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Prentice, I. C.

Publications and source records attributed to Prentice, I. C..

17 recordsLinked to original sources

An eco-evolutionary approach to defining wildfire regimes

Fuel availability and fuel dryness are consistently shown to be the primary drivers of wildfire intensity and burnt area. Here we hypothesise that differences in the timing of fuel build up and drying determine the optimal time for wildfire occurrence. We use gross primary production (GPP) as a measure of biomass production and hence fuel availability, and vapour pressure deficit (VPD) as a measure of fuel drying. We use the phase difference in the seasonal time course and magnitude of GPP and VPD to cluster regions that should therefore have distinct wildfire behaviour. We then show that each of the resultant clusters is distinctive in terms of one or more fire properties, specifically number of ignitions, burnt area, size, speed, duration, intensity, and length of the wildfire season. The emergence of distinct regimes as a function of two biophysical drivers reflects the fact that both vegetation and wildfire properties are a consequence of eco-evolutionary adaptions to environmental conditions. We then examine the degree to which human activities or vegetation properties modify these fire regimes within each of these clusters. Variability in GPP and VPD largely explains the within-cluster variation in fire properties. The type of vegetation cover has an influence on burnt area and carbon emissions in particular, while human activities are more important for fire properties such as size, rate of spread and duration largely through their influence of landscape fragmentation. Although both human activities and vegetation properties modify wildfire regimes, the ability to distinguish wildfire regimes using GPP and VPD alone emphasizes that land management, fire use and fire suppression are constrained by environmental conditions. This eco-evolutionary optimality approach to characterising wildfire regimes provides a basis for designing a simple fire model for Earth System modelling.

ecology↗

CN-model: A dynamic model for the coupled carbon and nitrogen cycles in terrestrial ecosystems

Reducing uncertainty in carbon cycle projections relies on reliable representations of interactions between the carbon and nutrient cycles. Here, we build on a set of principles and hypotheses, funded in established theoretical understanding and supported by empirical evidence, to formulate and implement a dynamic model of carbon-nitrogen cycle (C-N) interactions in terrestrial ecosystems. The model combines a representation of photosynthetic acclimation to the atmospheric environment with implications for canopy and leaf N, a representation of functional balance for modelling C allocation and growth different tissues, and a representation of plant N uptake based on a first principles-informed, simplified description of solute transport in the soil. Here, we provide a comprehensive description of the model and its implementation and demonstrate the stability and functionality of the model for simulating seasonal variations of ecosystem C and N fluxes and pools. This may provide a way forward for representing key ecosystem processes related to C and N cycling based on established theoretical concepts with strong empirical support.

ecology↗

Why models underestimate tropical forest productivity: a case study in West Africa

Tropical forests dominate terrestrial photosynthesis, yet there are major contradictions in our understanding due to a lack of field studies, especially outside the tropical Americas. A recent field study indicated that West African forests have among the highest forests gross primary productivity (GPP) yet observed, contradicting models that rank them lower than Amazonian forests. Here, we explore possible reasons for this data-model mismatch. We found the in situ GPP measurements higher than multiple global GPP products at the studied sites in Ghana. The underestimation of GPP by models largely disappears when a standard photosynthesis model is informed by local field-measured values of (a) fractional absorbed photosynthetic radiation (fAPAR), and (b) photosynthetic traits. Satellites systematically underestimate fAPAR in the tropics due to cloud contamination issues. The study highlights the potential widespread underestimation of tropical forests GPP and carbon cycling and hints at the ways forward for model and input data improvement. Related manuscriptThe recent field study mentioned above is a manuscript currently accepted by Nature Communications (manuscript id NCOMMS-23-37419), which is available as a preprint https://www.researchsquare.com/article/rs-3136892/v1 Codes and data availabilityAll data and codes underlying the study are currently shared via Github (link here) which will be made available through Zenodo upon acceptance.

ecology↗

Global variation in the ratio of sapwood to leaf area explained by optimality principles

The sapwood area supporting a given leaf area (vH) reflects a coordinated coupling between carbon uptake, water transport and loss at a whole plant level. Worldwide variation in vH reflects diverse plants strategies adapt to prevailing environments, and impact the evolution of global carbon and water cycles. Why such a variation has not been convincingly explained yet, thus hinder its representation in Earth System Models. Here we hypothesize that optimal vH tends to mediate between plant hydraulics and leaf photosynthesis so that leaf water loss matches water supply. By compiling and testing against two extensive datasets, we show that our hypothesis explains nearly 60% of vH variation responding to light, vapour pressure deficit, temperature, and sapwood conductance in a quantitively predictable manner. Sapwood conductance or warming-enhanced hydraulic efficiency reduces the demand on sapwood area for a given total leaf area and, whereas brightening and air dryness enhance photosynthetic capacities consequently increasing the demand. This knowledge can enrich Earth System Models where carbon allocation and water hydraulics play key roles in predicting future climate-carbon feedback.

