Staff Publications

Staff Publications

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    'Staff publications' is the digital repository of Wageningen University & Research

    'Staff publications' contains references to publications authored by Wageningen University staff from 1976 onward.

    Publications authored by the staff of the Research Institutes are available from 1995 onwards.

    Full text documents are added when available. The database is updated daily and currently holds about 240,000 items, of which 72,000 in open access.

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    Nitrogen deposition is the most important environmental driver of growth of pure, even-aged and managed European forests
    Etzold, Sophia ; Ferretti, Marco ; Reinds, Gert Jan ; Solberg, Svein ; Gessler, Arthur ; Waldner, Peter ; Schaub, Marcus ; Simpson, David ; Benham, Sue ; Hansen, Karin ; Ingerslev, Morten ; Jonard, Mathieu ; Karlsson, Per Erik ; Lindroos, Antti Jussi ; Marchetto, Aldo ; Manninger, Miklos ; Meesenburg, Henning ; Merilä, Päivi ; Nöjd, Pekka ; Rautio, Pasi ; Sanders, Tanja G.M. ; Seidling, Walter ; Skudnik, Mitja ; Thimonier, Anne ; Verstraeten, Arne ; Vesterdal, Lars ; Vejpustkova, Monika ; Vries, Wim de - \ 2020
    Forest Ecology and Management 458 (2020). - ISSN 0378-1127
    Air pollution - Climate change - Forest management - ICP Forests - Observational study - Ozone - Statistical modelling

    Changing environmental conditions may substantially interact with site quality and forest stand characteristics, and impact forest growth and carbon sequestration. Understanding the impact of the various drivers of forest growth is therefore critical to predict how forest ecosystems can respond to climate change. We conducted a continental-scale analysis of recent (1995–2010) forest volume increment data (ΔVol, m3 ha−1 yr−1), obtained from ca. 100,000 coniferous and broadleaved trees in 442 even-aged, single-species stands across 23 European countries. We used multivariate statistical approaches, such as mixed effects models and structural equation modelling to investigate how European forest growth respond to changes in 11 predictors, including stand characteristics, climate conditions, air and site quality, as well as their interactions. We found that, despite the large environmental gradients encompassed by the forests examined, stand density and age were key drivers of forest growth. We further detected a positive, in some cases non-linear effect of N deposition, most pronounced for beech forests, with a tipping point at ca. 30 kg N ha−1 yr−1. With the exception of a consistent temperature signal on Norway spruce, climate-related predictors and ground-level ozone showed much less generalized relationships with ΔVol. Our results show that, together with the driving forces exerted by stand density and age, N deposition is at least as important as climate to modulate forest growth at continental scale in Europe, with a potential negative effect at sites with high N deposition.

