Terrestrial nitrogen cycle simulation with a dynamic global vegetation model

A global scale Dynamic Nitrogen scheme (DyN) has been developed and incorporated into the Lund–Posdam–Jena (LPJ) dynamic global vegetation model (DGVM). The DyN is a comprehensive process‐based model of the cycling of N through and within terrestrial ecosystems, with fully interactive coupling to vegetation and C dynamics. The model represents the uptake, allocation and turnover of N in plants, and soil N transformations including mineralization, N₂ fixation, nitrification and denitrification, NH₃ volatilization, N leaching, and N₂, N₂O and NO production and emission. Modelled global patterns of site‐scale nitrogen fluxes and reservoirs are highly correlated to observations reported from different biomes. The simulation of site‐scale net primary production and soil carbon content was improved relative to the original LPJ, which lacked an interactive N cycle, especially in the temporal and boreal regions. Annual N uptake by global natural vegetation was simulated as 1.084 Pg N yr⁻¹, with lowest values <1 g N m⁻² yr⁻¹ (polar desert) and highest values in the range 24–36.5 g N m⁻² yr⁻¹ (tropical forests). Simulated global patterns of annual N uptake are consistent with previous model results by Melillo et al. The model estimates global total nitrogen storage potentials in vegetation (5.3 Pg N), litter (4.6 Pg N) and soil (≥67 Pg as organic N and 0.94 Pg as inorganic N). Simulated global patterns of soil N storage are consistent with the analysis by Post et al. although total simulated N storage is less. Deserts were simulated to store 460 Tg N (up to 0.262 kg N m⁻²) as NO₃⁻, contributing 80% of the global total NO₃⁻ inventory of 580 Tg N. This model result is in agreement with the findings of a large NO₃⁻ pool beneath deserts. Globally, inorganic soil N is a small reservoir, comprising only 1.6% of the global soil N content to 1.5 m soil depth, but the ratio has a very high spatial variability and in hot desert regions, inorganic NO₃⁻ is estimated to be the dominant form of stored N in the soil.     

Mycorrhizal‐mediated nitrogen acquisition in switchgrass under elevated temperatures and N enrichment

Arbuscular mycorrhizal fungi (AMF) can perform key roles in ecosystem functioning through improving host nutrient acquisition. Nitrogen (N) is an essential nutrient for plant growth, however, anthropogenic N loading (e.g. crop fertilization and deposition from combustion sources) is increasing so that N now threatens ecosystem sustainability around the world by causing terrestrial and aquatic eutrophication and acidification. It is important to better understand the capacity of AMF to directly uptake N from soils and transfer it to host plants because this process may increase N recycling and retention within ecosystems. In addition to understanding the role of AMF in the N cycle in the present day it is important to understand how AMF function may change as global change proceeds. Currently the net effects of N enrichment and elevated temperature predicted with global change on AMF are unknown. In this study, we examined the effects of N enrichment by simulated N‐deposition loading, elevated temperatures expected by future global changes and their interactions on growth and AMF‐mediated N acquisition of switchgrass (Panicum virgatum var. Alamo), an important species for biofuel production. Switchgrass plants were grown in microcosm units that divided mycorrhizal roots from AMF hyphae and organic residues enriched with ¹⁵N by compartments separated by an air gap to reduce N diffusion. While AMF did not enhance switchgrass biomass, mycorrhizas significantly increased ¹⁵N in shoots and total shoot N. Neither N enrichment nor elevated temperatures influenced this mycorrhizal‐mediated N uptake and transfer. Results from this study can aid in developing sustainable bioethanol and switchgrass production practices that are less reliant on synthetic fertilizers and more dependent on internal N recycling from AMF.     

