropical ecosystems. Using trees from 20 plots varying in elevation, precipitation, and ecosystem type (savanna-forest transitions) across the paleo- and neo-tropics, we test the relationship between crown dimensions and tree size. By analyzing these scaling relationships across environmental gradients, biogeographic regions, and phylogenetic distance, we extend Metabolic Scaling Theory (MST) predictions to include how local selective pressures shape variation in crown dimensions. Across all sites, we find strong agreement between mean trends and MST predictions for the scaling of crown size and shape, but large variation around the mean. While MST explained approximately half of the observed variation in tree crown dimensions, we find that local, ecosystem, and phylogenetic predictors account for the half of the residual variation. Crown scaling does not change significantly across regions, but does change across ecosystem types, where savanna tree crowns grow more quickly with tree girth than forest tree crowns. Crowns of legumes were wider and more voluminous than those of other taxa. Thus, while MST can accurately describe the central tendency of tree crown size, local ecological conditions and evolutionary history appear to modify the scaling of crown shape. Importantly, our extension of MST incorporating these differences accounts for the mechanisms driving variation in the scaling of crown dimensions across the tropics. We present allometric equations for the prediction of crown dimensions across tropical ecosystems. These results are critical when scaling the function of individual trees to larger spatial scales or incorporating the size and shape of tree crowns in global biogeochemical models.

Height-diameter allometric relationships for seedlings and trees across China

Height–diameter allometric (H-D) relationships play an important role in the light capture and stability of a tree, and it is poorly understood whether the relationships changes with growth stages or functional groups along a large scale environmental gradients. We present a comparative study of H-D relationships of 6810 seedlings and 19,707 trees from about 1000 species in 9 different sites across China. We showed that allometric exponents for seedlings and trees differ between different sites and do not represent a single, constant theoretical value (e.g. 2/3, 1/2 or 1). Seedlings had greater exponents than trees in most sites. The exponents varied between canopy and understory trees in 5 sites, while were indistinguishable from each other in the remaining 4 sites. Canopy-tree seedlings had smaller exponents than understory-tree seedlings in 3 sites but had indistinguishable exponents in the other 6 sites. Gymnosperm trees had exponents greater than or indistinguishable from angiosperm trees. Elevation alone, or together with mean temperature of the warmest month explained variation of tree allometries for canopy-tree and understory-tree seedlings. The exponents of canopy trees decreased with mean annual precipitation. Our results do not support the predictions from metabolic scaling theory or biomechanical models that height–diameter allometries are invariant. Our study provides insight into how ontogeny, adult stature, phyletic affiliations and environmental limitations affect height–diameter allometric relationships at biogeographical scales.

Effects of climate, biotic factors, and phylogeny on allometric relationships: testing the metabolic scaling theory in plantations and natural forests across China

BackgroundMetabolic scaling theory (MST) is still in debate because observed allometric exponents often deviate from MST predictions, and can change significantly depending on environment, phylogeny, and disturbance. We assembled published scaling exponents from literatures for three allometric relationships linked to biomass allocation: leaf biomass-diameter (L-D), stem biomass-diameter (S-D), and root biomass-diameter (R-D). We used data from natural forests and plantations across China to test the following hypotheses: 1) the allometric relationships of trees support the predictions of MST on a broad scale; 2) the observed deviations from MST predictions are caused by climate, biotic factors, and/or phylogeny; 3) abiotic and biotic factors influence allometric relationships in plantations and natural forests differently, and different allometric relationships (i.e. L-D, S-D, and R-D) are affected differently. We related these scaling exponents to geographic climate gradient, successional stage, stand density, leaf form and phenology, and phylogeny. We used mixed-effect models to examine the major factors affecting tree allometries.ResultsIn natural forests, S-D and R-D scaling exponents were consistent with MST predictions in primary forests, but were significantly lower in secondary forests. Both S-D and R-D scaling exponents in plantations had a medium value that fell between those of the secondary and primary forests, despite plantations being similar in species characteristics and age to secondary forests. The S-D and R-D exponents were significantly affected by factors that are not yet considered in MST, including winter coldness which explained 2.76% – 3.24% of variations, successional stage (7.91% – 8.20% of variations), density (a surrogate for competition, 5.86% – 8.54% of variations), and especially phylogeny (45.86% – 56.64% of variations explained). However, the L-D scaling exponents conformed to MST predictions in primary, secondary, and plantation forests, and was not strongly explained by most factors.ConclusionMST is only applicable to primary (steady-state) forests, and climate, biotic factors and phylogeny are causes of the observed deviations of allometric relationships from MST predictions. Forest management practices in plantations have a strong influence on tree allometries. L-D allometry is more strongly controlled by biophysical constraints than S-D and R-D allometries, however, the mechanisms behind this difference still need further examinations.

