Empirical evidence and theoretical understanding of ecosystem carbon and nitrogen cycle interactions.

Interactions between carbon (C) and nitrogen (N) cycles in terrestrial ecosystems are simulated in advanced vegetation models, yet methodologies vary widely, leading to divergent simulations of past land C balance trends. This underscores the need to reassess our understanding of ecosystem processes, given recent theoretical advancements and empirical data. We review current knowledge, emphasising evidence from experiments and trait data compilations for vegetation responses to CO2 and N input, alongside theoretical and ecological principles for modelling. N fertilisation increases leaf N content but inconsistently enhances leaf-level photosynthetic capacity. Whole-plant responses include increased leaf area and biomass, with reduced root allocation and increased aboveground biomass. Elevated atmospheric CO2 also boosts leaf area and biomass but intensifies belowground allocation, depleting soil N and likely reducing N losses. Global leaf traits data confirm these findings, indicating that soil N availability influences leaf N content more than photosynthetic capacity. A demonstration model based on the functional balance hypothesis accurately predicts responses to N and CO2 fertilisation on tissue allocation, growth and biomass, offering a path to reduce uncertainty in global C cycle projections.

Eco-evolutionary optimality as a means to improve vegetation and land-surface models.

Global vegetation and land-surface models embody interdisciplinary scientific understanding of the behaviour of plants and ecosystems, and are indispensable to project the impacts of environmental change on vegetation and the interactions between vegetation and climate. However, systematic errors and persistently large differences among carbon and water cycle projections by different models highlight the limitations of current process formulations. In this review, focusing on core plant functions in the terrestrial carbon and water cycles, we show how unifying hypotheses derived from eco-evolutionary optimality (EEO) principles can provide novel, parameter-sparse representations of plant and vegetation processes. We present case studies that demonstrate how EEO generates parsimonious representations of core, leaf-level processes that are individually testable and supported by evidence. EEO approaches to photosynthesis and primary production, dark respiration and stomatal behaviour are ripe for implementation in global models. EEO approaches to other important traits, including the leaf economics spectrum and applications of EEO at the community level are active research areas. Independently tested modules emerging from EEO studies could profitably be integrated into modelling frameworks that account for the multiple time scales on which plants and plant communities adjust to environmental change.

Forests buffer against variations in precipitation.

Atmospheric moisture recycling effectively increases the amount of usable water over land as the water can undergo multiple precipitation-evapotranspiration cycles. Differences in land cover and climate regulate the evapotranspiration flux. Forests can have deep roots that access groundwater facilitating transpiration throughout the dry season independent of precipitation. This stable transpiration buffers the forest against precipitation variability. However, it is not known whether the buffering effect, already modeled for tropical forests, is common to all forests globally. Here we apply a state-of-the-art Lagrangian moisture tracking model (UTrack) to study whether forest land cover in the upwind precipitationshed can lead to a reduction in monthly precipitation variability downwind. We found a significant buffering effect of forests in the precipitation variability of 10 out of 14 biomes globally. On average, if 50% of precipitation originates from forest, then we find a reduction in the coefficient of variation of monthly precipitation of 60%. We also observed that a high fraction of precipitation from non-forest land sources tends to have the opposite effect, that is, no buffering effect. The average variation of monthly precipitation was 69% higher in areas where 50% of precipitation originates from non-forest land sources in the precipitationshed. Our results emphasize the importance of land cover composition in the precipitationshed to buffer precipitation variability downwind, in particular forest cover. Understanding the influence of land cover in a precipitationshed on atmospheric moisture transport is key for evaluating an area's water-climate regulatory ecosystem services and may become increasingly important due to continued changes in land cover and climate change.

Organizing principles for vegetation dynamics.

Plants and vegetation play a critical-but largely unpredictable-role in global environmental changes due to the multitude of contributing processes at widely different spatial and temporal scales. In this Perspective, we explore approaches to master this complexity and improve our ability to predict vegetation dynamics by explicitly taking account of principles that constrain plant and ecosystem behaviour: natural selection, self-organization and entropy maximization. These ideas are increasingly being used in vegetation models, but we argue that their full potential has yet to be realized. We demonstrate the power of natural selection-based optimality principles to predict photosynthetic and carbon allocation responses to multiple environmental drivers, as well as how individual plasticity leads to the predictable self-organization of forest canopies. We show how models of natural selection acting on a few key traits can generate realistic plant communities and how entropy maximization can identify the most probable outcomes of community dynamics in space- and time-varying environments. Finally, we present a roadmap indicating how these principles could be combined in a new generation of models with stronger theoretical foundations and an improved capacity to predict complex vegetation responses to environmental change.