Mycorrhizal mediation of plant N acquisition and residue decomposition: Impact of mineral N inputs

Mycorrhizas are ubiquitous plant–fungus mutualists in terrestrial ecosystems and play important roles in plant resource capture and nutrient cycling. Sporadic evidence suggests that anthropogenic nitrogen (N) input may impact the development and the functioning of arbuscular mycorrhizal (AM) fungi, potentially altering host plant growth and soil carbon (C) dynamics. In this study, we examined how mineral N inputs affected mycorrhizal mediation of plant N acquisition and residue decomposition in a microcosm system. Each microcosm unit was separated into HOST and TEST compartments by a replaceable mesh screen that either prevented or allowed AM fungal hyphae but not plant roots to grow into the TEST compartments. Wild oat (Avena fatua L.) was planted in the HOST compartments that had been inoculated with either a single species of AM fungus, Glomus etunicatum, or a mixture of AM fungi including G. etunicatum. Mycorrhizal contributions to plant N acquisition and residue decomposition were directly assessed by introducing a mineral ¹⁵N tracer and ¹³C‐rich residues of a C₄ plant to the TEST compartments. Results from ¹⁵N tracer measurements showed that AM fungal hyphae directly transported N from the TEST soil to the host plant. Compared with the control with no penetration of AM fungal hyphae, AM hyphal penetration led to a 125% increase in biomass ¹⁵N of host plants and a 20% reduction in extractable inorganic N in the TEST soil. Mineral N inputs to the HOST compartments (equivalent to 5.0 g N m^?2 yr^?1) increased oat biomass and total root length colonized by mycorrhizal fungi by 189% and 285%, respectively, as compared with the no‐N control. Mineral N inputs to the HOST plants also reduced extractable inorganic N and particulate residue C proportion by 58% and 12%, respectively, in the corresponding TEST soils as compared to the no‐N control, by stimulating AM fungal growth and activities. The species mixture of mycorrhizal fungi was more effective in facilitating N transport and residue decomposition than the single AM species. These findings indicate that low‐level mineral N inputs may significantly enhance nutrient cycling and plant resource capture in terrestrial ecosystems via stimulation of root growth, mycorrhizal functioning, and residue decomposition. The long‐term effects of these observed alterations on soil C dynamics remain to be investigated.     

Soil warming opens the nitrogen cycle at the alpine treeline

Climate warming may alter ecosystem nitrogen (N) cycling by accelerating N transformations in the soil, and changes may be especially pronounced in cold regions characterized by N‐poor ecosystems. We investigated N dynamics across the plant–soil continuum during 6 years of experimental soil warming (2007–2012; +4 °C) at a Swiss high‐elevation treeline site (Stillberg, Davos; 2180 m a.s.l.) featuring Larix decidua and Pinus uncinata. In the soil, we observed considerable increases in the pool size in the first years of warming (by >50%), but this effect declined over time. In contrast, dissolved organic nitrogen (DON) concentrations in soil solutions from the organic layer increased under warming, especially in later years (maximum of +45% in 2012), suggesting enhanced DON leaching from the main rooting zone. Throughout the experimental period, foliar N concentrations showed species‐specific but small warming effects, whereas δ¹⁵N values showed a sustained increase in warmed plots that was consistent for all species analysed. The estimated total plant N pool size at the end of the study was greater (+17%) in warmed plots with Pinus but not in those containing Larix, with responses driven by trees. Irrespective of plot tree species identity, warming led to an enhanced N pool size of Vaccinium dwarf shrubs, no change in that of Empetrum hermaphroditum (dwarf shrub) and forbs, and a reduction in that of grasses, nonvascular plants, and fine roots. In combination, higher foliar δ¹⁵N values and the transient response in soil inorganic N indicate a persistent increase in plant‐available N and greater cumulative plant N uptake in warmer soils. Overall, greater N availability and increased DON concentrations suggest an opening of the N cycle with global warming, which might contribute to growth stimulation of some plant species while simultaneously leading to greater N losses from treeline ecosystems and possibly other cold biomes.     

