Stefano Manzoni

Stefano Manzoni

Universitetslektor, docent

Visa sidan på svenska
Works at Department of Physical Geography
Telephone 08-674 78 02
Visiting address Svante Arrhenius väg 8
Room T326
Postal address Inst för naturgeografi 106 91 Stockholm

About me

I am an associate professor in Ecohydrology, with research interests spanning soil-vegetation-atmosphere interactions, hydro-climatic impacts on carbon and nutrient cycling, sustainable resource use and management, and ecological stoichiometry. My approach is based on process-based, conceptual, and stochastic models of water, carbon, and nutrient fluxes, which are tested using global datasets.

Our team


Courses in the MSc Program in Hydrology, Hydrogeology and Water Resources:

  • Local to global water vulnerability and resilience (GE7025)
  • Ecohydrology – a Mediterranean perspective (GE7049)
  • Information and Modelling Systems for Land and Water Methods (GE7006, co-instructor)


New paper in Nature Plants!

Ongoing projects

Awards (most recent)

2018-2019 Highly Cited Researcher (category: cross-field), Clarivate Analytics

2017 Junior Marchi Lecturer, Gruppo Italiano di Idraulica

2014 Early Career Hydrologic Sciences Award, American Geophysical Union


A selection from Stockholm University publication database
  • 2019. Maurizio Mencuccini, Stefano Manzoni, Bradley Christoffersen. New Phytologist 222 (3), 1207-1222

    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.

  • 2019. Giulia Vico (et al.). Plant, Cell and Environment 42 (6), 1913-1928

    Under future climates, leaf temperature (T-l) will be higher and more variable. This will affect plant carbon (C) balance because photosynthesis and respiration both respond to short-term (subdaily) fluctuations in T-l and acclimate in the longer term (days to months). This study asks the question: To what extent can the potential and speed of photosynthetic acclimation buffer leaf C gain from rising and increasing variable T-l? We quantified how increases in the mean and variability of growth temperature affect leaf performance (mean net CO2 assimilation rates, A(net); its variability; and time under near-optimal photosynthetic conditions), as mediated by thermal acclimation. To this aim, the probability distribution of A(net) was obtained by combining a probabilistic description of short- and long-term changes in T-l with data on A(net) responses to these changes, encompassing 75 genera and 111 species, including both C3 and C4 species. Our results show that (a) expected increases in T-l variability will decrease mean A(net) and increase its variability, whereas the effects of higher mean T-l depend on species and initial T-l, and (b) acclimation reduces the effects of leaf warming, maintaining A(net) at >80% of its maximum under most thermal regimes.

  • 2019. Gabriele Messori (et al.). Environmental Research Letters 14 (6)

    The terrestrial biosphere is a key component of the global carbon cycle and is heavily influenced by climate. Climate variability can be diagnosed through metrics ranging from individual environmental variables, to collections of variables, to the so-called climate modes of variability. Similarly, the impact of a given climate variation on the terrestrial carbon cycle can be described using several metrics, including vegetation indices, measures of ecosystem respiration and productivity and net biosphere-atmosphere fluxes. The wide range of temporal (from sub-daily to paleoclimatic) and spatial (from local to continental and global) scales involved requires a scale-dependent investigation of the interactions between the carbon cycle and climate. However, a comprehensive picture of the physical links and correlations between climate drivers and carbon cycle metrics at different scales remains elusive, framing the scope of this contribution. Here, we specifically explore how climate variability metrics (from single variables to complex indices) relate to the variability of the carbon cycle at sub-daily to interannual scales (i.e. excluding long-term trends). The focus is on the interactions most relevant to the European terrestrial carbon cycle. We underline the broad areas of agreement and disagreement in the literature, and conclude by outlining some existing knowledge gaps and by proposing avenues for improving our holistic understanding of the role of climate drivers in modulating the terrestrial carbon cycle.

