Optimal Carbon Storage During Drought.

Allocation of non-structural carbohydrates (NSC) to storage allows plants to maintain a carbon pool in anticipation of future stress. However, to do so, plants must forego use of the carbon for growth, creating a trade-off between storage and growth. It is possible that plants actively regulate the storage pool to maximise fitness in a stress-prone environment.

Diel- and temperature-driven variation of leaf dark respiration rates and metabolite levels in rice.

Leaf respiration in the dark (R ) is often measured at a single time during the day, with hot-acclimation lowering R at a common measuring temperature. However, it is unclear whether the diel cycle influences the extent of thermal acclimation of R , or how temperature and time of day interact to influence respiratory metabolites. To examine these issues, we grew rice under 25°C : 20°C, 30°C : 25°C and 40°C : 35°C day : night cycles, measuring R and changes in metabolites at five time points spanning a single 24-h period.

Molecular and physiological responses during thermal acclimation of leaf photosynthesis and respiration in rice.

To further our understanding of how sustained changes in temperature affect the carbon economy of rice (Oryza sativa), hydroponically grown plants of the IR64 cultivar were developed at 30°C/25°C (day/night) before being shifted to 25/20°C or 40/35°C. Leaf messenger RNA and protein abundance, sugar and starch concentrations, and gas-exchange and elongation rates were measured on preexisting leaves (PE) already developed at 30/25°C or leaves newly developed (ND) subsequent to temperature transfer. Following a shift in growth temperature, there was a transient adjustment in metabolic gene transcript abundance of PE leaves before homoeostasis was reached within 24 hr, aligning with R (leaf dark respiratory CO release) and A (net CO assimilation) changes.

Drought-related tree mortality: addressing the gaps in understanding and prediction.

Increased tree mortality during and after drought has become a research focus in recent years. This focus has been driven by: the realisation that drought-related tree mortality is more widespread than previously thought; the predicted increase in the frequency of climate extremes this century; and the recognition that current vegetation models do not predict drought-related tree mortality and forest dieback well despite the large potential effects of these processes on species composition and biogeochemical cycling. To date, the emphasis has been on understanding the causal mechanisms of drought-related tree mortality, and on mechanistic models of plant function and vegetation dynamics, but a consensus on those mechanisms has yet to emerge.

Statistical patterns in tropical tree cover explained by the different water demand of individual trees and grasses.

Tree cover varies enormously across tropical ecosystems-from arid savannas to closed rain forests-and yet a general predictive theory of tropical tree cover remains elusive. Here we use the maximum-entropy method to predict the most likely sample frequency distribution of ecosystems with different tree and grass fractional cover if balance between water supply and demand were the dominant constraint on community assembly. Assuming a hierarchy of individual plant water demand in which trees require more water than grasses, we reproduce observed trends in the means and the upper and lower limits of tropical tree and grass cover across the entire spectrum of tropical ecosystem water supply.

New insights into carbon allocation by trees from the hypothesis that annual wood production is maximized.

Allocation of carbon (C) between tree components (leaves, fine roots and woody structures) is an important determinant of terrestrial C sequestration. Yet, because the mechanisms underlying C allocation are poorly understood, it is a weak link in current earth-system models. We obtain new theoretical insights into C allocation from the hypothesis (MaxW) that annual wood production is maximized.

Plant root distributions and nitrogen uptake predicted by a hypothesis of optimal root foraging.

CO(2)-enrichment experiments consistently show that rooting depth increases when trees are grown at elevated CO(2) (eCO(2)), leading in some experiments to increased capture of available soil nitrogen (N) from deeper soil. However, the link between N uptake and root distributions remains poorly represented in forest ecosystem and global land-surface models. Here, this link is modeled and analyzed using a new optimization hypothesis (MaxNup) for root foraging in relation to the spatial variability of soil N, according to which a given total root mass is distributed vertically in order to maximize annual N uptake.

Why does leaf nitrogen decline within tree canopies less rapidly than light? An explanation from optimization subject to a lower bound on leaf mass per area.

A long-established theoretical result states that, for a given total canopy nitrogen (N) content, canopy photosynthesis is maximized when the within-canopy gradient in leaf N per unit area (N(a)) is equal to the light gradient. However, it is widely observed that N(a) declines less rapidly than light in real plant canopies. Here we show that this general observation can be explained by optimal leaf acclimation to light subject to a lower-bound constraint on the leaf mass per area (m(a)).

Modeling carbon allocation in trees: a search for principles.

