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Evaluation of Methods to Estimate Production, Biomass and Turnover of Ectomycorrhizal Mycelium in Forests Soils - a Review H

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Evaluation of Methods to Estimate Production, Biomass and Turnover of Ectomycorrhizal Mycelium in Forests Soils - a Review H Evaluation of methods to estimate production, biomass and turnover of ectomycorrhizal mycelium in forests soils - A review H. Wallander, A. Ekblad, D. L. Godbold, D. Johnson, A. Bahr, P. Baldrian, R. G. Bjork, B. Kieliszewska-Rokicka, R. Kjoller, H. Kraigher, et al. To cite this version: H. Wallander, A. Ekblad, D. L. Godbold, D. Johnson, A. Bahr, et al.. Evaluation of methods to estimate production, biomass and turnover of ectomycorrhizal mycelium in forests soils - A review. Soil Biology and Biochemistry, Elsevier, 2013, 57, pp.1034 - 1047. 10.1016/j.soilbio.2012.08.027. hal-01268191 HAL Id: hal-01268191 https://hal.archives-ouvertes.fr/hal-01268191 Submitted on 29 May 2020 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Soil Biology & Biochemistry 57 (2013) 1034e1047 Contents lists available at SciVerse ScienceDirect Soil Biology & Biochemistry journal homepage: www.elsevier.com/locate/soilbio Review Evaluation of methods to estimate production, biomass and turnover of ectomycorrhizal mycelium in forests soils e A review H. Wallander a,*, A. Ekblad b, D.L. Godbold c, D. Johnson d, A. Bahr a, P. Baldrian e, R.G. Björk f, B. Kieliszewska-Rokicka g, R. Kjøller h, H. Kraigher i, C. Plassard j, M. Rudawska k a Department of Biology, Microbial Ecology Group, Ecology Building, Lund University, SE-223 62 Lund, Sweden b School of Science & Technology, Örebro University, SE-701 82 Örebro, Sweden c Institute of Forest Ecology, Universität für Bodenkultur, 1190 Vienna, Austria d Institute of Biological and Environmental Sciences, University of Aberdeen, Cruickshank Building, St. Machar Drive, Aberdeen AB24 3UU, UK e Laboratory of Environmental Microbiology, Institute of Microbiology ASCR, CZ-14220 Praha, Czech Republic f Department of Biological and Environmental Sciences, University of Gothenburg, P.O. Box 461, SE-405 30 Gothenburg, Sweden g Institute of Environmental Biology, Kazimierz Wielki University, Al. Ossolinskich 12, PL-85-093 Bydgoszcz, Poland h Terrestrial Ecology, Biological Institute, University of Copenhagen, Øster Farimagsgade 2D, DK-1353 Copenhagen, Denmark i Slovenian Forestry Institute, Vecna pot 2, 1000 Ljubljana, Slovenia j INRA, UMR Eco & Sols, 34060 Montpellier Cedex 02, France k Institute of Dendrology, Polish Academy of Sciences, Parkowa 5, 62-035 Kórnik, Poland article info abstract Article history: Mycorrhizal fungi constitute a considerable sink for carbon in most ecosystems. This carbon is used for Received 2 March 2012 building extensive mycelial networks in the soil as well as for metabolic activity related to nutrient Received in revised form uptake. A number of methods have been developed recently to quantify production, standing biomass 22 August 2012 and turnover of extramatrical mycorrhizal mycelia (EMM) in the field. These methods include mini- Accepted 23 August 2012 rhizotrons, in-growth mesh bags and cores, and indirect measurements of EMM based on classification of Available online 10 September 2012 ectomycorrhizal fungi into exploration types. Here we review the state of the art of this methodology and discuss how it can be developed and applied most effectively in the field. Furthermore, we also discuss Keywords: Chitin different ways to quantify fungal biomass based on biomarkers such as chitin, ergosterol and PLFAs, as Exploration type well as molecular methods, such as qPCR. The evidence thus far indicates that mycorrhizal fungi are key Ergosterol components of microbial biomass in many ecosystems. We highlight the need to extend the application Extramatrical mycelium of current methods to focus on a greater range of habitats and mycorrhizal types enabling incorporation In-growth bag of mycorrhizal fungal biomass and turnover into biogeochemical cycling models. Minirhizotron Ó 2012 Elsevier Ltd. Open access under CC BY-NC-ND license. PLFA Rhizomorphs Sampling design Turnover rates 1. Introduction activity in the forest canopy to the activity in the soil, and provides a flow of organic C from shoots to soil via fine roots and mycorrhizal A better understanding of below ground carbon (C) flux is of hyphae. The pathways by which this organic C can enter soils are fundamental importance to predict how changing climate will complex, involving both biomass turnover (Godbold et al., 2003), influence the C balance of forest (and other) ecosystems (Litton and biomass grazing (Setälä et al., 