ecology↗

Temperature effects on the global patterns of photosynthetic quantum efficiency

SummaryO_LIHistorically, terrestrial biosphere models (TBMs) have assigned the intrinsic (maximum) quantum yield of photosynthesis ({varphi}0) a constant value for each plant functional type. However, experimental studies have shown that {varphi}0- when measured on light-adapted leaves - depends on temperature. It is unclear whether this dependence is universal or biome-specific; how it is manifested at the ecosystem level; and how it should be represented in TBMs. C_LIO_LIBy fitting empirical light-response curves to a global set of eddy-covariance CO2 flux measurements and correcting for photorespiration, we inferred apparent, ecosystem-level {varphi}0values and their temperature responses across a wide range of environments. C_LIO_LIThe temperature response of apparent ecosystem-level {varphi}0 follows a universal bell-shaped curve. The shape of this curve does not markedly differ among biomes, but the maximum value of {varphi}0 decreases with increasing aridity, its temperature optimum increases with increasing growth temperature, and its sensitivity to temperature increases as growth temperature declines. C_LIO_LIOur model for {varphi}0(T) aligns with recent theory highlighting the role of cytochrome b6f in regulating the light reactions of photosynthesis. If implemented in TBMs, this model should allow better predictions of the responses of terrestrial ecosystem function to a warming climate. C_LI

ecology↗

Incorporating photosynthetic acclimation improves stomatal optimisation models

Stomatal opening in plant leaves is regulated through a balance of carbon and water exchange under different environmental conditions. Accurate estimation of stomatal regulation is crucial for understanding how plants respond to changing environmental conditions, particularly under climate change. A new generation of optimality-based modelling schemes determines instantaneous stomatal responses from a balance of trade-offs between carbon gains and hydraulic costs, but most such schemes do not account for biochemical acclimation in response to drought. Here, we compare the performance of seven instantaneous stomatal optimisation models with and without accounting for photosynthetic acclimation. Using experimental data from 38 plant species, we found that accounting for photosynthetic acclimation improves the prediction of carbon assimilation in a majority of the tested models. Non-stomatal mechanisms contributed significantly to the reduction of photosynthesis under drought conditions in all tested models. Drought effects on photosynthesis could not accurately be explained by the hydraulic impairment functions embedded in the stomatal models alone, indicating that photosynthetic acclimation must be considered to improve estimates of carbon assimilation during drought. Summary StatementAccounting for photosynthetic acclimation improves the predictions of carbon assimilation in all the stomatal optimization models evaluated. The influence of drought on photosynthesis cannot be fully explained by the hydraulic impairment function of the stomatal models alone.

biophysics↗

Contrasting carbon cycle along tropical forest aridity gradients in W Africa and Amazonia

2Tropical forests cover large areas of equatorial Africa and play a significant role in the global carbon cycle. However, there has been a lack of in-situ measurements to understand the forests gross and net primary productivity (GPP and NPP) and their allocation. Here we present the first detailed field assessment of the carbon budget of multiple forest sites in Africa, by monitoring 14 one-hectare plots along an aridity gradient in Ghana. When compared with an equivalent aridity gradient in Amazonia using the same measurement protocol, the studied West African forests generally had higher GPP and NPP and lower carbon use efficiency (CUE). The West African aridity gradient consistently shows the highest NPP, CUE, GPP, and autotrophic respiration at a medium-aridity site, Bobiri. Notably, NPP and GPP of the site are the highest yet reported anywhere in the tropics using similar methods. Widely used data products (MODIS and FLUXCOM) substantially underestimate productivity when compared to in situ measurements, in Amazonia and especially in Africa. Our analysis suggests that the high productivity of the African forests is linked to their large GPP allocation to canopy and semi-deciduous characteristics, which may be a result of a seasonal climate coupled with high soil fertility.

ecology↗

Competition for light can drive adverse species-composition shifts in the Amazon Forest under elevated CO2

The resilience of biodiverse forests to climate change depends on an interplay of adaptive processes operating at multiple temporal and organizational scales. These include short-term acclimation of physiological processes like photosynthesis and respiration, mid-term changes in forest structure due to competition, and long-term changes in community composition arising from competitive exclusion and genetic trait evolution. To investigate the roles of diversity and adaptation for forest resilience, we present Plant-FATE, a parsimonious eco-evolutionary vegetation model. Tested with data from a hyperdiverse Amazonian terra-firme forest, our model accurately predicts multiple emergent ecosystem properties characterizing forest structure and function. Under elevated CO2 conditions, we predict an increase in productivity, leaf area, and aboveground biomass, with the magnitude of this increase declining in nutrient-deprived soils if trees allocate more carbon to the rhizosphere to overcome nutrient limitation. Furthermore, increased aboveground productivity leads to greater competition for light and drives a shift in community composition towards fast-growing but short-lived species characterized by lower wood densities. Such a transition reduces the carbon residence time of woody biomass, dampening carbon-sink strength and potentially rendering the Amazon Forest more vulnerable to future climatic extreme events.