    Natuurrapportage Zeeland 2019
    Boddeke, Paul ; Boudewijn, Theo ; Helsdingen, Arda van; Lensink, Rob ; Röell, Ineke ; Roelofsen, Hans ; Sanders, Marlies ; Schütt, Jeroen ; Wamelink, Wieger ; Zee, Friso van der; Soomers, Hester - \ 2019
    Middelburg : Provincie Zeeland - 153
    Reduction of antibiotic use and antibiotic resistance in commercial poultry
    Veldman, K.T. ; Sanders, Pim ; Mevius, D.J. - \ 2019
    In: International Symposium on Avian Mycoplasmosis and Infectious Coryza 2019 (ISMC 2019). - Deventer : CVI / GD - p. 86 - 91.
    Use of antibiotics for companion animals and livestock in the Netherlands has reduced by more than 60% over the last 10 years (SDa 2019; MARAN-2019). This reduction is the result of a change in policy towards the use of antibiotics in veterinary practice and is characterized by a series of coherent political decisions which changed the playing field for farmers and veterinarians considerably. In the years before 2009 the Netherlands was a high consumer of antibiotics in veterinary practice (Grave et al., 2010). The ban of antimicrobial growth promoters (AGPs) did not result in a reduction in total use since in the Netherlands the AGPs were fully replaced by antibiotics licensed for therapy. The total sales of all antibiotics remained stable at around 600 tons from 2000 to 2009.This use pattern resulted in high levels of antimicrobial resistance in bacteria from livestock and food thereof and high prevalence of Livestock Associated MRSA and ESBL-producing E. coli and Salmonella (MARAN, 2019; RIVM, 2009). Specifically, ESBL-producing isolates in the food chain were considered a risk for public health and their high prevalences, predominantly but not solely in poultry and poultry meat products were the direct reason to initiate the change in policy towards antibiotic use in animals.In this manuscript the trends in antibiotic use in poultry will be explored in the context of total use in livestock and its effect on the occurrence and trends in ESBL-producers and antimicrobial resistance in other bacterial species from poultry.
    Basisbestand Natuur en Landschap
    Sanders, M.E. ; Meeuwsen, H.A.M. - \ 2019
    Wageningen : Wettelijke Onderzoekstaken Natuur & Milieu (WOt-technical report 158) - 101
    In its work for studies by the Netherlands Environmental Assessment Agency (PBL), such as the Assessment of the Dutch Human Environment, the Environmental Data Compendium, the Natuurpact reflexive evaluation and the Nature Outlook, the Statutory Research Tasks Unit for Nature & the Environment (WOt N&M) makes use of various base maps of natural and semi-natural habitats, such as the Nature Base Map (BKN), the Land Use Database of the Netherlands (LGN database) and the map of habitat management types (Beheertypenkaart). The problem is that information derived from these maps can influence the indicators and model results used in the PBL studies and lead to inconsistencies between them. This report describes the design of a standardised procedure for compiling a Geodatabase of Nature and Landscape (Basisbestand Natuur en Landschap – BNL) that can be regularly updated. The BNL is a raster database with a resolution of 2.5 metres. It consists of several layers and is generated using a GIS script. The geometric basis of the map is Top10NL, which is considered to be a reflection of the true situation in the field. The management type map produced by the provincial governments (IMNaB) was used as a source database for the areas and types of natural and semi-natural habitats. This map is a key resource in Dutch nature policy for the allocation of subsidies for conservation management and for determining ecological quality. The aim is to obtain ‘Status A’ for the BNL under the WOT Quality Control system.
    Towards a New Generation of Trait-Flexible Vegetation Models
    Berzaghi, Fabio ; Wright, Ian J. ; Kramer, Koen ; Oddou-Muratorio, Sylvie ; Bohn, Friedrich J. ; Reyer, Christopher P.O. ; Sabaté, Santiago ; Sanders, Tanja G.M. ; Hartig, Florian - \ 2019
    Trends in Ecology and Evolution (2019). - ISSN 0169-5347
    eco-evolution - intraspecific variation - plant genetics - plant traits - vegetation modeling

    Plant trait variability, emerging from eco-evolutionary dynamics that range from alleles to macroecological scales, is one of the most elusive, but possibly most consequential, aspects of biodiversity. Plasticity, epigenetics, and genetic diversity are major determinants of how plants will respond to climate change, yet these processes are rarely represented in current vegetation models. Here, we provide an overview of the challenges associated with understanding the causes and consequences of plant trait variability, and review current developments to include plasticity and evolutionary mechanisms in vegetation models. We also present a roadmap of research priorities to develop a next generation of vegetation models with flexible traits. Including trait variability in vegetation models is necessary to better represent biosphere responses to global change.

