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. Plant nutrients can be recycled through microbialdecomposition of organic matter but replacement of base cations andphosphorus, lost through harvesting of biomass/biofuels or leaching,requires de novo supply of fresh nutrients released through weathering of soilparent material (minerals and rocks). Weathering involves physical andchemical processes that are modified by biological activity of plants,microorganisms and animals. This article reviews recent progress made inunderstanding biological processes contributing to weathering. A perspectiveof increasing spatial scale is adopted, examining the consequences ofbiological activity for weathering from nanoscale interactions, through in vitro andin planta microcosm and mesocosm studies, to field experiments, and finally ecosystemand global level effects. The topics discussed include the physicalalteration of minerals and mineral surfaces; the composition, amounts,chemical properties, and effects of plant and microbial secretions; and therole of carbon flow (including stabilisation and sequestration of C in organicand inorganic forms). Although the predominant focus is on the effects offungi in forest ecosystems, the properties of biofilms, including bacterialinteractions, are also discussed. The implications of these biologicalprocesses for modelling are discussed, and we attempt to identify some keyquestions and knowledge gaps, as well as experimental approaches and areasof research in which future studies are likely to yield useful results. Aparticular focus of this article is to improve the representation of theways in which biological processes complement physical and chemicalprocesses that mobilise mineral elements, making them available for plantuptake. This is necessary to produce better estimates of weathering that arerequired for sustainable management of forests in a post-fossil-fueleconomy. While there are abundant examples of nanometre- and micrometre-scalephysical interactions between microorganisms and different minerals, opinionappears to be divided with respect to the quantitative significance of theseobservations for overall weathering. Numerous in vitro experiments and microcosmstudies involving plants and their associated microorganisms suggest thatthe allocation of plant-derived carbon, mineral dissolution and plantnutrient status are tightly coupled, but there is still disagreement aboutthe extent to which these processes contribute to field-scale observations.Apart from providing dynamically responsive pathways for the allocation ofplant-derived carbon to power dissolution of minerals, mycorrhizal myceliaprovide conduits for the long-distance transportation of weathering productsback to plants that are also quantitatively significant sinks for releasednutrients. These mycelial pathways bridge heterogeneous substrates, reducingthe influence of local variation in C:N ratios. The production ofpolysaccharide matrices by biofilms of interacting bacteria and/or fungi atinterfaces with mineral surfaces and roots influences patterns ofproduction of antibiotics and quorum sensing molecules, with concomitanteffects on microbial community structure, and the qualitative andquantitative composition of mineral-solubilising compounds and weatheringproducts. Patterns of carbon allocation and nutrient mobilisation from bothorganic and inorganic substrates have been studied at larger spatial andtemporal scales, including both ecosystem and global levels, and there is agenerally wider degree of acceptance of the “systemic” effects ofmicroorganisms on patterns of nutrient mobilisation. Theories about theevolutionary development of weathering processes have been advanced butthere is still a lack of information connecting processes at differentspatial scales. Detailed studies of the liquid chemistry of local weatheringsites at the micrometre scale, together with upscaling to soil-scaledissolution rates, are advocated, as well as new approaches involving stableisotopes.

Reviews and syntheses: Biological weathering and its consequences at different spatial levels – from nanoscale to global scale