  • 2019. Benjamin M. C. Fischer (et al.). Science of the Total Environment 657, 853-862

    There is an urgent need to develop agricultural methods that balance water supply and demand while at the same time improve resilience to climate variability. A promising instrument to address this need is biochar - a charcoal made from pyrolyzed organic material. However, it is often unclear how, if at all, biochar improves soil water availability, plant water consumption rates and crop yields. To address this question, we synthesized literature-derived observational data and evaluated the effects of biochar on evapotranspiration using a minimal soil water balance model. Results from the model were interpreted in the Budyko framework to assess how climatic conditions mediate the impacts of biochar on water fluxes. Our analysis of literature-derived observational data showed that while biochar addition generally increases the soil water holding capacity, it can have variable impacts on soil water retention relative to control conditions. Our modelling demonstrated that biochar increases long-term evapotranspiration rates, and therefore plant water availability, by increasing soil water retention capacity - especially in water-limited regions. Biochar amendments generally increased crop yields (75% of the compiled studies) and, in several cases (35% of the compiled studies), biochar amendments simultaneously increased crop yield and water use efficiencies. Hence, while biochar amendments are promising, the potential for variable impact highlights the need for targeted research on how biochar affects the soil-plant-water cycle.

  • 2019. Xue Feng (et al.). Plant, Cell and Environment 42 (4), 1104-1111

    Despite the appeal of the iso/anisohydric framework for classifying plant drought responses, recent studies have shown that such classifications can be strongly affected by a plant's environment. Here, we present measured in situ drought responses to demonstrate that apparent isohydricity can be conflated with environmental conditions that vary over space and time. In particular, we (a) use data from an oak species (Quercus douglasii) during the 2012-2015 extreme drought in California to demonstrate how temporal and spatial variability in the environment can influence plant water potential dynamics, masking the role of traits; (b) explain how these environmental variations might arise from climatic, topographic, and edaphic variability; (c) illustrate, through a common garden thought experiment, how existing trait-based or response-based isohydricity metrics can be confounded by these environmental variations, leading to Type-1 (false positive) and Type-2 (false negative) errors; and (d) advocate for the use of model-based approaches for formulating alternate classification schemes. Building on recent insights from greenhouse and vineyard studies, we offer additional evidence across multiple field sites to demonstrate the importance of spatial and temporal drivers of plants' apparent isohydricity. This evidence challenges the use of isohydricity indices, per se, to characterize plant water relations at the global scale.

  • 2019. Maurizio Mencuccini (et al.). New Phytologist

    Biomass and area ratios between leaves, stems and roots regulate many physiological and ecological processes. The Huber value H-v (sapwood area/leaf area ratio) is central to plant water balance and drought responses. However, its coordination with key plant functional traits is poorly understood, and prevents developing trait-based prediction models. Based on theoretical arguments, we hypothesise that global patterns in H-v of terminal woody branches can be predicted from variables related to plant trait spectra, that is plant hydraulics and size and leaf economics. Using a global compilation of 1135 species-averaged H-v, we show that H-v varies over three orders of magnitude. Higher H-v are seen in short small-leaved low-specific leaf area (SLA) shrubs with low K-s in arid relative to tall large-leaved high-SLA trees with high K-s in moist environments. All traits depend on climate but climatic correlations are stronger for explanatory traits than H-v. Negative isometry is found between H-v and K-s, suggesting a compensation to maintain hydraulic supply to leaves across species. This work identifies the major global drivers of branch sapwood/leaf area ratios. Our approach based on widely available traits facilitates the development of accurate models of above-ground biomass allocation and helps predict vegetation responses to drought.

  • 2019. Edmond Alavaisha, Stefano Manzoni, Regina Lindborg. Journal of Environmental Management 234, 159-166

    Converting natural and semi-natural vegetation to agriculture is currently the most significant land use change at global scale. This conversion leads to changes in soil nutrients and increased CO2 emissions. However, knowledge of how soil organic carbon and nutrients change under various farming management is still limited, especially for small scale farming systems. This study evaluated the effects of different farming systems on soil organic carbon (SOC), total nitrogen (TN) and total phosphorous (TP) in subsistence farming at Kilombero, Tanzania. We applied an in-situ experimental setup, comparing maize and rice farming with and without irrigation and difference in fertilizers, with replicated soil sampling at five soil depths to a depth of 60 cm. The results show that irrigation had a positive effect on profile-averaged concentrations of SOC and TN, while fertilization had a positive effect on TN. Higher concentrations and stocks of TN were found in maize field soils compered to rice fields. In the vertical profile, irrigation and fertilization had positive effects on concentrations of SOC and TN of top soil layers, and the interaction between irrigation and fertilization extended the effect to deeper soil layers. Our results indicate that moderate irrigation and fertilization can help to improve carbon storage and nutrient availability (TN) in small-scale farming soils in Africa.