Validation of demographic equilibrium theory against tree-size distributions and biomass density in Amazonia

Abstract. Predicting the response of forests to climate and land-use change depends on models that can simulate the time-varying distribution of different tree sizes within a forest – so-called forest demography models. A necessary condition for such models to be trustworthy is that they can reproduce the tree-size distributions that are observed within existing forests worldwide. In a previous study, we showed that demographic equilibrium theory (DET) is able to fit tree-diameter distributions for forests across North America, using a single site-specific fitting parameter (μ) which represents the ratio of the rate of mortality to growth for a tree of a reference size. We use a form of DET that assumes tree-size profiles are in a steady state resulting from the balance between a size-independent rate of tree mortality and tree growth rates that vary as a power law of tree size (as measured by either trunk diameter or biomass). In this study, we test DET against ForestPlots data for 124 sites across Amazonia, fitting, using maximum likelihood estimation, to both directly measured trunk diameter data and also biomass estimates derived from published allometric relationships. Again, we find that DET fits the observed tree-size distributions well, with best-fit values of the exponent relating growth rate to tree mass giving a mean of ϕ=0.71 (0.31 for trunk diameter). This finding is broadly consistent with exponents of ϕ=0.75 (ϕ=1/3 for trunk diameter) predicted by metabolic scaling theory (MST) allometry. The fitted ϕ and μ parameters also show a clear relationship that is suggestive of life-history trade-offs. When we fix to the MST value of ϕ=0.75, we find that best-fit values of μ cluster around 0.25 for trunk diameter, which is similar to the best-fit value we found for North America of 0.22. This suggests an as yet unexplained preferred ratio of mortality to growth across forests of very different types and locations.

The megabiota are disproportionately important for biosphere functioning

A prominent signal of the Anthropocene is the extinction and population reduction of the megabiota—the largest animals and plants on the planet. However, we lack a predictive framework for the sensitivity of megabiota during times of rapid global change and how they impact the functioning of ecosystems and the biosphere. Here, we extend metabolic scaling theory and use global simulation models to demonstrate that (i) megabiota are more prone to extinction due to human land use, hunting, and climate change; (ii) loss of megabiota has a negative impact on ecosystem metabolism and functioning; and (iii) their reduction has and will continue to significantly decrease biosphere functioning. Global simulations show that continued loss of large animals alone could lead to a 44%, 18% and 92% reduction in terrestrial heterotrophic biomass, metabolism, and fertility respectively. Our findings suggest that policies that emphasize the promotion of large trees and animals will have disproportionate impact on biodiversity, ecosystem processes, and climate mitigation.

A General Model for Seed and Seedling Respiratory Metabolism.

The ontogeny of seed plants usually involves a dormant dehydrated state and the breaking of dormancy and germination, which distinguishes it from that of most organisms. Seed germination and seedling establishment are critical ontogenetic stages in the plant life cycle, and both are fueled by respiratory metabolism. However, the scaling of metabolic rate with respect to individual traits remains poorly understood. Here, we tested metabolic scaling theory during seed germination and early establishment growth using a recently developed model and empirical data collected from 41 species. The results show that (i) the mass-specific respiration rate (Rm) was weakly correlated with body mass, mass-specific N content, and mass-specific C content; (ii) Rm conformed to a single Michaelis-Menten curve as a function of tissue water content; and (iii) the central parameters in the model were highly correlated with DNA content and critical enzyme activities. The model offers new insights and a more integrative scaling theory that quantifies the combined effects of tissue water content and body mass on respiratory metabolism during early plant ontogeny.

