Ecosystem photosynthesis in land-surface models: a first-principles approach

Vegetation regulates land-atmosphere water and energy exchanges and is an essential component of land-surface models (LSMs). However, LSMs have been handicapped by assumptions that equate acclimated photosynthetic responses to environment with fast responses observable in the laboratory. These time scales can be distinguished by including specific representations of acclimation, but at the cost of further increasing parameter requirements. Here we develop an alternative approach based on optimality principles that predict the acclimation of carboxylation and electron-transport capacities, and a variable controlling the response of leaf-level carbon dioxide drawdown to vapour pressure deficit (VPD), to variations in growth conditions on a weekly to monthly time scale. In the "P model", an optimality-based light-use efficiency model for gross primary production (GPP) on this time scale, these acclimated responses are implicit. Here they are made explicit, allowing fast and slow response time-scales to be separated and GPP to be simulated at sub-daily timesteps. The resulting model mimics diurnal cycles of GPP recorded by eddy-covariance flux towers in a temperate grassland and boreal, temperate and tropical forests, with no parameter changes between biomes. Best performance is achieved when biochemical capacities are adjusted to match recent midday conditions. This model suggests a simple and parameter-sparse method to include both instantaneous and acclimated responses within an LSM framework, with many potential applications in weather, climate and carbon - cycle modelling. Plain Language SummaryVegetation regulates the exchanges of energy, water and carbon dioxide between the land and the atmosphere. Numerical climate models represent these processes, focusing mainly on their rapid variations in response to changes in the environment (including temperature and light) on timescales of seconds to hours. However, plants also adjust their physiology to environmental changes over longer periods within the season. Here we have adapted a simple model that formulates plant behaviour in terms of optimal trade-offs between different processes, so it simulates processes on both time scales. This model correctly reproduces the daily cycle of carbon dioxide uptake by plants, as recorded in different kinds of vegetation. We show that plants optimize their behaviour for midday conditions, when the light is greatest, and adjust to longer-term environmental variations on a timescale of about a week to a month. The model conveniently avoids the need to give specific, fixed values to physiological variables (such as photosynthetic capacity) for different types of plants. The optimality assumptions mean that the model gives equally good results in tropical, temperate and boreal forests, and in grasslands, using the same equations, and a very small number of input variables that are constant across the world. Key PointsO_LIOptimality theory is used to develop a simple model incorporating fast and acclimated responses of photosynthesis and stomatal conductance C_LIO_LIBiogeochemical photosynthetic capacities adjust to midday light conditions C_LIO_LIThe new model simulates gross primary production on sub-daily timesteps across a range of different vegetation types and climate C_LI