  • 2019. Martin Thurner (et al.). Global Ecology and Biogeography 28 (5), 640-660

    Aim Information on the amount of carbon stored in the living tissue of tree stems (sapwood) is crucial for carbon and water cycle applications. Here, we aim to investigate sapwood-to-stem proportions and differences therein between tree genera and derive a sapwood biomass map. Location Northern Hemisphere boreal and temperate forests. Time period 2010. Major taxa studied Twenty-five common tree genera. Methods First, we develop a theoretical framework to estimate sapwood biomass for a given stem biomass by applying relationships between sapwood cross-sectional area (CSA) and stem CSA and between stem CSA and stem biomass. These measurements are extracted from a biomass and allometry database (BAAD), an extensive literature review and our own studies. The established allometric relationships are applied to a remote sensing-based stem biomass product in order to derive a spatially continuous sapwood biomass map. The application of new products on the distribution of stand density and tree genera facilitates the synergy of satellite and forest inventory data. Results Sapwood-to-stem CSA relationships can be modelled with moderate to very high modelling efficiency for different genera. The total estimated sapwood biomass equals 12.87 +/- 6.56 petagrams of carbon (PgC) in boreal (mean carbon density: 1.13 +/- 0.58 kgC m(-2)) and 15.80 +/- 9.10 PgC in temperate (2.03 +/- 1.17 kgC m(-2)) forests. Spatial patterns of sapwood-to-stem biomass proportions are crucially driven by the distribution of genera (spanning from 20-30% in Larix to > 70% in Pinus and Betula forests). Main conclusions The presented sapwood biomass map will be the basis for large-scale estimates of plant respiration and transpiration. The enormous spatial differences in sapwood biomass proportions reveal the need to consider the functionally more important sapwood instead of the entire stem biomass in global carbon and water cycle studies. Alterations in tree species distribution, induced by forest management or climate change, can strongly affect the available sapwood biomass even if stem biomass remains unchanged.

  • 2019. Giorgos Maneas (et al.). Water 11 (2)

    Human interventions during the last 70 years have altered the characteristics of the Gialova Lagoon, a coastal wetland that is part of a wider Natura 2000 site. In this study, we explore how human interventions and climate altered the wetland's hydrological conditions and habitats, leading to changing wetland functions over time. Our interpretations are based on a mixed methodological approach combining conceptual hydrologic models, analysis of aerial photographs, local knowledge, field observations, and GIS (Geographic Information System) analyses. The results show that the combined effects of human interventions and climate have led to increased salinity in the wetland over time. As a result, the fresh and brackish water marshes have gradually been turned into open water or replaced by halophytic vegetation with profound ecological implications. Furthermore, current human activities inside the Natura 2000 area and in the surrounding areas could further impact on the water quantity and quality in the wetland, and on its sensitive ecosystems. We suggest that a more holistic understanding of the broader socio-ecological system is needed to understand the dynamics of the wetland and to achieve sustainable long-term management and conservation strategies.