    Methodenontwikkeling evaluatie (kosten)effectiviteit : Tussenrapportage WOT-04-010-037.12
    Wit-de Vries, E. de; Mattijssen, T.J.M. ; Nuesink, Nienke ; Michels, R. ; Koeijer, T.J. de; Schütt, Jeroen ; Sanders, M.E. ; Hinsberg, Arjen van - \ 2019
    WOT Natuur & Milieu (WOt-interne notitie 293)
    Voorbereiding rapportage Balans. : Tussenrapportage WOT-04-010-033
    Sanders, M.E. ; Langers, F. ; Schütt, Jeroen ; Mensing, Victor - \ 2019
    Data from: Virus Shedding of Avian Influenza in Poultry: A Systematic Review and Meta-Analysis
    Germeraad, Evelien ; Sanders, Pim ; Gonzales Rojas, Jose - \ 2019
    Wageningen University & Research
    avian influenza - meta-analysis - poultry - systematic review - virus shedding
    These systematic review and meta-analysis were performed to summarize qualitative and quantitative information on virus shedding levels and duration for different AIV strains in experimentally infected poultry species.
    Method for producing inositol and inorganic phosphate
    Sanders, J.P.M. ; Boeriu, C.G. ; Yilmaz, G. - \ 2019
    Octrooinummer: WO2019221602, gepubliceerd: 2019-11-21.
    The present invention relates to a method for producing inositol and inorganic phosphate, comprising the steps of (a) providing a composition comprising phytate, phytic acid, and/or phytin; (b) hydrolysing the phytate, phytic acid, and/or phytin to provide a composition comprising inositol phosphate and inorganic phosphate; (c) separating inorganic phosphate from the composition provided in step (b) to provide a composition enriched in inositol phosphate and a composition comprising inorganic phosphate; (d) hydrolysing the inositol phosphate obtained in step (c) in order to provide a composition comprising inositol and inorganic phosphate.
    Convention on biological diversity : Sixth national report of the Kingdom of the Netherlands
    Sanders, M.E. ; Henkens, R.J.H.G. ; Slijkerman, D.M.E. - \ 2019
    Wageningen : Statutory Research Tasks Unit for Nature & the Environment (WOt technical report 156) - 136
    The Netherlands is one of the most densely populated countries in the world. Most of the country’s land area is highly productive agricultural land. There is little or no area of natural ecosystem that could be called pristine,although high biodiversity values are certainly found. TheKingdom of the Netherlands also comprises six Caribbeanislands with, among other things, tropical rainforests,mangrove forests and coral reefs. The main cause of the long-term decline biodiversity has been the intensification of agricultural production, including the reclamation of semi-natural areas, the drainage of wet areas, the use of artificial fertilizer, etc. But also unsustainable fishing,pollution, overgrazing, climate change and invasive alien species cause a decline in biodiversity. To reverse this decline, the Netherlands has prepared and implemented action plans and targets for biodiversity since 1990. This report presents the most important efforts to achieve the biodiversity targets.
    Virus Shedding of Avian Influenza in Poultry: A Systematic Review and Meta-Analysis
    Germeraad, Evelien A. ; Sanders, Pim ; Hagenaars, Thomas J. ; Jong, Mart C.M. de; Beerens, Nancy ; Gonzales, Jose L. - \ 2019
    Viruses 11 (2019)9. - ISSN 1999-4915
    avian influenza - meta-analysis - poultry - systematic review - virus shedding

    Understanding virus shedding patterns of avian influenza virus (AIV) in poultry is important for understanding host-pathogen interactions and developing effective control strategies. Many AIV strains were studied in challenge experiments in poultry, but no study has combined data from those studies to identify general AIV shedding patterns. These systematic review and meta-analysis were performed to summarize qualitative and quantitative information on virus shedding levels and duration for different AIV strains in experimentally infected poultry species. Methods were designed based on the Preferred Reporting Items for Systematic Reviews and Meta-Analyses (PRISMA) guidelines. Four electronic databases were used to collect literature. A total of 1155 abstract were screened, with 117 studies selected for the qualitative analysis and 71 studies for the meta-analysis. A large heterogeneity in experimental methods was observed and the quantitative analysis showed that experimental variables such as species, virus origin, age, inoculation route and dose, affect virus shedding (mean, peak and duration) for highly pathogenic AIV (HPAIV), low pathogenic AIV (LPAIV) or both. In conclusion, this study highlights the need to standardize experimental procedures, it provides a comprehensive summary of the shedding patterns of AIV strains by infected poultry and identifies the variables that influence the level and duration of AIV shedding.

    Validation of an ultra-high-performance liquid chromatography-mass spectrometry method for the quantification of cysteinylated aldehydes and application to malt and beer samples
    Bustillo Trueba, P. ; Jaskula-Goiris, B. ; Clippeleer, J. De; Goiris, K. ; Praet, T. ; Sharma, U.K. ; Eycken, E. Van der; Sanders, M.G. ; Vincken, J.P. ; Brabanter, J. De; Rouck, G. De; Aerts, G. ; Cooman, L. De - \ 2019
    Journal of Chromatography. A, Including electrophoresis and other separation methods 1604 (2019). - ISSN 0021-9673
    2-substituted 1,3-thiazolidine-4-carboxylic acids - Beer - Cysteinylated aldehydes - Liquid chromatography-mass spectrometry - Malt - Method validation

