Microbes are the unseen majority in soil and comprise a large portion of lifes genetic diversity. Despite their abundance, the impact of soil microbes on ecosystem processes is still poorly understood. Here we explore the various roles that soil microbes play in terrestrial ecosystems with special emphasis on their contribution to plant productivity and diversity. Soil microbes are important regulators of plant productivity, especially in nutrient poor ecosystems where plant symbionts are responsible for the acquisition of limiting nutrients. Mycorrhizal fungi and nitrogenfixing bacteria are responsible for c. 5‐20% (grassland and savannah) to 80% (temperate and boreal forests) of all nitrogen, and up to 75% of phosphorus, that is acquired by plants annually. Free-living microbes also strongly regulate plant productivity, through the mineralization of, and competition for, nutrients that sustain plant productivity. Soil microbes, including microbial pathogens, are also important regulators of plant community dynamics and plant diversity, determining plant abundance and, in some cases, facilitating invasion by exotic plants. Conservative estimates suggest that c. 20 000 plant species are completely dependent on microbial symbionts for growth and survival pointing to the importance of soil microbes as regulators of plant species richness on Earth. Overall, this review shows that soil microbes must be considered as important drivers of plant diversity and productivity in terrestrial ecosystems.

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The unseen majority: Soil microbes as drivers of plant diversity and productivity in terrestrial ecosystems

Publisher Summary In this chapter, the advances that have been made in understanding the ecology of the mineral nutrition of wild plants from terrestrial ecosystems have been reviewed. This chapter is organized along three lines. First, the issues of nutrient-limited plant growth and nutrient uptake, with special emphasis on the importance of the uptake of nutrients in organic form—both by mycorrhizal and by non-mycorrhizal plants—and the importance of symbiotic nitrogen fixation is treated. In addition, the influence of allocation patterns on mineral nutrient uptake is described. Next, a few of the nutritional aspects of leaf functioning and how nutrients are used for biomass production by the plant are explored. That is done by studying the nutrient use efficiency (NUE) of plants and the various components of NUE. Finally, the feedback of plant species to soil nutrient availability by reviewing patterns in litter decomposition and nutrient mineralization is investigated. The chapter concludes with a synthesis of the various aspects of the mineral nutrition of wild plants. The chapter ends with a conceptual description of plant strategies with respect to mineral nutrition.

The Mineral Nutrition of Wild Plants Revisited: A Re-evaluation of Processes and Patterns

Until recently, the common view of the terrestrial nitrogen cycle had been driven by two core assumptions—plants use only inorganic N and they compete poorly against soil microbes for N. Thus, plants were thought to use N that microbes “left over,” allowing the N cycle to be divided cleanly into two pieces—the microbial decomposition side and the plant uptake and use side. These were linked by the process of net mineralization. Over the last decade, research has changed these views. N cycling is now seen as being driven by the depolymerization of N-containing polymers by microbial (including mycorrhizal) extracellular enzymes. This releases organic N-containing monomers that may be used by either plants or microbes. However, a complete new conceptual model of the soil N cycle needs to incorporate recent research on plant–microbe competition and microsite processes to explain the dynamics of N across the wide range of N availability found in terrestrial ecosystems. We discuss the evolution of thinking abou...

Nitrogen mineralization: challenges of a changing paradigm

Nutrient limitation to primary productivity and other biological processes is widespread in terrestrial ecosystems, and nitrogen (N) and phosphorus (P) are the most common limiting elements, both individually and in combination. Mechanisms that drive P limitation, and their interactions with the N cycle, have received less attention than mechanisms causing N limitation. We identify and discuss six mechanisms that could drive P limitation in terrestrial ecosystems. The best known of these is depletion-driven limitation, in which accumulated P losses during long-term soil and ecosystem development contribute to what Walker and Syers termed a "terminal steady state" of profound P depletion and limitation. The other mechanisms are soil barriers that prevent access to P; transactional limitation, in which weathering of P-containing minerals does not keep pace with the supply of other resources; low-P parent materials; P sinks; and anthropogenic changes that increase the supply of other resources (often N) relative to P. We distinguish proximate nutrient limitation (which occurs where additions of a nutrient stimulate biological processes, especially productivity) from ultimate nutrient limitation (where additions of a nutrient can transform ecosystems). Of the mechanisms that drive P limitation, we suggest that depletion, soil barriers, and low-P parent material often cause ultimate limitation because they control the ecosystem mass balance of P. Similarly, demand-independent losses and constraints to N fixation can control the ecosystem-level mass balance of N and cause it to be an ultimate limiting nutrient.