  • 2019. Simone Fatichi (et al.). Global Biogeochemical Cycles 33 (6), 620-648

    Present gaps in the representation of key soil biogeochemical processes such as the partitioning of soil organic carbon among functional components, microbial biomass and diversity, and the coupling of carbon and nutrient cycles present a challenge to improving the reliability of projected soil carbon dynamics. We introduce a new soil biogeochemistry module linked with a well-tested terrestrial biosphere model T&C. The module explicitly distinguishes functional soil organic carbon components. Extracellular enzymes and microbial pools are differentiated based on the functional roles of bacteria, saprotrophic, and mycorrhizal fungi. Soil macrofauna is also represented. The model resolves the cycles of nitrogen, phosphorus, and potassium. Model simulations for 20 sites compared favorably with global patterns of litter and soil stoichiometry, microbial and macrofaunal biomass relations with soil organic carbon, soil respiration, and nutrient mineralization rates. Long-term responses to bare fallow and nitrogen addition experiments were also in agreement with observations. Some discrepancies between predictions and observations are appreciable in the response to litter manipulation. Upon successful model reproduction of observed general trends, we assessed patterns associated with the carbon cycle that were challenging to address empirically. Despite large site-to-site variability, fine root, fungal, bacteria, and macrofaunal respiration account for 33%, 40%, 24%, and 3% on average of total belowground respiration, respectively. Simulated root exudation and carbon export to mycorrhizal fungi represent on average about 13% of plant net primary productivity. These results offer mechanistic and general estimates of microbial biomass and its contribution to respiration fluxes and to soil organic matter dynamics.

  • 2019. John Livsey (et al.). Environmental Research Letters 14 (7)

    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.

  • 2019. Quan Zhang (et al.). Environmental Research Letters 14 (7)

    Elevated vapor pressure deficit (VPD) due to drought and warming is well-known to limit canopy stomatal and surface conductance, but the impacts of elevated VPD on ecosystem gross primary productivity (GPP) are less clear. The intrinsic water use efficiency (iWUE), defined as the ratio of carbon (C) assimilation to stomatal conductance, links vegetation C gain and water loss and is a key determinant of how GPP will respond to climate change. While it is well-established that rising atmospheric CO2 increases ecosystem iWUE, historic and future increases in VPD caused by climate change and drought are often neglected when considering trends in ecosystem iWUE. Here, we synthesize long-term observations of C and water fluxes from 28 North American FLUXNET sites, spanning eight vegetation types, to demonstrate that ecosystem iWUE increases consistently with rising VPD regardless of changes in soil moisture. Another way to interpret this result is that GPP decreases less than surface conductance with increasing VPD. We also project how rising VPD will impact iWUE into the future. Results vary substantially from one site to the next; in a majority of sites, future increases in VPD (RCP 8.5, highest emission scenario) are projected to increase iWUE by 5%-15% by 2050, and by 10%-35% by the end of the century. The increases in VPD owing to elevated global temperatures could be responsible for a 0.13% year(-1) increase in ecosystem iWUE in the future. Our results highlight the importance of considering VPD impacts on iWUE independently of CO2 impacts.

  • 2019. Benjamin M. C. Fischer (et al.). Frontiers in Earth Science 7

    The stable isotope composition of water (delta O-18 and delta H-2) is an increasingly utilized tool to distinguish between different pools of water along the soil-plant-atmosphere continuum (SPAC) and thus provides information on how plants use water. Clear bottlenecks for the ubiquitous application of isotopic analysis across the SPAC are the relatively high-energy and specialized materials required to extract water from plant materials. Could simple and cost-effective do-it-yourself MacGyver methods be sufficient for extracting plant water for isotopic analysis? This study develops a suite of novel techniques for plant water extraction and compares them to a standard research-grade water extraction method. Our results show that low-tech methods using locally-sourced materials can indeed extract plant water consistently and comparably to what is done with other state-of-the-art methods. Further, our findings show that other factors play a larger role than water extraction methods in achieving the desired accuracy and precision of stable isotope composition: (1) appropriate transport, (2) fast sample processing and (3) efficient workflows. These results are methodologically promising for the rapid expansion of isotopic investigations, especially for citizen science and/or school projects or in remote areas, where improved SPAC understanding could help manage water resources to fulfill agricultural and other competing water needs.

  • 2018. P. T. Capek (et al.). Nature Ecology & Evolution 2 (10), 1588-1596

    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.