    This paper describes the method validation for the simultaneous determination of seven cysteinylated aldehydes, i.e. 2-substituted 1,3-thiazolidines-4-carboxylic acids, using ultra-high-performance liquid chromatography-mass spectrometry (UHPLC–MS). Authentic reference compounds were first synthesized for identification and quantification purposes. Moreover, nuclear magnetic resonance (1H NMR and 13C NMR) was applied for verification of their structure, while ultra-high-performance liquid chromatography–mass spectrometry (UHPLC–MS) was applied for estimation of the purity. The method for quantification of cysteinylated aldehydes in model solutions has been validated according to the criteria and procedures described in international standards. The synthesized compounds were successfully identified via UHPLC–MS by comparing retention time and MS spectra with the commercial reference compounds. Method validation revealed good linearity (R2 > 0.995) over the range of 0.4–2.2 µg/L to approximately 1000 µg/L, depending on the analyte. The limits of quantification varied from 0.9 to 4.3 µg/L depending on the nature of the compound. Furthermore, evaluation of the method showed good accuracy and stability of the standard solutions. Reported chromatographic recoveries ranged from 112 to 120%. Consequently, the currently described method was applied on malt and beer samples. For the first time, quantification of cysteinylated aldehydes was obtained in malt. In contrast, in fresh beers unambiguous identification of these compounds was not achieved.

    Pilot grondgebondenheid West-Achterhoek : Over huiskavels en eiwitvoorziening op het eigen melkveebedrijf
    Sanders, Willem Jan ; Hubeek, Alwin ; Vogelzang, Theo ; Doornewaard, Gerben ; Prins, Henry - \ 2019
    Kadaster - 66 p.
    ‘Vergroening chemische industrie vereist radicale keuzes’: hernieuwbare grondstoffen, maar ook nieuwe processen
    Sanders, J.P.M. - \ 2019
    biobased economy - chemical industry - netherlands - innovation adoption - renewable energy - biomass
    Joint sequencing of human and pathogen genomes reveals the genetics of pneumococcal meningitis
    Lees, John A. ; Ferwerda, Bart ; Kremer, Philip H.C. ; Wheeler, Nicole E. ; Valls Serón, Mercedes ; Croucher, Nicholas J. ; Gladstone, Rebecca A. ; Bootsma, Hester J. ; Rots, Nynke Y. ; Wijmega-Monsuur, Alienke J. ; Sanders, Elisabeth A.M. ; Trzciński, Krzysztof ; Wyllie, Anne L. ; Zwinderman, Aeilko H. ; Berg, Leonard H. van den; Rheenen, Wouter van; Veldink, Jan H. ; Harboe, Zitta B. ; Lundbo, Lene F. ; Groot, Lisette C.P.G.M. de; Schoor, Natasja M. van; Velde, Nathalie van der; Ängquist, Lars H. ; Sørensen, Thorkild I.A. ; Nohr, Ellen A. ; Mentzer, Alexander J. ; Mills, Tara C. ; Knight, Julian C. ; Plessis, Mignon du; Nzenze, Susan ; Weiser, Jeffrey N. ; Parkhill, Julian ; Madhi, Shabir ; Benfield, Thomas ; Gottberg, Anne von; Ende, Arie van der; Brouwer, Matthijs C. ; Barrett, Jeffrey C. ; Bentley, Stephen D. ; Beek, Diederik van de - \ 2019
    Nature Communications 10 (2019)1. - ISSN 2041-1723 - 14 p.

    Streptococcus pneumoniae is a common nasopharyngeal colonizer, but can also cause life-threatening invasive diseases such as empyema, bacteremia and meningitis. Genetic variation of host and pathogen is known to play a role in invasive pneumococcal disease, though to what extent is unknown. In a genome-wide association study of human and pathogen we show that human variation explains almost half of variation in susceptibility to pneumococcal meningitis and one-third of variation in severity, identifying variants in CCDC33 associated with susceptibility. Pneumococcal genetic variation explains a large amount of invasive potential (70%), but has no effect on severity. Serotype alone is insufficient to explain invasiveness, suggesting other pneumococcal factors are involved in progression to invasive disease. We identify pneumococcal genes involved in invasiveness including pspC and zmpD, and perform a human-bacteria interaction analysis. These genes are potential candidates for the development of more broadly-acting pneumococcal vaccines.