Effect of plant nitrogen status on the contribution of arbuscular mycorrhizal hyphae to plant nitrogen uptake

The contribution of hyphae of Glomus mosseae (Nicol. and Gerd.) Gerd. and Trappe (BEG 107) to the acquisition of mineral nitrogen by Triticum aestivum L. cv. Hano (wheat) was tested under conditions of low P and high N (+N−P) or low N (−N−P). Mycorrhizal colonisation increased the shoot dry weight and plant tissue concentrations of P and cations. However, N tissue concentrations of mycorrhizal plants were not increased, although nitrate reductase activities were significantly higher (in vivo activity) in +N−P mycorrhizal compared to non-mycorrhizal roots. Severe plant N deficiency reduced the percentage root length colonised (but not the percentage viable colonisation), hyphal length, total 15 N uptake by hyphae and dry weight of mycorrhizal plants. Although mycorrhizal colonisation did not affect the overall plant N status, hyphae transported 1% (−N−P) and 7% (+N−P) of the 15 N-labelled NH₄NO₃ to mycorrhizal plants over 48 h. The higher rate of hyphal N uptake was apparently related to the more extensive hyphal growth at the higher level of plant N supply. However, the hyphal N supply was not sufficiently high to sustain adequate N nutrition of the plants supplied with very low amounts of N to the roots. Conversely, a sufficient N supply to the roots was important for the development of an extensive mycelium.     

High nitrogen deposition alters the decomposition of bog plant litter and reduces carbon accumulation

Bogs are globally important sinks of atmospheric carbon (C) due to the accumulation of partially decomposed litter that forms peat. Because bogs receive their nutrients from the atmosphere, the world‐wide increase of nitrogen (N) deposition is expected to affect litter decomposition and, ultimately, the rate of C accumulation. However, the mechanism of such biogeochemical alteration remains unclear and quantification of the effect of N addition on litter accumulation has yet to be done. Here, we show that 7 years of N addition to a bog decreased the C : N ratio, increased the bacterial biomass and stimulated the activity of hydrolytic and oxidative enzymes in surface peat. Furthermore, N addition modified nutrient limitation of microbes during litter decomposition so that phosphorus became a primary limiting nutrient. Alteration of N release from decomposing litter affected bog water chemistry and the competitive balance between peat‐forming mosses and vascular plants. We estimate that deposition of about 4 g N m^?2 yr^?1 will cause a mean annual reduction of fresh litter C accumulation of about 40 g m^?2 primarily as a consequence of decreased litter production from peat‐forming mosses. Our findings show that N deposition interacts with both above and below ground components of biodiversity to threaten the ability of bogs to act as N‐sinks, which may offset the positive effects of N on C accumulation seen in other ecosystems.     

Disentangling effects of air and soil temperature on C allocation in cold environments: A 14C pulse‐labelling study with two plant species

Carbon cycling responses of ecosystems to global warming will likely be stronger in cold ecosystems where many processes are temperature‐limited. Predicting these effects is difficult because air and soil temperatures will not change in concert, and will affect above and belowground processes differently. We disentangled above and belowground temperature effects on plant C allocation and deposition of plant C in soils by independently manipulating air and soil temperatures in microcosms planted with either Leucanthemopsis alpina or Pinus mugo seedlings. Daily average temperatures of 4 or 9°C were applied to shoots and independently to roots, and plants pulse‐labelled with ¹⁴CO₂. We traced soil CO₂ and ¹⁴CO₂ evolution for 4 days, after which microcosms were destructively harvested and ¹⁴C quantified in plant and soil fractions. In microcosms with L. alpina, net ¹⁴C uptake was higher at 9°C than at 4°C soil temperature, and this difference was independent of air temperature. In warmer soils, more C was allocated to roots at greater soil depth, with no effect of air temperature. In P. mugo microcosms, assimilate partitioning to roots increased with air temperature, but only when soils were at 9°C. Higher soil temperatures also increased the mean soil depth at which ¹⁴C was allocated. Our findings highlight the dependence of C uptake, use, and partitioning on both air and soil temperature, with the latter being relatively more important. The strong temperature‐sensitivity of C assimilate use in the roots and rhizosphere supports the hypothesis that cold limitation on C uptake is primarily mediated by reduced sink strength in the roots. We conclude that variations in soil rather than air temperature are going to drive plant responses to warming in cold environments, with potentially large changes in C cycling due to enhanced transfer of plant‐derived C to soils.     