A regional-scale assessment of using metabolic scaling theory to predict ecosystem properties.

Metabolic scaling theory (MST) is one of ecology's most high-profile general models and can be used to link size distributions and productivity in forest systems. Much of MST's foundation is based on size distributions following a power law function with a scaling exponent of -2, a property assumed to be consistent in steady-state ecosystems. We tested the theory's generality by comparing actual size distributions with those predicted using MST parameters assumed to be general. We then used environmental variables and functional traits to explain deviation from theoretical expectations. Finally, we compared values of relative productivity predicted using MST with a remote-sensed measure of productivity. We found that fire-prone heath communities deviated from MST-predicted size distributions, whereas fire-sensitive rainforests largely agreed with the theory. Scaling exponents ranged from -1.4 to -5.3. Deviation from the power law assumption was best explained by specific leaf area, which varies along fire frequency and moisture gradients. While MST may hold in low-disturbance systems, we show that it cannot be applied under many environmental contexts. The theory should remain general, but understanding the factors driving deviation from MST and subsequent refinements is required if it is to be applied robustly across larger scales.

Current-year shoot hydraulic structure in two boreal conifers-implications of growth habit on water potential.

Metabolic scaling theory allows us to link plant hydraulic structure with metabolic rates in a quantitative framework. In this theoretical framework, we considered the hydraulic structure of current-year shoots in Pinus sylvestris and Picea abies, focusing on two properties unaccounted for by metabolic scaling theories: conifer needles are attached to the entire length of shoots, and the shoot as a terminal element does not display invariant properties. We measured shoot length and diameter as well as conduit diameter and density in two locations of 14 current-year non-leader shoots of pine and spruce saplings, and calculated conductivities of shoots from measured conduit properties. We evaluated scaling exponents for the hydraulic structure of shoots at the end of the water transport pathway from the data and applied the results to simulate water potential of shoots in the crown. Shoot shape was intermediate between cylindrical and paraboloid. Contrary to previous findings, we found that conduit diameter scaled with relative, not absolute, distance from the apex and absolute under-bark shoot diameter independently of species within the first-year shoots. Shoot hydraulic conductivity scaled with shoot diameter and hydraulic diameter. Larger shoots had higher hydraulic conductance. We further demonstrate by novel model calculations that ignoring foliage distribution along the hydraulic pathway overestimates water potential loss in shoots and branches and therefore overestimates related water stress effects. Scaling of hydraulic properties with shoot size enhances apical dominance and may contribute to the decline of whole-tree conductance in large trees.

Linking scaling laws across eukaryotes

Scaling laws relating body mass to species characteristics are among the most universal quantitative patterns in biology. Within major taxonomic groups, the 4 key ecological variables of metabolism, abundance, growth, and mortality are often well described by power laws with exponents near 3/4 or related to that value, a commonality often attributed to biophysical constraints on metabolism. However, metabolic scaling theories remain widely debated, and the links among the 4 variables have never been formally tested across the full domain of eukaryote life, to which prevailing theory applies. Here we present datasets of unprecedented scope to examine these 4 scaling laws across all eukaryotes and link them to test whether their combinations support theoretical expectations. We find that metabolism and abundance scale with body size in a remarkably reciprocal fashion, with exponents near ±3/4 within groups, as expected from metabolic theory, but with exponents near ±1 across all groups. This reciprocal scaling supports "energetic equivalence" across eukaryotes, which hypothesizes that the partitioning of energy in space across species does not vary significantly with body size. In contrast, growth and mortality rates scale similarly both within and across groups, with exponents of ±1/4. These findings are inconsistent with a metabolic basis for growth and mortality scaling across eukaryotes. We propose that rather than limiting growth, metabolism adjusts to the needs of growth within major groups, and that growth dynamics may offer a viable theoretical basis to biological scaling.