    Method for extracting salt from a composition comprising organic matter
    Sanders, J.P.M. ; Meesters, K.P.H. - \ 2019
    Octrooinummer: WO2019011976, gepubliceerd: 2019-01-17.
    The present invention relates to a method for extracting salt and optionally one or more organic compounds from a composition comprising organic matter, and in particular from a dry or solid composition comprising organic matter. The present invention further relates to the use of the extracted salt and optionally one or more organic compounds, and/or of the remaining organic matter, for preparing foodstuff, a fertilizer, animal feed and/or a chemical.
    Dynamics of the Gut Microbiota in Children Receiving Selective or Total Gut Decontamination Treatment during Hematopoietic Stem Cell Transplantation
    Bekker, Vincent ; Zwittink, Romy D. ; Knetsch, Cornelis W. ; Sanders, Ingrid M.J.G. ; Berghuis, Dagmar ; Heidt, Peter J. ; Vossen, Jaak M.J.J. ; Vos, Willem M. de; Belzer, Clara ; Bredius, Robbert G.M. ; van‘t Hof, Peter J. ; Lankester, Arjan C. ; Kuijper, Ed J. - \ 2019
    Biology of Blood and Marrow Transplantation 25 (2019)6. - ISSN 1083-8791 - p. 1164 - 1171.
    Graft-versus-host disease - Gut decontamination - Hematopoietic stem cell transplantation - Microbiota - Pediatrics

    Bloodstream infections and graft-versus-host disease are common complications after hematopoietic stem cell transplantation (HSCT) procedures, associated with the gut microbiota that acts as a reservoir for opportunistic pathogens. Selective gut decontamination (SGD) and total gut decontamination (TGD) during HSCT have been associated with a decreased risk of developing these complications after transplantation. However, because studies have shown conflicting results, the use of these treatments remains subject of debate. In addition, their impact on the gut microbiota is not well studied. The aim of this study was to elucidate the dynamics of the microbiota during and after TGD and to compare these with the dynamics of SGD. In this prospective, observational, single-center study fecal samples were longitudinally collected from 19 children eligible for allogenic HSCT (TGD, n=12; SGD, n=7), weekly during hospital admission and monthly after discharge. In addition, fecal samples were collected from 3 family stem cell donors. Fecal microbiota structure of patients and donors was determined by 16S rRNA gene amplicon sequencing. Microbiota richness and diversity markedly decreased during SGD and TGD and gradually increased after cessation of decontamination treatment. During SGD, gut microbiota composition was relatively stable and dominated by Bacteroides, whereas it showed high inter- and intraindividual variation and low Bacteroides abundance during TGD. In some children TGD allowed the genera Enterococcus and Streptococcus to thrive during treatment. A gut microbiota dominated by Bacteroides was associated with increased predicted activity of several metabolic processes. Comparing the microbiota of recipients and their donors indicated that receiving an SCT did not alter the patient's microbiota to become more similar to that of its donor. Overall, our findings indicate that SGD and TGD affect gut microbiota structure in a treatment-specific manner. Whether these treatments affect clinical outcomes via interference with the gut microbiota needs to be further elucidated.

    Iron-polyphenol complexes cause blackening upon grinding Hermetia illucens (black soldier fly) larvae
    Janssen, Renske H. ; Canelli, Greta ; Sanders, Mark G. ; Bakx, Edwin J. ; Lakemond, Catriona M.M. ; Fogliano, Vincenzo ; Vincken, Jean Paul - \ 2019
    Scientific Reports 9 (2019). - ISSN 2045-2322

    Insects are a promising alternative protein source. One of the bottlenecks in applying insects in food is the fast darkening initiated during grinding. Besides enzymatic browning, non-enzymatic factors can cause off-colour formation, which differs between species. This study investigates the impact of iron, phenoloxidase, and polyphenols on off-colour formation in insect larvae. Hermetia illucens showed a blackish colour, whereas Tenebrio molitor turned brown and Alphitobius diaperinus remained the lightest. This off-colour formation appeared correlated with the iron content in the larvae, which was 61 ± 9.71, 54 ± 1.72 and 221 ± 6.07 mg/kg dw for T. molitor, A. diaperinus and H. illucens, respectively. In model systems, the formation of iron-L-3,4-dihydroxyphenylalanine (L-DOPA) bis- and tris-complexes were evidenced by direct injection into ESI-TOF-MS, based on their charges combined with iron isotope patterns. The reversibility of the binding of iron to phenolics, and thereby loss of blackening, was confirmed by EDTA addition. Besides complex formation, oxidation of L-DOPA by redox reactions with iron occurred mainly at low pH, whereas auto-oxidation of L-DOPA mainly occurred at pH 10. Tyrosinase (i.e. phenoloxidase) activity did not change complex formation. The similarity in off-colour formation between the model system and insects indicated an important role for iron-phenolic complexation in blackening.