Inclusion of ecologically based trait variation in plant functional types reduces the projected land carbon sink in an earth system model

Earth system models demonstrate large uncertainty in projected changes in terrestrial carbon budgets. The lack of inclusion of adaptive responses of vegetation communities to the environment has been suggested to hamper the ability of modeled vegetation to adequately respond to environmental change. In this study, variation in functional responses of vegetation has been added to an earth system model (ESM) based on ecological principles. The restriction of viable mean trait values of vegetation communities by the environment, called ‘habitat filtering’, is an important ecological assembly rule and allows for determination of global scale trait–environment relationships. These relationships were applied to model trait variation for different plant functional types (PFTs). For three leaf traits (specific leaf area, maximum carboxylation rate at 25 °C, and maximum electron transport rate at 25 °C), relationships with multiple environmental drivers, such as precipitation, temperature, radiation, and CO₂, were determined for the PFTs within the Max Planck Institute ESM. With these relationships, spatiotemporal variation in these formerly fixed traits in PFTs was modeled in global change projections (IPCC RCP8.5 scenario). Inclusion of this environment‐driven trait variation resulted in a strong reduction of the global carbon sink by at least 33% (2.1 Pg C yr^?1) from the 2nd quarter of the 21st century onward compared to the default model with fixed traits. In addition, the mid‐ and high latitudes became a stronger carbon sink and the tropics a stronger carbon source, caused by trait‐induced differences in productivity and relative respirational costs. These results point toward a reduction of the global carbon sink when including a more realistic representation of functional vegetation responses, implying more carbon will stay airborne, which could fuel further climate change.     

Is nitrogen transfer among plants enhanced by contrasting nutrient‐acquisition strategies?

Nitrogen (N) transfer among plants has been found where at least one plant can fix N₂. In nutrient‐poor soils, where plants with contrasting nutrient‐acquisition strategies (without N₂ fixation) co‐occur, it is unclear if N transfer exists and what promotes it. A novel multi‐species microcosm pot experiment was conducted to quantify N transfer between arbuscular mycorrhizal (AM), ectomycorrhizal (EM), dual AM/EM, and non‐mycorrhizal cluster‐rooted plants in nutrient‐poor soils with mycorrhizal mesh barriers. We foliar‐fed plants with a K¹⁵NO₃ solution to quantify one‐way N transfer from ‘donor’ to ‘receiver’ plants. We also quantified mycorrhizal colonization and root intermingling. Transfer of N between plants with contrasting nutrient‐acquisition strategies occurred at both low and high soil nutrient levels with or without root intermingling. The magnitude of N transfer was relatively high (representing 4% of donor plant N) given the lack of N₂ fixation. Receiver plants forming ectomycorrhizas or cluster roots were more enriched compared with AM‐only plants. We demonstrate N transfer between plants of contrasting nutrient‐acquisition strategies, and a preferential enrichment of cluster‐rooted and EM plants compared with AM plants. Nutrient exchanges among plants are potentially important in promoting plant coexistence in nutrient‐poor soils.     

Plant–fungus competition for nitrogen erases mycorrhizal growth benefits of Andropogon gerardii under limited nitrogen supply