A global estimate of terrestrial net secondary production of primary consumers

AbstractAimNet secondary production (NSP) emerges from the consumption of net primary production (NPP) by all heterotrophic organisms. There has been sporadic interest in the importance of NSP, but no global estimates have been produced. Here, we examine NSP and attempt a global estimate using contemporary NPP data combined with modern metabolic scaling theory for consumption rates. We distinguish between potential NSP, as the amount of secondary production that could be supported by NPP, and realized NSP, as the amount remaining after anthropogenic habitat disruption.LocationGlobal.Time period2000–2014.MethodsWe present a metabolic model of NSP, implementing a type II functional response for consumption rates, wherein search efficiency and handling time are calculated based on consumer mass and ambient temperature. We solve this model for each 0.05‐decimal‐degree pixel in the global terrestrial biosphere, using as data inputs NPP (MOD17A3) and land‐surface temperature (MOD11C3). We aggregate estimates within global land‐cover classifications (MCD12C1) to obtain cover‐specific and global estimates of NSP. We also correct our estimates based on declines in consumer abundance reported in Living planet report 2014.ResultsWe estimate that potential NSP is 4.74 Pg C/year globally (95% CI: 3.75–5.75). When we correct for global consumer population declines, realized NSP is estimated at 2.37 Pg C/year (95% CI: 1.86–2.89), a loss of 50% in the rate of carbon flux through secondary consumers.Main conclusionsOur estimates should not be viewed as the last word on NSP but are sufficient to suggest that the flux of carbon through consumers is of a similar magnitude to many other fluxes crucial to the global carbon cycle. We view this as a hypothesis to be tested that suggests NSP deserves significantly more attention in Earth systems, macroecology and biogeochemical research.

A metabolic scaling theory-driven remote sensing approach to map spatiotemporal dynamics of litterfall in a tropical montane cloud forest

Litter production, or litterfall, is a predominant part of the carbon and nutrient cycles of forest ecosystems. As a major and relatively invariable aspect of primary production, litterfall could be a predictor of total plant production. However, the conventional field approach for its estimation is laborious and costly. High temporal resolution optical satellite imagery may strengthen our ability to estimate litterfall over a vast region, but this remote sensing method is limited in some tropical and near-tropical mountainous regions, mainly due to coarse spatial resolution and frequent cloud coverage. The metabolic scaling theory (MST) states that forest productivity (including litterfall) is directly proportional to the mass of the photosynthesis tissue (leaves), and spatially, it may be isometrically scaled using the size of the largest individual in the system. In this study, we investigated the temporal dimension of the MST and hypothesized that the maximum leaf abundance over a year could also be a key determinant of annual litter production. The relationship between the leaf mass during the peak growing season and annual litterfall could facilitate large-scale litterfall mapping in mountainous terrains, since vegetation indices such as the Enhanced Vegetation Index (EVI) or the Normalized Difference Vegetation Index (NDVI), commonly utilized as a surrogate for leaf abundance, can be derived from relatively high spatial resolution satellite imagery. We correlated the summer growing season Landsat Enhanced Thematic Mapper plus (ETM+) EVI and NDVI for 2016 and 2017 and the temporally corresponding annual accumulated field litterfall data acquired from hinoki (Chamaecyparis spp.) dominant tropical montane cloud forests in northeastern Taiwan (23.98 N, 120.97 E). We found that the summer growing season EVI and NDVI may be salient variables to explain the spatial variation of annual litter production ( r2 = 0.39–0.67, p ≤ 0.01). The results have profound implications for applying MST to the regional mapping of ecosystem production across space and time in humid tropical regions, and these implications may facilitate the remote assessment of the terrestrial carbon budget.

Asymmetric Competition Can Shape the Size Distribution of Trees in a Natural Tropical Forest

AbstractThe architecture (here, the size distribution combined with the spatial pattern of individuals) of natural forest at demographic equilibrium can be used to infer the demographic processes that drive the forest dynamics. In particular, a constant growth rate and a constant mortality rate for all trees would generate an exponential distribution of their size, whereas the metabolic scaling theory predicts a power distribution. In an undisturbed tropical rainforest in French Guiana, the diameter distribution was significantly steeper than the best-fit exponential distribution and significantly flatter than the best-fit power distribution. A simple individual-based model of forest dynamics with asymmetric competition between trees, where the strength of competition was regulated by a single parameter, was able to predict the observed distribution. Competition drove the forest into a self-organized state with stronger inequalities of size among trees, a lower mean competition index, and a spatial pattern of trees that deviated from complete spatial randomness.

