No abstract

The worldwide leaf economics spectrum relates leaf lifespan (LL) to leaf dry mass per unit area (LMA). By combining three well-supported principles, we show that an isometric relationship between these two quantities maximizes the leaf net carbon gain. This theory predicts a spectrum of equally competent LMA-LL combinations in any given environment, and how their optimal ratio varies across environments. By analysing two large, independent leaf-trait datasets for woody species, we provide quantitative empirical support for the predicted dependencies of LL on LMA and environment in evergreen plants, and for the distinct predicted dependencies of LMA on light, temperature, growing season length and aridity in evergreen and deciduous plants. We thereby resolve the long standing question of why deciduous LMA tends to increase (with increasing LL) towards the equator, while evergreen LMA and LL decrease. We also show how the statistical distribution of LMA within communities can be modelled as an outcome of environmental selection on the global pool of species with diverse values of LMA and LL.

The life span of leaves increases with their mass per unit area (LMA). It is unclear why. Here, we show that this empirical generalization (the foundation of the worldwide leaf economics spectrum) is a consequence of natural selection, maximizing average net carbon gain over the leaf life cycle. Analyzing two large leaf trait datasets, we show that evergreen and deciduous species with diverse construction costs (assumed proportional to LMA) are selected by light, temperature, and growing-season length in different, but predictable, ways. We quantitatively explain the observed divergent latitudinal trends in evergreen and deciduous LMA and show how local distributions of LMA arise by selection under different environmental conditions acting on the species pool. These results illustrate how optimality principles can underpin a new theory for plant geography and terrestrial carbon dynamics.

Climate exerts a major influence on crop development and yield. Despite extensive modelling efforts, there is still considerable uncertainty about the consequences of a changing climate for the yields of major crops. Existing crop models are complex and rely on many assumptions and parameters, motivating a quest for more parsimonious models with stronger theoretical and empirical foundations. This paper presents a prototype of such a model for wheat, informed by measurements of gross primary production (GPP), biomass and yield at research sites across the wheat-growing regions of China. First, GPP was predicted using a recently developed first-principles model driven only by climate, carbon dioxide (CO 2 ) concentration, and light absorbed by leaves. Modelled GPP was shown to agree well with eddy-covariance measurements. Second, the data were used to show that above-ground biomass (AB) is proportional to time-integrated GPP, and that grain yield shows a saturating relationship with AB. Simple empirical equations based on these findings were combined with modelled GPP to predict yield, including propagated errors due to parameter uncertainty in both the GPP model and the empirical equations. The resulting 'hybrid' model, applied in a variety of climates, successfully predicted measured interannual variations in AB and yield. Third, the model was extended to include a phenology scheme, a mass-balance equation relating mean leaf area index to accumulated GPP over the growth phase, and an independently observed response of leaf massper-area to CO 2 . Sensitivity analyses and scenario runs with this extended model showed a positive but saturating (at ~600 ppm) response of yield to rising CO 2 , consistent with experimental evidence. This positive effect was partially counteracted by a net negative response of yield to increasing temperature, caused by increasing photorespiration and an accelerated growth cycle.

Evaluation of potential crop yields is important for global food security assessment because it represents the biophysical ‘ceiling’ determined by variety, climate and ambient CO2. Statistical approaches have limitations when assessing future potential yields, while large differences between results obtained using process-based models reflect uncertainties in model parameterisations. Here we simulate the potential yield of wheat across the present-day wheat-growing areas, using a new global model that couples a parameter-sparse, optimality-based representation of gross primary production (GPP) to empirical functions relating GPP, biomass production and yield. The model reconciles the transparency and parsimony of statistical models with a mechanistic grounding in the standard model of C3 photosynthesis, and seamlessly integrates photosynthetic acclimation and CO2 fertilization effects. The model accurately predicted the CO2 response observed in FACE experiments, and captured the magnitude and spatial pattern of EARTHSTAT ‘attainable yield’ data in 2000 CE better than process-based models in ISIMIP. Global simulations of potential yield during 1981–2016 were analysed in parallel with global historical data on actual yield, in order to test the hypothesis that environmental effects on modelled potential yields would also be shown in observed actual yields. Higher temperatures are thereby shown to have negatively affected (potential and actual) yields over much of the world. Greater solar radiation is associated with higher yields in humid regions, but lower yields in semi-arid regions. Greater precipitation is associated with higher yields in semi-arid regions. The effect of rising CO2 is reflected in increasing actual yield, but trends in actual yield are stronger than the CO2 effect in many regions, presumably because they also include effects of crop breeding and improved management. We present this hybrid modelling approach as a useful addition to the toolkit for assessing global environmental change impacts on the growth and yield of arable crops.