Stefano ManzoniUniversitetslektor, docent
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I urval från Stockholms universitets publikationsdatabas
Persistence of soil organic carbon caused by functional complexity
2020. Johannes Lehmann (et al.). Nature Geoscience 13, 529-534Artikel
Soil organic carbon management has the potential to aid climate change mitigation through drawdown of atmospheric carbon dioxide. To be effective, such management must account for processes influencing carbon storage and re-emission at different space and time scales. Achieving this requires a conceptual advance in our understanding to link carbon dynamics from the scales at which processes occur to the scales at which decisions are made. Here, we propose that soil carbon persistence can be understood through the lens of decomposers as a result of functional complexity derived from the interplay between spatial and temporal variation of molecular diversity and composition. For example, co-location alone can determine whether a molecule is decomposed, with rapid changes in moisture leading to transport of organic matter and constraining the fitness of the microbial community, while greater molecular diversity may increase the metabolic demand of, and thus potentially limit, decomposition. This conceptual shift accounts for emergent behaviour of the microbial community and would enable soil carbon changes to be predicted without invoking recalcitrant carbon forms that have not been observed experimentally. Functional complexity as a driver of soil carbon persistence suggests soil management should be based on constant care rather than one-time action to lock away carbon in soils. Dynamic interactions between chemical and biological controls govern the stability of soil organic carbon and drive complex, emergent patterns in soil carbon persistence.
Organizing principles for vegetation dynamics
2020. Oskar Franklin (et al.). Nature plants 6 (5), 444-453Artikel
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. Integrating natural selection and other organizing principles into next-generation vegetation models could render them more theoretically sound and useful for earth system applications and modelling climate impacts.
Dynamic upscaling of decomposition kinetics for carbon cycling models
2020. Arjun Chakrawal (et al.). Geoscientific Model Development 13 (3), 1399-1429Artikel
The distribution of organic substrates and microorganisms in soils is spatially heterogeneous at the microscale. Most soil carbon cycling models do not account for this microscale heterogeneity, which may affect predictions of carbon (C) fluxes and stocks. In this study, we hypothesize that the mean respiration rate (R) over bar at the soil core scale (i) is affected by the microscale spatial heterogeneity of substrate and microorganisms and (ii) depends upon the degree of this heterogeneity. To theoretically assess the effect of spatial heterogeneities on (R) over bar, we contrast heterogeneous conditions with isolated patches of substrate and microorganisms versus spatially homogeneous conditions equivalent to those assumed in most soil C models. Moreover, we distinguish between biophysical heterogeneity, defined as the nonuniform spatial distribution of substrate and microorganisms, and full heterogeneity, defined as the nonuniform spatial distribution of substrate quality (or accessibility) in addition to biophysical heterogeneity. Four common formulations for decomposition kinetics (linear, multiplicative, Michaelis-Menten, and inverse Michaelis-Menten) are considered in a coupled substrate-microbial biomass model valid at the microscale. We start with a 2-D domain characterized by a heterogeneous substrate distribution and numerically simulate organic matter dynamics in each cell in the domain. To interpret the mean behavior of this spatially explicit system, we propose an analytical scale transition approach in which microscale heterogeneities affect (R) over bar through the second-order spatial moments (spatial variances and covariances). The model assuming homogeneous conditions was not able to capture the mean behavior of the heterogeneous system because the second-order moments cause (R) over bar to be higher or lower than in the homogeneous system, depending on the sign of these moments. This effect of spatial heterogeneities appears in the upscaled nonlinear decomposition formulations, whereas the upscaled linear decomposition model deviates from homogeneous conditions only when substrate quality is heterogeneous. Thus, this study highlights the inadequacy of applying at the macroscale the same decomposition formulations valid at the microscale and proposes a scale transition approach as a way forward to capture microscale dynamics in core-scale models.