Considered to play an important role in plant mineral nutrition, arbuscular mycorrhizal (AM) symbiosis is a common relationship between the roots of a great majority of plant species and glomeromycotan fungi. Its effects on the plant host are highly context dependent, with the greatest benefits often observed in phosphorus (P)‐limited environments. Mycorrhizal contribution to plant nitrogen (N) nutrition is probably less important under most conditions. Moreover, inasmuch as both plant and fungi require substantial quantities of N for their growth, competition for N could potentially reduce net mycorrhizal benefits to the plant under conditions of limited N supply. Further compounded by increased belowground carbon (C) drain, the mycorrhizal costs could outweigh the benefits under severe N limitation. Using a field AM fungal community or a laboratory culture of Rhizophagus irregularis as mycorrhizal inoculants, we tested the contribution of mycorrhizal symbiosis to the growth, C allocation, and mineral nutrition of Andropogon gerardii growing in a nutrient‐poor substrate under variable N and P supplies. The plants unambiguously competed with the fungi for N when its supply was low, resulting in no or negative mycorrhizal growth and N‐uptake responses under such conditions. The field AM fungal communities manifested their potential to improve plant P nutrition only upon N fertilization, whereas the R. irregularis slightly yet significantly increased P uptake of its plant host (but not the host's growth) even without N supply. Coincident with increasing levels of root colonization by the AM fungal structures, both inoculants invariably increased nutritional and growth benefits to the host with increasing N supply. This, in turn, resulted in relieving plant P deficiency, which was persistent in non‐mycorrhizal plants across the entire range of nutrient supplies.     

The European carbon balance. Part 4: integration of carbon and other trace‐gas fluxes

Overviewing the European carbon (C), greenhouse gas (GHG), and non‐GHG fluxes, gross primary productivity (GPP) is about 9.3 Pg yr^?1, and fossil fuel imports are 1.6 Pg yr^?1. GPP is about 1.25% of solar radiation, containing about 360 × 10¹⁸ J energy – five times the energy content of annual fossil fuel use. Net primary production (NPP) is 50%, terrestrial net biome productivity, NBP, 3%, and the net GHG balance, NGB, 0.3% of GPP. Human harvest uses 20% of NPP or 10% of GPP, or alternatively 1‰ of solar radiation after accounting for the inherent cost of agriculture and forestry, for production of pesticides and fertilizer, the return of organic fertilizer, and for the C equivalent cost of GHG emissions. C equivalents are defined on a global warming potential with a 100‐year time horizon. The equivalent of about 2.4% of the mineral fertilizer input is emitted as N₂O. Agricultural emissions to the atmosphere are about 40% of total methane, 60% of total NO‐N, 70% of total N₂O‐N, and 95% of total NH₃‐N emissions of Europe. European soils are a net C sink (114 Tg yr^?1), but considering the emissions of GHGs, soils are a source of about 26 Tg CO₂ C‐equivalent yr^?1. Forest, grassland and sediment C sinks are offset by GHG emissions from croplands, peatlands and inland waters. Non‐GHGs (NH₃, NOx) interact significantly with the GHG and the C cycle through ammonium nitrate aerosols and dry deposition. Wet deposition of nitrogen (N) supports about 50% of forest timber growth. Land use change is regionally important. The absolute flux values total about 50 Tg C yr^?1. Nevertheless, for the European trace‐gas balance, land‐use intensity is more important than land‐use change. This study shows that emissions of GHGs and non‐GHGs significantly distort the C cycle and eliminate apparent C sinks.     

Feedbacks between plant N demand and rhizosphere priming depend on type of mycorrhizal association

Ecosystem carbon (C) balance is hypothesised to be sensitive to the mycorrhizal strategies that plants use to acquire nutrients. To test this idea, we coupled an optimality‐based plant nitrogen (N) acquisition model with a microbe‐focused soil organic matter (SOM) model. The model accurately predicted rhizosphere processes and C–N dynamics across a gradient of stands varying in their relative abundance of arbuscular mycorrhizal (AM) and ectomycorrhizal (ECM) trees. When mycorrhizal dominance was switched – ECM trees dominating plots previously occupied by AM trees, and vice versa – legacy effects were apparent, with consequences for both C and N stocks in soil. Under elevated productivity, ECM trees enhanced decomposition more than AM trees via microbial priming of unprotected SOM. Collectively, our results show that ecosystem responses to global change may hinge on the balance between rhizosphere priming and SOM protection, and highlight the importance of dynamically linking plants and microbes in terrestrial biosphere models.     