Mapping Maximum Tree Height of the Great Khingan Mountain, Inner Mongolia Using the Allometric Scaling and Resource Limitations Model

Maximum tree height is an important indicator of forest vegetation in understanding the properties of plant communities. In this paper, we estimated regional maximum tree heights across the forest of the Great Khingan Mountain in Inner Mongolia with the allometric scaling and resource limitations model. The model integrates metabolic scaling theory and the water–energy balance equation (Penman–Monteith equation) to predict maximum tree height constrained by local resource availability. Monthly climate data, including precipitation, wind speed, vapor pressure, air temperature, and solar radiation are inputs of this model. Ground measurements, such as tree heights, diameters at breast height, and crown heights, have been used to compute the parameters of the model. In addition, Geoscience Laser Altimeter System (GLAS) data is used to verify the results of model prediction. We found that the prediction of regional maximum tree heights is highly correlated with the GLAS tree heights (R2 = 0.64, RMSE = 2.87 m, MPSE = 12.45%). All trees are between 10 to 40 m in height, and trees in the north are taller than those in the south of the region of research. Furthermore, we analyzed the sensitivity of the input variables and found the model predictions are most sensitive to air temperature and vapor pressure.

Equilibrium forest demography explains the distribution of tree sizes across North America

The distribution of tree sizes within a forest strongly influences how it will respond to disturbances and environmental changes such as future climate change and increases in atmospheric CO2. This means that global vegetation models must include variation in tree size to accurately represent carbon sinks, such as that seen in North America. Here we use an analytical model of large-scale forest demography which assumes tree growth varies as a power of tree diameter whilst tree mortality is independent of size. The equilibrium solutions of this model are able to accurately reproduce the tree-size distributions, for 61 species and four plant functional types, measured across North America, using just a single species-specific fitting parameter, μ, which determines the ratio of mortality to growth. The predictions of metabolic scaling theory for tree-size distributions are also tested and found to deviate significantly from observations and that maybe explained by the assumptions made about how individual trees fill the available space. We show that equilibrium forest demography implies a single curve that relates mean tree diameter to μ, and that this can be used to make reasonable estimates of the whole dataset mean trunk diameter by fitting only to the larger trees. Our analysis suggests that analytical solutions such as those in this paper may have a role in aiding the understanding and development of next-generation Dynamic Global Vegetation Models based on ecosystem demography.

Traits drive global wood decomposition rates more than climate.

Wood decomposition is a major component of the global carbon cycle. Decomposition rates vary across climate gradients, which is thought to reflect the effects of temperature and moisture on the metabolic kinetics of decomposers. However, decomposition rates also vary with wood traits, which may reflect the influence of stoichiometry on decomposer metabolism as well as geometry relating the surface areas that decomposers colonize with the volumes they consume. In this paper, we combined metabolic and geometric scaling theories to formalize hypotheses regarding the drivers of wood decomposition rates, and assessed these hypotheses using a global compilation of data on climate, wood traits, and wood decomposition rates. Our results are consistent with predictions from both metabolic and geometric scaling theories. Approximately half of the global variation in decomposition rates was explained by wood traits (nitrogen content and diameter), whereas only a fifth was explained by climate variables (air temperature, precipitation, and relative humidity). These results indicate that global variation in wood decomposition rates is best explained by stoichiometric and geometric wood traits. Our findings suggest that inclusion of wood traits in global carbon cycle models can improve predictions of carbon fluxes from wood decomposition.

Linking phytoplankton community metabolism to the individual size distribution.

Quantifying variation in ecosystem metabolism is critical to predicting the impacts of environmental change on the carbon cycle. We used a metabolic scaling framework to investigate how body size and temperature influence phytoplankton community metabolism. We tested this framework using phytoplankton sampled from an outdoor mesocosm experiment, where communities had been either experimentally warmed (+ 4 °C) for 10 years or left at ambient temperature. Warmed and ambient phytoplankton communities differed substantially in their taxonomic composition and size structure. Despite this, the response of primary production and community respiration to long‐ and short‐term warming could be estimated using a model that accounted for the size‐ and temperature dependence of individual metabolism, and the community abundance‐body size distribution. This work demonstrates that the key metabolic fluxes that determine the carbon balance of planktonic ecosystems can be approximated using metabolic scaling theory, with knowledge of the individual size distribution and environmental temperature.