Quantifying Microbial Metabolism in Soils Using Calorespirometry — A Bioenergetics Perspective
2020. Arjun Chakrawal (et al.). Soil Biology and Biochemistry 148Artikel
Microbial carbon use efficiency (CUE) measures the partitioning between anabolic and catabolic processes. While most work on CUE has been based on carbon (C) mass flows, the roles of organic C energy contents and microbial energy demand on CUE have been rarely considered. Thus, a bioenergetics perspective could provide new insights on how microorganisms utilize C and ultimately allow evaluating their role in C stabilization in soils. Recently, the calorespirometric ratio (CR)— the ratio of heat dissipation and respiration— has been used to characterize the efficiency of microbial growth in soils. Here, we formulate a coupled mass and energy balance model for microbial growth and provide a generalized relationship between CUE and CR. In the model, we consider two types of organic C in soils: an added substrate (e.g., glucose) and the native soil organic matter (SOM), to also account for priming effects. Furthermore, we consider both aerobic and fermentation metabolic pathways. We use this model as a framework to generalize previous formulations and generate hypotheses on the expected variations in CR as a function of substrate quality, metabolic pathways, and microbial traits (specifically CUE). In turn, the same equations can be used to estimate CUE from measured CR. Our results confirm previous findings on CR and show that without microbial growth, CR depends only on the rates of the different metabolic pathways, while CR is also a function of the growth yields for these metabolic pathways when microbial growth occurs. Under strictly aerobic conditions, CUE increases with increasing CR for substrates with a higher degree of reduction than that of the microbial biomass, while CUE decreases with increasing CR for substrates with a lower degree of reduction than the microbial biomass. When aerobic reactions and fermentation occur simultaneously, the relation between CUE and CR is mediated by (i) the degree of reduction of the substrates, (ii) the rates and growth yields of all metabolic pathways, and (iii) the contribution of SOM priming to microbial growth. Using the proposed framework, calorespirometry can be used to evaluate CUE and the role of different metabolic pathways in soil systems.
Rainfall intensification increases the contribution of rewetting pulses to soil heterotrophic respiration
2020. Stefano Manzoni (et al.). Biogeosciences 17 (15), 4007-4023Artikel
Soil drying and wetting cycles promote carbon (C) release through large heterotrophic respiration pulses at rewetting, known as the Birch effect. Empirical evidence shows that drier conditions before rewetting and larger changes in soil moisture at rewetting cause larger respiration pulses. Because soil moisture varies in response to rainfall, these respiration pulses also depend on the random timing and intensity of precipitation. In addition to rewetting pulses, heterotrophic respiration continues during soil drying, eventually ceasing when soils are too dry to sustain microbial activity. The importance of respiration pulses in contributing to the overall soil heterotrophic respiration flux has been demonstrated empirically, but no theoretical investigation has so far evaluated how the relative contribution of these pulses may change along climatic gradients or as precipitation regimes shift in a given location. To fill this gap, we start by assuming that heterotrophic respiration rates during soil drying and pulses at rewetting can be treated as random variables dependent on soil moisture fluctuations, and we develop a stochastic model for soil heterotrophic respiration rates that analytically links the statistical properties of respiration to those of precipitation. Model results show that both the mean rewetting pulse respiration and the mean respiration during drying increase with increasing mean precipitation. However, the contribution of respiration pulses to the total heterotrophic respiration increases with decreasing precipitation frequency and to a lesser degree with decreasing precipitation depth, leading to an overall higher contribution of respiration pulses under future more intermittent and intense precipitation. Specifically, higher rainfall intermittency at constant total rainfall can increase the contribution of respiration pulses up to similar to 10 % or 20 % of the total heterotrophic respiration in mineral and organic soils, respectively. Moreover, the variability of both components of soil heterotrophic respiration is also predicted to increase under these conditions. Therefore, with future more intermittent precipitation, respiration pulses and the associated nutrient release will intensify and become more variable, contributing more to soil biogeochemical cycling.
Hydro-climatic controls explain variations in catchment-scale nitrogen use efficiency
2020. Anna Scaini (et al.). Environmental Research Letters 15 (9)Artikel
The efficiency of fertilizer conversion to harvestable products is often low in annual crops such that large amounts of nutrients are lost from fields with negative consequences for the environment. Focusing on nitrogen (N) use efficiency (NUE: the ratio of N in harvested products over the sum of all N inputs), we propose that hydrological controls can explain variations in NUE, because water mediates both the uptake of N by plants and N leaching. We assess these controls at the catchment scale, at which the water balance can be constrained by precipitation and runoff data and NUE can be quantified with census data. With this approach we test the hypotheses that a higher evaporative ratio (ET/P: the ratio of evapotranspiration over precipitation) increases N retention, thereby increasing NUE both across catchments at a given time and through time. With data from 73 catchments in the United States, encompassing a wide range of pedoclimatic conditions for the period 1988-2007, we apply a linear mixed effect model to test the effect of ET/P on NUE. Supporting our hypotheses, ET/P was positively related to NUE, and NUE increased through time. Moreover, we found an interaction between ET/P and time, such that the ET/P effect on NUE decreased in the period 1998-2007. We conclude that climatic changes that increase ET/P without negatively affecting yields, will increase N retention in the examined catchments.