Robust biological nitrogen fixation in a model grass–bacterial association

Nitrogen‐fixing rhizobacteria can promote plant growth; however, it is controversial whether biological nitrogen fixation (BNF) from associative interaction contributes to growth promotion. The roots of Setaria viridis, a model C₄ grass, were effectively colonized by bacterial inoculants resulting in a significant enhancement of growth. Nitrogen‐13 tracer studies provided direct evidence for tracer uptake by the host plant and incorporation into protein. Indeed, plants showed robust growth under nitrogen‐limiting conditions when inoculated with an ammonium‐excreting strain of Azospirillum brasilense. ¹¹C‐labeling experiments showed that patterns in central carbon metabolism and resource allocation exhibited by nitrogen‐starved plants were largely reversed by bacterial inoculation, such that they resembled plants grown under nitrogen‐sufficient conditions. Adoption of S. viridis as a model should promote research into the mechanisms of associative nitrogen fixation with the ultimate goal of greater adoption of BNF for sustainable crop production.     

Ectomycorrhizal impacts on plant nitrogen nutrition: emerging isotopic patterns,latitudinal variation and hidden mechanisms

Ectomycorrhizal (EcM)‐mediated nitrogen (N) acquisition is one main strategy used by terrestrial plants to facilitate growth. Measurements of natural abundance nitrogen isotope ratios (denoted as δ¹⁵N relative to a standard) increasingly serve as integrative proxies for mycorrhiza‐mediated N acquisition due to biological fractionation processes that alter ¹⁵N:¹⁴N ratios. Current understanding of these processes is based on studies from high‐latitude ecosystems where plant productivity is largely limited by N availability. Much less is known about the cause and utility of ecosystem δ¹⁵N patterns in the tropics. Using structural equation models, model selection and isotope mass balance we assessed relationships among co‐occurring soil, mycorrhizal plants and fungal N pools measured from 40 high‐ and 9 low‐latitude ecosystems. At low latitudes ¹⁵N‐enrichment caused ecosystem components to significantly deviate from those in higher latitudes. Collectively, δ¹⁵N patterns suggested reduced N‐dependency and unique sources of EcM ¹⁵N‐enrichment under conditions of high N availability typical of the tropics. Understanding the role of mycorrhizae in global N cycles will require reevaluation of high‐latitude perspectives on fractionation sources that structure ecosystem δ¹⁵N patterns, as well as better integration of EcM function with biogeochemical theories pertaining to climate‐nutrient cycling relationships.     

Arbuscular mycorrhizas and their role in plant growth, nitrogen interception and soil gas efflux in an organic production system

Background and aims

Roots and mycorrhizas play an important role in not only plant nutrient acquisition, but also ecosystem nutrient cycling.

Methods

A field experiment was undertaken in which the role of arbuscular mycorrhizas (AM) in the growth and nutrient acquisition of tomato plants was investigated. A mycorrhiza defective mutant of tomato (Solanum lycopersicum L.) (named rmc) and its mycorrhizal wild type progenitor (named 76R) were used to control for the formation of AM. The role of roots and AM in soil N cycling was studied by injecting a ¹⁵N-labelled nitrate solution into surface soil at different distances from the 76R and rmc genotypes of tomato, or in plant free soil. The impacts of mycorrhizal and non-mycorrhizal root systems on soil greenhouse gas (CO₂ and ¹⁴⁺¹⁵N₂O and ¹⁵N₂O) emissions, relative to root free soils, were also studied.

Results

The formation of AM significantly enhanced plant growth and nutrient acquisition, including interception of recently applied NO ₃ ^? . Whereas roots caused a small but significant decrease in ¹⁵N₂O emissions from soils at 23?h after labeling, compared to the root-free treatment, arbuscular mycorrhizal fungi (AMF) had little effect on N₂O emissions. In contrast soil CO₂ emissions were higher in plots containing mycorrhizal root systems, where root biomass was also greater.

Conclusions

Taken together, these data indicate that roots and AMF have an important role to play in plant nutrient acquisition and ecosystem N cycling.     