1999) and turnover of low molecular Giardina, 2008). Litton et al. (2007) reported below ground C allo- weight exudates from roots and fungal hyphae (van Hees et al., cation in forest ecosystems can represent 25e63% of GPP on a global 2005). The fate of C entering soil systems is also complex. Much of scale, and this C has a large influence on the physical, chemical and this C is lost as respiration (Janssens et al., 2001) and a small but biological properties of soils. The below ground allocation of C links significant fraction enters the soil organic matter (SOM) pool. Determination of the pools and fluxes of biomass inputs in isolation from fine roots and mycorrhiza provides a major scientific challenge. Some studies (e.g. Wallander et al., 2004) suggest that biomass pools * Corresponding author. MEMEG, Department of Biology, Ecology Building, Lund and inputs from fine roots and mycorrhizal hyphae are in the same University, SE-223 62 Lund, Sweden. Tel.: þ46 46 222 4247; fax: þ46 46 222 4158. E-mail address: [email protected] (H. Wallander). order of magnitude. However, estimates of fungal inputs rely on 0038-0717 Ó 2012 Elsevier Ltd. Open access under CC BY-NC-ND license. http://dx.doi.org/10.1016/j.soilbio.2012.08.027 H. Wallander et al. / Soil Biology & Biochemistry 57 (2013) 1034e1047 1035 methods and conversion factors that contain a certain degree of assess the turnover of fungal hyphae. This area needs clearly to be inaccuracy that needs to be considered. developed in future research as it is a key process in C sequestration Precise measurements of production, standing biomass and of forest soils. We also include aspects of sampling strategies and turnover of extramatrical mycelium (EMM) of mycorrhizal fungi are indirect estimates of EMM production that have great potential for essential in order to accurately describe the C cycle of terrestrial the future. The mechanisms through which EMM regulate C cycling ecosystems. Although several techniques are available for this, they in terrestrial ecosystems have been considered recently in another all have limitations that need to be taken into consideration. review (Cairney, 2012). Existing biogeochemical models often treat the uptake appa- ratus as a single organ, meaning that there is no distinction between 2. Measurements of mycorrhizal hyphal production roots and mycorrhizal hyphae. It is possible, and probably neces- sary, to amend this by allocating carbon and nutrients specifically A key problem in the determination of mycorrhizal hyphal for the fine roots and mycorrhizal hyphae respectively. This would production is lack of methods to distinguish growth of mycorrhizal require the development of dynamic allocation routines responsive hyphae from that of saprotrophic fungi. As ECM fungi do not form to carbon, nutrients and water availability (Jönsson, 2006), and a monophyletic clade (Hibbett et al., 2000; Tedersoo et al., 2010)no would allow the models to simulate nutrient uptake and carbon flux single biochemical or DNA based marker can be found to quantify this dynamically. In this review, we will discuss and compare available group from the complex soil environment. Therefore various methods methods to estimate production, standing biomass and turnover of are needed to distinguish the biomass of EMM from that of other mycorrhizal mycelia (summarized in Tables 1e3). We focus on fungal mycelia. Mycelial growth can be estimated by direct observa- temperate and boreal forests, in which the dominant plants asso- tion in minirhizotrons (Treseder et al., 2005; Pritchard et al., 2008; ciate with ectomycorrhizal (ECM) fungi. From a methodological Vargas and Allen, 2008a) and by the use of root free in-growth bags or perspective, greatest progress has been made in quantification of cores, which is the most commonly applied method to measure EMM production, biomass and turnover of ECM fungi compared to the production in forests (Wallander et al., 2001; Godbold et al., 2006; other main mycorrhizal types (arbuscular and ericoid mycorrhizas). Hendricks et al., 2006; Kjøller, 2006; Korkama et al., 2007; Parrent and This progress has been driven partly by technical reasons but more Vilgalys, 2007; Hedh et al., 2008; Majdi et al., 2008). importantly because of the recognition of the key roles boreal and temperate forests play in the global C cycle. However, we emphasise 2.1. Observational methods from the outset that greater effort must be applied to other ecosystems in which plants are primarily colonised by arbuscular The first observational studies used plastic sheets placed at the and ericoid mycorrhizal fungi, which also have important roles in litter/soil interface above
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