Terrestrial phosphorus limitation: mechanisms, implications, and nitrogen–phosphorus interactions

Nature And Properties Of Soils

Plant growth in the boreal forest, the largest terrestrial biome, is generally limited by the availability of nitrogen. The presumed cause of this limitation is slow mineralization of soil organic nitrogen1,2. Here we demonstrate, to our knowledge for the first time, the uptake of organic nitrogen in the field by the trees Pinus sylvestris and Picea abies, the dwarf shrub Vaccinium myrtillus and the grass Deschampsia flexuosa. These results show that these plants, irrespective of their different types of root–fungal associations (mycorrhiza), bypass nitrogen mineralization. A trace of the amino acid glycine, labelled with the stable isotopes 13C and 15N, was injected into the organic (mor) layer of an old successional boreal coniferous forest. Ratios of 13C:15N in the roots showed that at least 91, 64 and 42% of the nitrogen from the absorbed glycine was taken up in intact glycine by the dwarf shrub, the grass and the trees, respectively. Rates of glycine uptake were similar to those of 15N-ammonium. Our data indicate that organic nitrogen is important for these different plants, even when they are competing with each other and with non-symbiotic microorganisms. This has major implications for our understanding of the effects of nitrogen deposition, global warming and intensified forestry.

Boreal forest plants take up organic nitrogen

Weatherable minerals under many European coniferous forests contain a network of numerous tubular pores, formed by organic acids exuded by fungi. We believe that symbiotic mycorrhizal hyphae translocate dissolved minerals from the isolated micropores directly to their host plants, bypassing competition for nutrient uptake by other organisms. The discovery of this pathway challenges current ideas about nutrient uptake from the bulk soil solution and criteria for critical loads of acidic deposition on forests.

Rock-eating fungi

The boreal forest is expected to experience the greatest warming of all forest biomes, raising concerns that some of the large quantities of soil carbon in these systems may be added to the atmosphere as CO2. However, nitrogen deposition or fertilization has the potential to increase boreal forest production and retard the decomposition of soil organic matter, hence increasing both tree stand and soil C storage.

The major contributors to soil-surface CO2 effluxes are autotrophic and heterotrophic respiration. To evaluate the effect of nutrient additions on the relative contributions from autotrophic and heterotrophic respiration, a large-scale girdling experiment was performed in a long-term nutrient optimization experiment in a 40-year-old stand of Norway spruce in northern Sweden. Trees on three nonfertilized plots and three fertilized plots were girdled in early summer 2002, and three nonfertilized and three fertilized plots were used as control plots. Each plot was 0.1 ha and contained around 230 trees. Soil-surface CO2 fluxes, soil moisture, and soil temperature were monitored in both girdled and nongirdled plots.

In late July, the time of the seasonal maximum in soil-surface CO2 efflux, the total soil-CO2 efflux in nongirdled plots was 40% lower in the fertilized than in the nonfertilized plots, while the efflux in girdled fertilized and nonfertilized plots was 50% and 60% lower, respectively, than in the corresponding nongirdled controls. We attribute these reductions to losses of the autotrophic component of the total soil-surface CO2 efflux. The estimates of autotrophic respiration are conservative as root starch reserves were depleted more rapidly in roots of girdled than in nongirdled trees. Thus, heterotrophic activity was overestimated.

Calculated on a unit area basis, both the heterotrophic and autotrophic soil respiration was significantly lower in fertilized plots, which is especially noteworthy given that aboveground production was around three times higher in fertilized than in nonfertilized plots.

Fertilization of boreal forest reduces both autotrophic and heterotrophic soil respiration

Low molecular weight organic acids and their Al-complexes in soil solution—composition, distribution and seasonal variation in three podzolized soils

Weatherable minerals in all podzol surface soils and shallow granitic rock under European coniferous forests studied hitherto are criss-crossed by numerous open, tubular pores, 3-10 μm in width. We ...

Reiner Giesler

Papers

Boreal forest plants take up organic nitrogen

Rock-eating fungi

Fertilization of boreal forest reduces both autotrophic and heterotrophic soil respiration

Low molecular weight organic acids and their Al-complexes in soil solution—composition, distribution and seasonal variation in three podzolized soils

Mycorrhizal weathering: A true case of mineral plant nutrition?