丛枝菌根真菌与氮添加对不同根形态基因型水稻氮吸收的影响

植物主要依赖自身根系从土壤中获取矿质养分; 具有不同根形态的植物对于养分的吸收能力存在差异。丛枝菌根真菌(AMF)能与陆地植物根系形成共生关系, 帮助植物吸收矿质养分。但是, AMF对于植物根系养分吸收的促进效应是否会受根形态的影响还鲜有研究。该研究选取4种不同根形态基因型水稻(根毛缺陷突变体rhl1、侧根缺陷突变体iaa11、不定根缺失突变体arl1和野生型Kas)为研究对象, 设置2种施氮水平处理(低氮: 20 mg·kg^-1氨氮; 高氮: 100 mg·kg^-1氨氮), 利用稳定同位素¹⁵N示踪标记技术, 探究AMF和氮添加对不同根形态植物氮吸收的影响。研究结果发现, 相比低氮处理, 高氮处理下, rhl1、Kas、iaa11与arl1的茎叶¹⁵N浓度分别提高了60%、72%、128%与118%, 说明氮添加显著促进了水稻氮吸收, 且iaa11与arl1对氮添加的响应更强烈。在低氮水平下, AMF对rhl1、Kas、iaa11与arl1氮吸收的平均效应值分别为17%、31%、42%、51%, 表明AMF对于植物氮吸收的促进效应受根形态影响, iaa11与arl1对AMF的响应明显高于Kas与rhl1; 相较于低氮水平, 高氮水平下AMF对于不同根形态水稻氮吸收的促进效应都会显著降低, 表明氮添加削弱了AMF对植物氮吸收的促进效应。该研究阐明了4种不同根形态基因型水稻氮养分吸收存在显著差异, 其中氮吸收能力较弱的基因型水稻对AMF的响应更强, 该结果补充了植物与AMF在养分吸收上存在功能互补的控制实验证据。     

C allocation to the fungus is not a cost to the plant in ectomycorrhizae

Mycorrhizal benefit to plants is most frequently evaluated through growth differences between mycorrhizal (M) and non‐mycorrhizal (NM) plants. These growth differences are often considered to be due to differences in belowground C expenditure, or in cost efficiency, i.e. amount of nutrients acquired per C expended. We searched published reports for relations between plant growth and belowground C allocation, C use efficiency, or nutrient uptake, in ectomycorrhizal (ECM) versus non‐mycorrhizal plants. We found a similar number of cases of negative, null or positive effects of ECM on plant growth. These effects were not correlated with differences on belowground C allocation or C use efficiency between M and NM plants. In contrast, they were very strongly correlated with mycorrhizal effects on plant N gain. A comprehensive analysis of the published data therefore provided evidence that C is an excess, rather than a costly, resource, and that the outcome of the symbiosis depends only on whether mycorrhizae result in increased or decreased nutrient acquisition compared with NM plants, and not on cost efficiency differences between M and NM plants. Consequences of this finding for the regulation of resource exchange between symbionts and the nature of the symbiosis are discussed.     

Organic enrichment of sediments reduces arbuscular mycorrhizal fungi in oligotrophic lake plants

1. Arbuscular mycorrhizal fungi (AMF) commonly colonise isoetid species inhabiting oxygenated sediments in oligotrophic lakes but are usually absent in other submerged plants. We hypothesised that organic enrichment of oligotrophic lake sediments reduces AMF colonisation and hyphal growth because of sediment O₂ depletion and low carbon supply from stressed host plants. 2. We added organic matter to sediments inhabited by isoetids and measured pore‐water chemistry (dissolved O₂, inorganic carbon, Fe²⁺ and ), colonisation intensity of roots and hyphal density after 135 days of exposure. 3. Addition of organic matter reduced AMF colonisation of roots of both Lobelia dortmanna and Littorella uniflora, and high additions stressed the plants. Even small additions of organic matter almost stopped AMF colonisation of initially un‐colonised L. uniflora, though without reducing plant growth. Mean hyphal density in sediments was high (6 and 15 m cm^?3) and comparable with that in terrestrial soils (2–40 m cm^?3). Hyphal density was low in the upper 1 cm of isoetid sediments, high in the main root zone between 1 and 8 cm and positively related to root density. Hyphal surface area exceeded root surface area by 1.7–3.2 times. 4. We conclude that AMF efficiently colonise isoetids in oligotrophic sediments and form extensive hyphal networks. Small additions of organic matter to sediments induce sediment anoxia and reduce AMF colonisation of roots but cause no apparent plant stress. High organic addition induces night‐time anoxia in both the sediment and the plant tissue. Tissue anoxia reduces root growth and AMF colonisation, probably because of restricted translocation of nutrient ions and organic solutes between roots and leaves. Isoetids should rely on AMF for P uptake on nutrient‐poor mineral sediments but are capable of growing without AMF on organic sediments.     