ecology↗

A unifying principle for global greenness patterns and trends

Vegetation cover regulates the exchanges of energy, water and carbon between land and atmosphere. Remotely-sensed fractional absorbed photosynthetically active radiation (fAPAR), a land-surface greenness metric, depends on carbon allocation to foliage while also controlling photon flux for photosynthesis. Greenness is thus both a driver and an outcome of gross primary production (GPP). An equation with just two (globally) fitted parameters describes annual maximum fAPAR (fAPARmax) as the smaller of a water-limited value, transpiring a constant fraction of annual precipitation, and an energy-limited value, maximizing annual plant growth. This minimalist description reproduces global greenness patterns, and the consistent temporal trends among remote-sensing products, as accurately as the best-performing dynamic global vegetation models. Widely observed greening is attributed to the influence of rising carbon dioxide on the light- and water-use efficiencies of GPP, augmented by wetting in some dry regions and warming in high latitudes. Limited regions show browning, attributed to drying.

ecology↗

Global terrestrial nitrogen uptake and nitrogen use efficiency

Plant biomass production (BP), nitrogen uptake (Nup) and their ratio, nitrogen use efficiency (NUE), must be quantified to understand how nitrogen (N) cycling constrains terrestrial carbon (C) uptake. But the controls of key plant processes determining Nup and NUE, including BP, C and N allocation, tissue C:N ratios and N resorption efficiency (NRE), remain poorly known. We compiled measurements from 804 forest and grassland sites and derived regression models for each of these processes with growth temperature, vapour pressure deficit, stand age, soil C:N ratio, fAPAR (remotely sensed fraction of photosynthetically active radiation absorbed by green vegetation) and growing-season average daily incident photosynthetic photon flux density (gPPFD) (effectively the seasonal concentration of light availability, which increases polewards) as predictors. An empirical model for leaf N was based on optimal photosynthetic capacity (a function of gPPFD and climate) and observed leaf mass-per-area. The models were used to produce global maps of Nup and NUE. Global BP was estimated as 72 Pg C/yr; Nup as 950 Tg N/yr; and NUE as 76 gC/gN. Forest BP was found to increase with growth temperature and fAPAR and to decrease with stand age, soil C:N ratio and gPPFD. Forest NUE is controlled primarily by climate through its effect on C allocation - especially to leaves, being richer in N than other tissues. NUE is greater in colder climates, where N is less readily available, because belowground allocation is increased. NUE is also greater in drier climates because leaf allocation is reduced. NRE is enhanced (further promoting NUE) in both cold and dry climates. These findings can provide observationally based benchmarks for model representations of C-N cycle coupling. State-of-the-art vegetation models in the TRENDY ensemble showed variable performance against these benchmarks, and models including coupled C-N cycling produced relatively poor simulations of Nup and NUE.

ecology↗

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

ecology↗

Plant hydraulics coordinated with photosynthetic traits and climate

The coupling between water loss and carbon dioxide uptake drives the coordination of plant hydraulic and photosynthetic traits. Analysing multi-species measurements on a 3000 m elevation gradient, we found that hydraulic and leaf-economic traits were less plastic, and more closely associated with phylogeny, than photosynthetic traits. The two trait sets are linked by the sapwood-to-leaf area ratio (Huber value, vH), shown here to be codetermined by sapwood hydraulic conductance (KS), leaf mass-per-area (LMA) and photosynthetic capacity (Vcmax). Substantial hydraulic diversity was related to the trade-off between KS and vH. Leaf drought tolerance (inferred from turgor loss point, -{pi}tlp) increased with wood density, but the trade-off between hydraulic efficiency (KS) and -{pi}tlp was weak. The least-cost optimality framework was extended to predict trait (KS-dominated) and environmental (temperature-dominated) effects on vH. These results suggest an approach to include photosynthetic-hydraulic coordination in land-surface models; however, prediction of non-plastic trait distributions remains a challenge.

ecology↗

Leaf economics explained by optimality principles

The worldwide leaf economics spectrum relates leaf lifespan (LL) to leaf dry mass per unit area (LMA)1. By combining three well-supported principles2-4, we show that an isometric relationship between these two quantities maximizes the leafs 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 species1,5, 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 decrease6. 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.