Understanding coastal wetland conditions and futures by closing their hydrologic balance
2020. Stefano Manzoni (et al.). Hydrology and Earth System Sciences 24 (7), 3557-3571Artikel
Coastal wetlands and lagoons are under pressure due to competing demands for freshwater resources and climatic changes, which may increase salinity and cause a loss of ecological functions. These pressures are particularly high in Mediterranean regions with high evaporative demand compared to precipitation. To manage such wetlands and maximize their provision of ecosystem services, their hydrologic balance must be quantified. However, multiple channels, diffuse surface water exchanges, and diverse groundwater pathways complicate the quantification of different water balance components. To overcome this difficulty, we developed a mass balance approach based on coupled water and salt balance equations to estimate currently unknown water exchange fluxes through the Gialova lagoon, southwestern Peloponnese, Greece. Our approach facilitates quantification of both saline and freshwater exchange fluxes, using measured precipitation, water depth and salinity, and estimated evaporation rates over a study period of 2 years (2016-2017). While water exchanges were dominated by evaporation and saline water inputs from the sea during the summer, precipitation and freshwater inputs were more important during the winter. About 40 % and 60 % of the freshwater inputs were from precipitation and lateral freshwater flows, respectively. Approximately 70 % of the outputs was due to evaporation, with the remaining 30 % being water flow from the lagoon to the sea. Under future drier and warmer conditions, salinity in the lagoon is expected to increase, unless freshwater inputs are enhanced by restoring hydrologic connectivity between the lagoon and the surrounding freshwater bodies. This restoration strategy would be fundamental to stabilizing the current wide seasonal fluctuations in salinity and maintain ecosystem functionality but could be challenging to implement due to expected reductions in water availability in the freshwater bodies supporting the lagoon.
Modelling water fluxes in plants
2019. Maurizio Mencuccini, Stefano Manzoni, Bradley Christoffersen. New Phytologist 222 (3), 1207-1222Artikel
Models of plant water fluxes have evolved from studies focussed on understanding the detailed structure and functioning of specific components of the soil-plant-atmosphere (SPA) continuum to architectures often incorporated inside eco-hydrological and terrestrial biosphere (TB) model schemes. We review here the historical evolution of this field, examine the basic structure of a simplified individual-based model of plant water transport, highlight selected applications for specific ecological problems and conclude by examining outstanding issues requiring further improvements in modelling vegetation water fluxes. We particularly emphasise issues related to the scaling from tissue-level traits to individual-based predictions of water transport, the representation of nonlinear and hysteretic behaviour in soil-xylem hydraulics and the need to incorporate knowledge of hydraulics within broader frameworks of plant ecological strategies and their consequences for predicting community demography and dynamics.
Do alternative irrigation strategies for rice cultivation decrease water footprints at the cost of long-term soil health?
2019. John Livsey (et al.). Environmental Research Letters 14 (7)Artikel
The availability of water is a growing concern for flooded rice production. As such, several water-saving irrigation practices have been developed to reduce water requirements. Alternate wetting and drying and mid-season drainage have been shown to potentially reduce water requirements while maintaining rice yields when compared to continuous flooding. With the removal of permanently anaerobic conditions during the growing season, water-saving irrigation can also reduce CO2 equivalent (CO2eq) emissions, helping reduce the impact of greenhouse gas (GHG) emissions. However, the long-term impact of water-saving irrigation on soil organic carbon (SOC)-used here as an indicator of soil health and fertility-has not been explored. We therefore conducted a meta-analysis to assess the effects of common water-saving irrigation practices (alternate wetting and drying and mid-season drainage) on (i) SOC, and (ii) GHG emissions. Despite an extensive literature search, only 12 studies were found containing data to constrain the soil C balance in both continuous flooding and water-saving irrigation plots, highlighting the still limited understanding of long-term impacts of water-saving irrigation on soil health and GHG emissions. Water-saving irrigation was found to reduce emissions of CH4 by 52.3% and increased those of CO2 by 44.8%. CO2eq emissions were thereby reduced by 18.6% but the soil-to-atmosphere carbon (C) flux increased by 25% when compared to continuous flooding. Water-saving irrigation was also found to have a negative effect on both SOC-reducing concentrations by 5.2%-and soil organic nitrogen-potentially depleting stocks by more than 100 kgN/ha per year. While negative effects of water-saving irrigation on rice yield may not be visible in short-term experiments, care should be taken when assessing the long-term sustainability of these irrigation practices because they can decrease soil fertility. Strategies need to be developed for assessing the more long-term effects of these irrigation practices by considering trade-offs between water savings and other ecosystem services.