Will forest size structure follow the −2 power-law distribution under ideal demographic equilibrium state?

Scaling relations formed in forest development processes are fairly important for understanding and predicting forest dynamics. During self-thinning of a relatively even-sized forest, tree abundance will decrease with an increase in average tree size, forming the size-abundance relation (SAR); while for a size-structured forest under the demographic equilibrium state, the frequency of trees also varies with size classes in a similar, decreasing pattern, manifesting as the size-frequency distribution (SFD). In the metabolic scaling theory (MST), the two scaling relations are considered to be consistent. However, in this paper, we proved that SFD can never be equivalent to SAR unless the growth rate of tree diameters is a constant. The reason derives from the time differences of transition between different size classes, which are influenced in SFD maintenance but not in SAR formation. Demographic equilibrium of a size structured forest requires a different resource allocation among different size classes at the same time, which contradicts the resource conservation during SAR formation in the self-thinning process. Consequently, if the rate of resource use per individual scales as a +2 power with its diameter according to MST, which led to the −2 power SAR, SFD can never be a −2 power-law distribution. The previous confusion between SFD of a size-structured forest and SAR formed during self-thinning processes may lead to many misunderstandings and unreliable predictions on forest structure and dynamics.

Adaptive diversification of growth allometry in the plant Arabidopsis thaliana

Seed plants vary tremendously in size and morphology; however, variation and covariation in plant traits may be governed, at least in part, by universal biophysical laws and biological constants. Metabolic scaling theory (MST) posits that whole-organismal metabolism and growth rate are under stabilizing selection that minimizes the scaling of hydrodynamic resistance and maximizes the scaling of resource uptake. This constrains variation in physiological traits and in the rate of biomass accumulation, so that they can be expressed as mathematical functions of plant size with near-constant allometric scaling exponents across species. However, the observed variation in scaling exponents calls into question the evolutionary drivers and the universality of allometric equations. We have measured growth scaling and fitness traits of 451 Arabidopsis thaliana accessions with sequenced genomes. Variation among accessions around the scaling exponent predicted by MST was correlated with relative growth rate, seed production, and stress resistance. Genomic analyses indicate that growth allometry is affected by many genes associated with local climate and abiotic stress response. The gene with the strongest effect, PUB4, has molecular signatures of balancing selection, suggesting that intraspecific variation in growth scaling is maintained by opposing selection on the trade-off between seed production and abiotic stress resistance. Our findings suggest that variation in allometry contributes to local adaptation to contrasting environments. Our results help reconcile past debates on the origin of allometric scaling in biology and begin to link adaptive variation in allometric scaling to specific genes.

Diverse scaling relationships of tree height and diameter in five tree species

ABSTRACTBackground: Allometric scaling relationships are important in ecology and forestry. The metabolic scaling theory (MST) predicts a universal invariant scaling relationship for tree growth, relating height and diameter to each other.Aims: Data on five tree species (Pinus taeda L., Pinus virginiana Mill., Liquidambar styraciflua L., Liriodendron tulipifera L., and Pinus palustris Mill.) were used to test the predictions from MST on the scaling of height–diameter and also diameter growth.Methods: Data on tree height and diameter for five tree species from both natural forests and plantations were collected to study two types of scaling relationships: tree height–diameter and stem diameter growth. A reduced major axis of regression analysis model type II was used to determine scaling exponents from each species under different environmental conditions.Results: No universal invariant scaling exponent was found in height–diameter and diameter growth for the five species. The scaling varied across natural forests, plantations and scales (e.g., time and number of measured trees). However, in some situations the scaling exponents failed to show significant difference with the predicted values (e.g., 2/3 or 1).Conclusions: Diverse scaling exponents were observed for the five tree species with the scaling relationships varying with environmental settings.