  • 2018. Loic D'Orangeville (et al.). Global Change Biology 24 (6), 2339-2351

    Projected changes in temperature and drought regime are likely to reduce carbon (C) storage in forests, thereby amplifying rates of climate change. While such reductions are often presumed to be greatest in semi-arid forests that experience widespread tree mortality, the consequences of drought may also be important in temperate mesic forests of Eastern North America (ENA) if tree growth is significantly curtailed by drought. Investigations of the environmental conditions that determine drought sensitivity are critically needed to accurately predict ecosystem feedbacks to climate change. We matched site factors with the growth responses to drought of 10,753 trees across mesic forests of ENA, representing 24 species and 346 stands, to determine the broad-scale drivers of drought sensitivity for the dominant trees in ENA. Here we show that two factors-the timing of drought, and the atmospheric demand for water (i.e., local potential evapotranspiration; PET)-are stronger drivers of drought sensitivity than soil and stand characteristics. Droughtinduced reductions in tree growth were greatest when the droughts occurred during early-season peaks in radial growth, especially for trees growing in the warmest, driest regions (i.e., highest PET). Further, mean species trait values (rooting depth and psi(50)) were poor predictors of drought sensitivity, as intraspecific variation in sensitivity was equal to or greater than interspecific variation in 17 of 24 species. From a general circulation model ensemble, we find that future increases in earlyseason PET may exacerbate these effects, and potentially offset gains in C uptake and storage in ENA owing to other global change factors.

  • 2018. P. M. Homyak (et al.). Ecology 99 (10), 2348-2362

    Soil moisture controls microbial activity and soil carbon cycling. Because microbial activity decreases as soils dry, decomposition of soil organic matter (SOM) is thought to decrease with increasing drought length. Yet, microbial biomass and a pool of water‐extractable organic carbon (WEOC) can increase as soils dry, perhaps implying microbes may continue to break down SOM even if drought stressed. Here, we test the hypothesis that WEOC increases as soils dry because exoenzymes continue to break down litter, while their products accumulate because they cannot diffuse to microbes. To test this hypothesis, we manipulated field plots by cutting off litter inputs and by irrigating and excluding precipitation inputs to extend or shorten the length of the dry season. We expected that the longer the soils would remain dry, the more WEOC would accumulate in the presence of litter, whereas shortening the length of the dry season, or cutting off litter inputs, would reduce WEOC accumulation. Lastly, we incubated grass roots in the laboratory and measured the concentration of reducing sugars and potential hydrolytic enzyme activities, strictly to understand the mechanisms whereby exoenzymes break down litter over the dry season. As expected, extending dry season length increased WEOC concentrations by 30% above the 108 μg C/g measured in untreated plots, whereas keeping soils moist prevented WEOC from accumulating. Contrary to our hypothesis, excluding plant litter inputs actually increased WEOC concentrations by 40% above the 105 μg C/g measured in plots with plants. Reducing sugars did not accumulate in dry senesced roots in our laboratory incubation. Potential rates of reducing sugar production by hydrolytic enzymes ranged from 0.7 to 10 μmol·g−1·h−1 and far exceeded the rates of reducing sugar accumulation (~0.001 μmol·g−1·h−1). Our observations do not support the hypothesis that exoenzymes continue to break down litter to produce WEOC in dry soils. Instead, we develop the argument that physical processes are more likely to govern short‐term WEOC dynamics via slaking of microaggregates that stabilize SOM and through WEOC redistribution when soils wet up, as well as through less understood effects of drought on the soil mineral matrix.

  • 2018. Stefano Manzoni (et al.). Biogeosciences 15 (19), 5929-5949

    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.

  • 2018. Xue Feng (et al.). Ecology Letters 21 (11), 1723-1736

    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.

  • 2018. Valentin Couvreur (et al.). Plant, Cell and Environment 41 (8), 1821-1839

    Trees grow by vertically extending their stems, so accurate stem hydraulic models are fundamental to understanding the hydraulic challenges faced by tall trees. Using a literature survey, we showed that many tree species exhibit continuous vertical variation in hydraulic traits. To examine the effects of this variation on hydraulic function, we developed a spatially explicit, analytical water transport model for stems. Our model allows Huber ratio, stem-saturated conductivity, pressure at 50% loss of conductivity, leaf area, and transpiration rate to vary continuously along the hydraulic path. Predictions from our model differ from a matric flux potential model parameterized with uniform traits. Analyses show that cavitation is a whole-stem emergent property resulting from non-linear pressure-conductivity feedbacks that, with gravity, cause impaired water transport to accumulate along the path. Because of the compounding effects of vertical trait variation on hydraulic function, growing proportionally more sapwood and building tapered xylem with height, as well as reducing xylem vulnerability only at branch tips while maintaining transport capacity at the stem base, can compensate for these effects. We therefore conclude that the adaptive significance of vertical variation in stem hydraulic traits is to allow trees to grow tall and tolerate operating near their hydraulic limits.