ecology↗

Towards a unified theory of plant photosynthesis and hydraulics

The global carbon and water cycles are governed by the coupling of CO2 and water vapour exchanges through the leaves of terrestrial plants, controlled by plant adaptations to balance carbon gains and hydraulic risks. We introduce a trait-based optimality theory that unifies the treatment of stomatal responses and biochemical acclimation of plants to environments changing on multiple timescales. Tested with experimental data from 18 species, our model successfully predicts the simultaneous decline in carbon assimilation rate, stomatal conductance, and photosynthetic capacity during progressive soil drought. It also correctly predicts the dependencies of gas exchange on atmospheric vapour pressure deficit, temperature, and CO2. Model predictions are also consistent with widely observed empirical patterns, such as the distribution of hydraulic strategies. Our unified theory opens new avenues for reliably modelling the interactive effects of drying soil and rising atmospheric CO2 on global photosynthesis and transpiration.

plant biology↗

Forest production efficiency increases with growth temperature

Introductory paragraphWe present a global analysis of the relationship of forest production efficiency (FPE) to stand age and climate, based on a large compilation of data on gross primary production and either biomass production or net primary production. FPE is important for both forest production and atmospheric carbon dioxide uptake. Earlier findings - FPE declining with age - are supported by this analysis. However, FPE also increases with absolute latitude, precipitation and (all else equal) with temperature. The temperature effect is opposite to what would be expected based on the short-term physiological response of respiration rates to temperature. It implies top-down regulation of forest carbon loss, perhaps reflecting the higher carbon costs of nutrient acquisition in colder climates. Current ecosystem models do not reproduce this phenomenon. They consistently predict lower FPE in warmer climates, and are therefore likely to overestimate carbon losses in a warming climate.

plant biology↗

Drivers of vascular plant species area relationships in British broadleaved woodlands and their effects on the species area curve

The loss of plant biodiversity in Great Britain is a major concern, with a fifth of species endangered or vulnerable according to the latest IUCN Red List. The Governments 25 Year Plan for the environment aims to halt this loss and build new habitats, including new woodlands. To ensure that biodiversity loss is halted in existing woodlands and gain is maximised in new ones, we need to better understand which drivers have been most influential in controlling biodiversity. Here we focus on vascular plant species richness. Previous attempts to explain plant species richness have mainly focussed on alpha diversity in a consistent, fixed unit area. Here, we additionally undertake a novel analysis of the effects of environmental heterogeneity and abiotic factors on species-area relationships derived from 16 randomly placed quadrats in each of 103 semi-natural, broad-leaved woodlands across Britain. Species-area relationships were examined at two scales (4m2 to 200m2 and 200m2 to 3200m2) to explore the relationship between the drivers of species richness and the exponent z, of the canonical species-area curve, S = cAz. We also explore the use of a new metric {zeta}r, based on zeta diversity to quantify heterogeneity. Zeta diversity quantifies the number of species shared between multiple combinations of plots. Habitat heterogeneity increased species richness, as did the proximity of the woodlands to surrounding natural habitats. Higher levels of soil organic matter and the progression of woodlands to later successional stages, decreased species richness. Richness was also seen to have a unimodal response to soil acidity with a peak around pH 6. At the smaller scale, heterogeneity elements in the woodland such as riparian zones or coppicing led to an increase in the value of the exponent of the species area curve. At the larger scale, species turnover led to an increase in the exponent of the curves while succession led to a decrease. At both scales, soil organic matter content had a negative effect. {zeta}r was found to be a significant and important variable and to affect both species richness and the slope of the species accumulation curves at larger scales. SynthesisHabitat heterogeneity measures included the presence of coppicing, open areas such as rides and riparian zones and the difference between species assemblages in different plots in the woodland. Results suggest that to maximize vascular plant diversity, woodlands should be managed for heterogeneity. In addition, the increase in richness with exposure to surrounding natural habitats suggests that woodlands benefit from being embedded in more benign habitats and further, that land management surrounding woodlands has a clear role to play in supporting biodiversity within woodlands. This is an area were Agri-environment schemes have an important role.

ecology↗

Plant respiration: controlled by photosynthesis or biomass?

Two simplifying hypotheses have been proposed for whole-plant respiration. One links respiration to photosynthesis; the other to biomass. Using a first-principles carbon balance model with a prescribed live woody biomass turnover, applied at a forest research site where multidecadal measurements are available for comparison, we show that if turnover is fast the accumulation of respiring biomass is low and respiration depends primarily on photosynthesis; while if turnover is slow the accumulation of respiring biomass is high and respiration depends primarily on biomass. But the first scenario is inconsistent with evidence for substantial carryover of fixed carbon between years, while the second implies far too great an increase in respiration during stand development - leading to depleted carbohydrate reserves and an unrealistically high mortality risk. These two mutually incompatible hypotheses are thus both incorrect. Respiration is not linearly related either to photosynthesis or to biomass, but it is more strongly controlled by recent photosynthates (and reserve availability) than by total biomass.

ecology↗