    Prediction of soil pH patterns in nature areas on a national scale
    Wamelink, G.W.W. ; Walvoort, Dennis J.J. ; Sanders, Marlies E. ; Meeuwsen, Henk A.M. ; Wegman, Ruut M.A. ; Pouwels, Rogier ; Knotters, Martin - \ 2019
    Applied Vegetation Science 22 (2019)2. - ISSN 1402-2001 - p. 189 - 199.
    acidity - kriging - pH - relevé - soil type - vegetation

    Question: To assess the acidification process, nationwide information about soil pH on a site level is called for. Measurements of soil pH may be used, however there are not sufficient measurements available to map soil pH nationwide on site level. Instead we developed a soil pH map based on vegetation data. Location: Natural terrestrial areas in The Netherlands. Methods: 271,693 vegetation plots were used to estimate average soil pH per plot with indicator values, based on field measurements, of plant species. By spatial interpolation average pH values between the plots, with the soil type, groundwater table and vegetation management type as ancillary explanatory variables we created a soil pH map. The map covers all terrestrial nature areas (all areas that are not built up areas, agricultural areas and infrastructural areas) in the Netherlands with a map resolution of 25 × 25 m2 raster cells. Results: The predicted pH of the map varied between 3.0 and 8.6 with standard errors between 0.13 and 0.93. Most of the standard errors range from 0.4 to 0.55, with an average just below 0.5 pH unit. Cross-validation shows that for 33% the difference between observed and predicted is between −0.1 and 0.1 pH-unit and for 83% the difference is between −0.5 and 0.5 pH-unit. Validation shows that the pH map is unbiased (mean error is almost zero), accurate (root mean squared error is 0.64) and nicely captures spatial patterns (r = 0.77). We applied the pH map to assess the impact of acidification on the abiotic quality of nature areas in the Netherlands. Conclusions: The model fit in the predicted soil pH is in good resulting in a low standard error and a high correlation. The measures taken to prevent acidic deposition causing further acidifying of nature areas can be considered as successful.

    Responses of forest ecosystems in Europe to decreasing nitrogen deposition
    Schmitz, Andreas ; Sanders, Tanja G.M. ; Bolte, Andreas ; Bussotti, Filippo ; Dirnböck, Thomas ; Johnson, Jim ; Peñuelas, Josep ; Pollastrini, Martina ; Prescher, Anne Katrin ; Sardans, Jordi ; Verstraeten, Arne ; Vries, Wim de - \ 2019
    Environmental Pollution 244 (2019). - ISSN 0269-7491 - p. 980 - 994.
    Air pollution - Emission reduction - Forest monitoring - Nitrogen deposition - Recovery

    Average nitrogen (N) deposition across Europe has declined since the 1990s. This resulted in decreased N inputs to forest ecosystems especially in Central and Western Europe where deposition levels are highest. While the impact of atmospheric N deposition on forests has been receiving much attention for decades, ecosystem responses to the decline in N inputs received less attention. Here, we review observational studies reporting on trends in a number of indicators: soil acidification and eutrophication, understory vegetation, tree nutrition (foliar element concentrations) as well as tree vitality and growth in response to decreasing N deposition across Europe. Ecosystem responses varied with limited decrease in soil solution nitrate concentrations and potentially also foliar N concentrations. There was no large-scale response in understory vegetation, tree growth, or vitality. Experimental studies support the observation of a more distinct reaction of soil solution and foliar element concentrations to changes in N supply compared to the three other parameters. According to the most likely scenarios, further decrease of N deposition will be limited. We hypothesize that this expected decline will not cause major responses of the parameters analysed in this study. Instead, future changes might be more strongly controlled by the development of N pools accumulated within forest soils, affected by climate change and forest management. We find limited indication for response of Europe's forests to declining N deposition. Reactions have been reported for soil solution NO3 and potentially foliar N concentrations but not for other indicators.

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