  • 2017. Giulia Vico (et al.). Environmental Research Letters 12 (6)

    Drought-deciduous and evergreen species are both common in tropical forests, where there is the need to cope with water shortages during periodic dry spells and over the course of the dry season. Which phenological strategy is favored depends on the long-term balance of carbon costs and gains that leaf phenology imposes as a result of the alternation of wet and dry seasons and the unpredictability of rainfall events. This study integrates a stochastic eco-hydrological framework with key plant economy traits to derive the long-term average annual net carbon gain of trees exhibiting different phenological strategies in tropical forests. The average net carbon gain is used as a measure of fitness to assess which phenological strategies are more productive and more evolutionarily stable (i.e. not prone to invasion by species with a different strategy). The evergreen strategy results in a higher net carbon gain and more evolutionarily stable communities with increasing wet season lengths. Reductions in the length of the wet season or the total rainfall, as predicted under climate change scenarios, should promote a shift towards more drought-deciduous communities, with ensuing implications for ecosystem functioning.

  • 2017. Preetisri Baskaran (et al.). New Phytologist 213 (3), 1452-1465

    Tree growth in boreal forests is limited by nitrogen (N) availability. Most boreal forest trees form symbiotic associations with ectomycorrhizal (ECM) fungi, which improve the uptake of inorganic N and also have the capacity to decompose soil organic matter (SOM) and to mobilize organic N (ECM decomposition'). To study the effects of ECM decomposition' on ecosystem carbon (C) and N balances, we performed a sensitivity analysis on a model of C and N flows between plants, SOM, saprotrophs, ECM fungi, and inorganic N stores. The analysis indicates that C and N balances were sensitive to model parameters regulating ECM biomass and decomposition. Under low N availability, the optimal C allocation to ECM fungi, above which the symbiosis switches from mutualism to parasitism, increases with increasing relative involvement of ECM fungi in SOM decomposition. Under low N conditions, increased ECM organic N mining promotes tree growth but decreases soil C storage, leading to a negative correlation between C stores above- and below-ground. The interplay between plant production and soil C storage is sensitive to the partitioning of decomposition between ECM fungi and saprotrophs. Better understanding of interactions between functional guilds of soil fungi may significantly improve predictions of ecosystem responses to environmental change.

  • 2017. Stefano Manzoni (et al.). Ecology Letters 20 (9), 1182-1191

    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.

  • 2017. C. A. Sierra (et al.). Global Change Biology 23 (5), 1763-1773

    Comparisons among ecosystem models or ecosystem dynamics along environmental gradients commonly rely on metrics that integrate different processes into a useful diagnostic. Terms such as age, turnover, residence, and transit times are often used for this purpose; however, these terms are variably defined in the literature and in many cases, calculations ignore assumptions implicit in their formulas. The aim of this opinion piece was i) to make evident these discrepancies and the incorrect use of formulas, ii) highlight recent results that simplify calculations and may help to avoid confusion, and iii) propose the adoption of simple and less ambiguous terms.

  • 2015. Allen G. Hunt, Manzoni Stefano.

    This book presents research into the physical rules that can underlie the behaviour of biota, as well as the geochemistry of soil development. It addresses both nutrient and water transport limitations of processes from chemical weathering to vascular plant growth. It attempts to bring order to the question of the extent to which soils can facilitate plant growth, and what limitations on plant sizes, metabolism, occurrence and correlations can be formulated thereby.

