Over the last 50 years, the use of nitrogen (N) and phosphorus (P) fertilizers has increased at a faster rate than global food production, resulting in substantial decreases of N and P efficiency in agriculture (Fig. 1). Thus, considerable amounts of N and P fertilizers have been wasted in agroecosystems, and the alteration of N and P biogeochemical cycles is among the most visible impacts of human activities on ecosystem services, far exceeding the hypothesized "planetary boundaries" for N and approaching those for P (Rockström et al., 2009). Projections for the future suggest that a substantial further increase in N and P fertilizer use will occur to cope with increasing food demand (Tilman et al., 2001;Vance et al., 2003). Tilman et al. (2001) predicted that global consumption of fertilizer P will increase from 34.
There is growing evidence of the importance of extramatrical mycelium (EMM) of mycorrhizal fungi in carbon (C) cycling in ecosystems. However, our understanding has until recently been mainly based on laboratory experiments, and knowledge of such basic parameters as variations in mycelial production, standing biomass and turnover as well as the regulatory mechanisms behind such variations in forest soils is limited. Presently, the production of EMM by ectomycorrhizal (EM) fungi has been Plant Soil (2013) at~140 different forest sites to be up to several hundreds of kg per ha per year, but the published data are biased towards Picea abies in Scandinavia. Little is known about the standing biomass and turnover of EMM in other systems, and its influence on the C stored or lost from soils. Here, focussing on ectomycorrhizas, we discuss the factors that regulate the production and turnover of EMM and its role in soil C dynamics, identifying important gaps in this knowledge. C availability seems to be the key factor determining EMM production and possibly its standing biomass in forests but direct effects of mineral nutrient availability on the EMM can be important. There is great uncertainty about the rate of turnover of EMM. There is increasing evidence that residues of EM fungi play a major role in the formation of stable N and C in SOM, which highlights the need to include mycorrhizal effects in models of global soil C stores.
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