We review approaches to predicting carbon and nitrogen allocation in forest models in terms of their underlying assumptions and their resulting strengths and limitations. Empirical and allometric methods are easily developed and computationally efficient, but lack the power of evolution-based approaches to explain and predict multifaceted effects of environmental variability and climate change. In evolution-based methods, allocation is usually determined by maximization of a fitness proxy, either in a fixed environment, which we call optimal response (OR) models, or including the feedback of an individual's strategy on its environment (game-theoretical optimization, GTO).

Predictions of single-nucleotide polymorphism differentiation between two populations in terms of mutual information.

Mutual information (I) provides a robust measure of genetic differentiation for the purposes of estimating dispersal between populations. At present, however, there is little predictive theory for I. The growing importance in population biology of analyses of single-nucleotide and other single-feature polymorphisms (SFPs) is a potent reason for developing an analytic theory for I with respect to a single locus.

Leaf-trait variation explained by the hypothesis that plants maximize their canopy carbon export over the lifespan of leaves.

Measured values of four key leaf traits (leaf area per unit mass, nitrogen concentration, photosynthetic capacity, leaf lifespan) co-vary consistently within and among diverse biomes, suggesting convergent evolution across species. The same leaf traits co-vary consistently with the environmental conditions (light intensity, carbon-dioxide concentration, nitrogen supply) prevailing during leaf development. No existing theory satisfactorily explains all of these trends.

Maximum entropy production and plant optimization theories.

Plant ecologists have proposed a variety of optimization theories to explain the adaptive behaviour and evolution of plants from the perspective of natural selection ('survival of the fittest'). Optimization theories identify some objective function--such as shoot or canopy photosynthesis, or growth rate--which is maximized with respect to one or more plant functional traits. However, the link between these objective functions and individual plant fitness is seldom quantified and there remains some uncertainty about the most appropriate choice of objective function to use.

Why is plant-growth response to elevated CO amplified when water is limiting, but reduced when nitrogen is limiting? A growth-optimisation hypothesis.

Experimental evidence indicates that the stomatal conductance and nitrogen concentration ([N]) of foliage decline under CO enrichment, and that the percentage growth response to elevated CO is amplified under water limitation, but reduced under nitrogen limitation. We advance simple explanations for these responses based on an optimisation hypothesis applied to a simple model of the annual carbon-nitrogen-water economy of trees growing at a CO-enrichment experiment at Oak Ridge, Tennessee, USA. The model is shown to have an optimum for leaf [N], stomatal conductance and leaf area index (LAI), where annual plant productivity is maximised.

Statistical mechanics unifies different ecological patterns.

Recently there has been growing interest in the use of maximum relative entropy (MaxREnt) as a tool for statistical inference in ecology. In contrast, here we propose MaxREnt as a tool for applying statistical mechanics to ecology. We use MaxREnt to explain and predict species abundance patterns in ecological communities in terms of the most probable behaviour under given environmental constraints, in the same way that statistical mechanics explains and predicts the behaviour of thermodynamic systems.

Sustainable stemwood yield in relation to the nitrogen balance of forest plantations: a model analysis.

We used an existing analytical model of stemwood growth in relation to nitrogen supply, which we describe in an accompanying paper, to examine the long-term effects of harvesting and fire on tree growth. Our analysis takes into account the balance between nitrogen additions from deposition, fixation, and fertilizer applications, and nitrogen losses from stemwood harvesting, regeneration burning, leaching and gaseous emissions. Using a plausible set of parameter values for Eucalyptus, we conclude that nitrogen loss through fire is the main factor limiting sustainable yield, defined as the maximum mean annual stemwood volume increment obtained in the steady state, if management practices are continued indefinitely.

Analytical model of stemwood growth in relation to nitrogen supply.

We derived a simplified version of a previously published process-based model of forest productivity and used it to gain information about the dependence of stemwood growth on nitrogen supply. The simplifications we made led to the following general expression for stemwood carbon (c(w)) as a function of stand age (t), which shows explicitly the main factors involved: c(w)(t) = eta(w)G*/ micro (w)(1 - lambdae(- micro (w)t) - micro (w)e(-lambdat)/lambda - micro (w)), where eta(w) is the fraction of total carbon production (G) allocated to stemwood, G* is the equilibrium value of G at canopy closure, lambda describes the rate at which G approaches G*, and micro (w) is the combined specific rate of stemwood maintenance respiration and senescence. According to this equation, which describes a sigmoidal growth curve, c(w) is zero initially and asymptotically approaches eta(w)G*/ micro (w) with the rate of approach dependent on lambda and micro (w).