Global patterns of nitrogen limitation: confronting two global biogeochemical models with observations

Projections of future changes in land carbon (C) storage using biogeochemical models depend on accurately modeling the interactions between the C and nitrogen (N) cycles. Here, we present a framework for analyzing N limitation in global biogeochemical models to explore how C‐N interactions of current models compare to field observations, identify the processes causing model divergence, and identify future observation and experiment needs. We used a set of N‐fertilization simulations from two global biogeochemical models (CLM‐CN and O‐CN) that use different approaches to modeling C‐N interactions. On the global scale, net primary productivity (NPP) in the CLM‐CN model was substantially more responsive to N fertilization than in the O‐CN model. The most striking difference between the two models occurred for humid tropical forests, where the CLM‐CN simulated a 62% increase in NPP at high N addition levels (30 g N m^?2 yr^?1), while the O‐CN predicted a 2% decrease in NPP due to N fertilization increasing plant respiration more than photosynthesis. Across 35 temperate and boreal forest sites with field N‐fertilization experiments, we show that the CLM‐CN simulated a 46% increase in aboveground NPP in response to N, which exceeded the observed increase of 25%. In contrast, the O‐CN only simulated a 6% increase in aboveground NPP at the N‐fertilization sites. Despite the small response of NPP to N fertilization, the O‐CN model accurately simulated ecosystem retention of N and the fate of added N to vegetation when compared to empirical ¹⁵N tracer application studies. In contrast, the CLM‐CN predicted lower total ecosystem N retention and partitioned more losses to volatilization than estimated from observed N budgets of small catchments. These results point to the need for model improvements in both models in order to enhance the accuracy with which global C‐N cycle feedbacks are simulated.     

Crassulacean acid metabolism (CAM) offers sustainable bioenergy production and resilience to climate change

Biomass production on low‐grade land is needed to meet future energy demands and minimize resource conflicts. This, however, requires improvements in plant water‐use efficiency (WUE) that are beyond conventional C3 and C4 dedicated bioenergy crops. Here we present the first global‐scale geographic information system (GIS)‐based productivity model of two highly water‐efficient crassulacean acid metabolism (CAM) candidates: Agave tequilana and Opuntia ficus‐indica. Features of these plants that translate to WUE advantages over C3 and C4 bioenergy crops include nocturnal stomatal opening, rapid rectifier‐like root hydraulic conductivity responses to fluctuating soil water potential and the capacity to buffer against periods of drought. Yield simulations for the year 2070 were performed under the four representative concentration pathway (RCPs) scenarios presented in the IPCC's 5th Assessment Report. Simulations on low‐grade land suggest that O. ficus‐indica alone has the capacity to meet ‘extreme’ bioenergy demand scenarios (>600 EJ yr^?1) and is highly resilient to climate change (?1%). Agave tequilana is moderately impacted (?11%). These results are significant because bioenergy demand scenarios >600 EJ yr^?1 could be met without significantly increasing conflicts with food production and contributing to deforestation. Both CAM candidates outperformed the C4 bioenergy crop, Panicum virgatum L. (switchgrass) in arid zones in the latitudinal range 30°S–30°N.