A plant-microbe interaction framework explaining nutrient effects on primary production
2018. P. T. Capek (et al.). Nature Ecology & Evolution 2 (10), 1588-1596Artikel
In most terrestrial ecosystems, plant growth is limited by nitrogen and phosphorus. Adding either nutrient to soil usually affects primary production, but their effects can be positive or negative. Here we provide a general stoichiometric framework for interpreting these contrasting effects. First, we identify nitrogen and phosphorus limitations on plants and soil microorganisms using their respective nitrogen to phosphorus critical ratios. Second, we use these ratios to show how soil microorganisms mediate the response of primary production to limiting and non-limiting nutrient addition along a wide gradient of soil nutrient availability. Using a meta-analysis of 51 factorial nitrogen-phosphorus fertilization experiments conducted across multiple ecosystems, we demonstrate that the response of primary production to nitrogen and phosphorus additions is accurately predicted by our stoichiometric framework. The only pattern that could not be predicted by our original framework suggests that nitrogen has not only a structural function in growing organisms, but also a key role in promoting plant and microbial nutrient acquisition. We conclude that this stoichiometric framework offers the most parsimonious way to interpret contrasting and, until now, unresolved responses of primary production to nutrient addition in terrestrial ecosystems.
The ecohydrological context of drought and classification of plant responses
2018. Xue Feng (et al.). Ecology Letters 21 (11), 1723-1736Artikel
Many recent studies on drought‐induced vegetation mortality have explored how plant functional traits, and classifications of such traits along axes of, for example, isohydry–anisohydry, might contribute to predicting drought survival and recovery. As these studies proliferate, the consistency and predictive value of such classifications need to be carefully examined. Here, we outline the basis for a systematic classification of plant drought responses that accounts for both environmental conditions and functional traits. We use non‐dimensional analysis to integrate plant traits and metrics of environmental variation into groups that can be associated with alternative drought stress pathways (hydraulic failure and carbon limitation), and demonstrate that these groupings predict physiological drought outcomes using both synthetic and measured data. In doing so, we aim to untangle some confounding effects of environment and trait variations that undermine current classification schemes, advocate for more careful treatment of the environmental context within which plants experience and respond to drought, and outline a pathway towards a general classification of drought vulnerability.
Reviews and syntheses
2018. Stefano Manzoni (et al.). Biogeosciences 15 (19), 5929-5949Artikel
The cycling of carbon (C) between the Earth surface and the atmosphere is controlled by biological and abiotic processes that regulate C storage in biogeochemical compartments and release to the atmosphere. This partitioning is quantified using various forms of C-use efficiency (CUE) - the ratio of C remaining in a system to C entering that system. Biological CUE is the fraction of C taken up allocated to biosynthesis. In soils and sediments, C storage depends also on abiotic processes, so the term C-storage efficiency (CSE) can be used. Here we first review and reconcile CUE and CSE definitions proposed for autotrophic and heterotrophic organisms and communities, food webs, whole ecosystems and watersheds, and soils and sediments using a common mathematical framework. Second, we identify general CUE patterns; for example, the actual CUE increases with improving growth conditions, and apparent CUE decreases with increasing turnover. We then synthesize > 5000CUE estimates showing that CUE decreases with increasing biological and ecological organization - from uni-cellular to multicellular organisms and from individuals to ecosystems. We conclude that CUE is an emergent property of coupled biological-abiotic systems, and it should be regarded as a flexible and scale-dependent index of the capacity of a given system to effectively retain C.
Optimal metabolic regulation along resource stoichiometry gradients
2017. Stefano Manzoni (et al.). Ecology Letters 20 (9), 1182-1191Artikel
Most heterotrophic organisms feed on substrates that are poor in nutrients compared to their demand, leading to elemental imbalances that may constrain their growth and function. Flexible carbon (C)-use efficiency (CUE, C used for growth over C taken up) can represent a strategy to reduce elemental imbalances. Here, we argue that metabolic regulation has evolved to maximise the organism growth rate along gradients of nutrient availability and translated this assumption into an optimality model that links CUE to substrate and organism stoichiometry. The optimal CUE is predicted to decrease with increasing substrate C-to-nutrient ratio, and increase with nutrient amendment. These predictions are generally confirmed by empirical evidence from a new database of c. 2200 CUE estimates, lending support to the hypothesis that CUE is optimised across levels of organisation (microorganisms and animals), in aquatic and terrestrial systems, and when considering nitrogen or phosphorus as limiting nutrients.