  • 2020. Stefano Manzoni, Tanya Strydom. Global Change Biology 26 (1), 119-188

    Plant traits-the morphological, anatomical, physiological, biochemical and phenological characteristics of plants-determine how plants respond to environmental factors, affect other trophic levels, and influence ecosystem properties and their benefits and detriments to people. Plant trait data thus represent the basis for a vast area of research spanning from evolutionary biology, community and functional ecology, to biodiversity conservation, ecosystem and landscape management, restoration, biogeography and earth system modelling. Since its foundation in 2007, the TRY database of plant traits has grown continuously. It now provides unprecedented data coverage under an open access data policy and is the main plant trait database used by the research community worldwide. Increasingly, the TRY database also supports new frontiers of trait-based plant research, including the identification of data gaps and the subsequent mobilization or measurement of new data. To support this development, in this article we evaluate the extent of the trait data compiled in TRY and analyse emerging patterns of data coverage and representativeness. Best species coverage is achieved for categorical traits-almost complete coverage for 'plant growth form'. However, most traits relevant for ecology and vegetation modelling are characterized by continuous intraspecific variation and trait-environmental relationships. These traits have to be measured on individual plants in their respective environment. Despite unprecedented data coverage, we observe a humbling lack of completeness and representativeness of these continuous traits in many aspects. We, therefore, conclude that reducing data gaps and biases in the TRY database remains a key challenge and requires a coordinated approach to data mobilization and trait measurements. This can only be achieved in collaboration with other initiatives.

  • 2019. Andreas Hagenbo (et al.). Journal of Ecology 107 (6), 2808-2822

    In boreal forest soils, mycelium of mycorrhizal fungi is pivotal for regulating soil carbon (C) cycling and storage. The carbon use efficiency (CUE), a key parameter in C cycling models, can inform on the partitioning of C between microbial biomass, and potential soil storage, and respiration. Here, we test the dependency of mycorrhizal mycelial CUE on stand age and seasonality in managed boreal forest stands. Based on mycelial production and respiration estimates, derived from sequentially incubated ingrowth mesh bags, we estimated CUE on an ecosystem scale during a seasonal cycle and across a chronosequence of eight, 12- to 158-year-old, managed Pinus sylvestris forest stands characterized by decreasing pH and nitrogen (N) availability with increasing age. Mycelial respiration was related to total soil respiration, and by using eddy covariance flux measurements, primary production (GPP) was estimated in the 12- and 100-year-old forests, and related to mycelial respiration and CUE. As hypothesized, mycelial CUE decreased significantly with increasing forest age by c. 65%, supposedly related to a shift in mycorrhizal community composition and a metabolic adjustment reducing their own biomass N demand with declining soil N availability. Furthermore, mycelial CUE increased by a factor of five over the growing season; from 0.03 in May to 0.15 in November, and we propose that the seasonal change in CUE is regulated by a decrease in photosynthate production and temperature. The respiratory contribution of mycorrhizal mycelium ranged from 14% to 26% of total soil respiration, and was on average 17% across all sites and occasions. Synthesis. Carbon is retained more efficiently in mycorrhizal mycelium late in the growing season, when fungi have access to a more balanced C and nutrient supplies. Earlier in the growing season, at maximum host plant photosynthesis, when below-ground C availability is high in relation to N, the fungi respire excess C resulting in lower mycelial carbon use efficiency (CUE). Additionally, C is retained less efficiently in mycorrhizal fungal biomass in older forest stands characterized by more nutrient depleted soils than younger forest stands.

  • 2019. Guoyi Zhou (et al.).

    Soil organic carbon (SOC) plays critical roles in stabilizing atmospheric CO2 concentration, but the mechanistic controls on the amount and distribution of SOC on global scales are not well understood. In turn, this has hampered the ability to model global C budgets and to find measures to mitigate climate change. Here, based on the data from a large field survey campaign with 2600 plots across China's forest ecosystems and a global collection of published data from forested land, we find that a low litter carbon-to-nitrogen ratio (C/N) and high wetness index (P/PET, precipitation-to-potential-evapotranspiration ratio) are the two factors that promote SOC accumulation, with only minor contributions of litter quantity and soil texture. The field survey data demonstrated that high plant diversity decreased litter C/N and thus indirectly promoted SOC accumulation by increasing the litter quality. We conclude that any changes in plant-community composition, plant-species richness and environmental factors that can reduce the litter C/N ratio, or climatic changes that increase wetness index, may promote SOC accumulation. The study provides a guideline for modeling the carbon cycle of various ecosystem scales and formulates the principle for land-based actions for mitigating the rising atmospheric CO2 concentration.

Show all publications by Stefano Manzoni at Stockholm University

Last updated: May 11, 2020

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