Resilience of ecosystem service delivery in grasslands in response to single and compound extreme weather events.

Extreme weather events are increasing in frequency and magnitude with profound effects on ecosystem functioning. Further, there is now a greater likelihood that multiple extreme events are occurring within a single year. Here we investigated the effect of a single drought, flood or compound (flood + drought) extreme event on temperate grassland ecosystem processes in a field experiment. To assess system resistance and resilience, we studied changes in a wide range of above- and below-ground indicators (plant diversity and productivity, greenhouse gas emissions, soil chemical, physical and biological metrics) during the 8 week stress events and then for 2 years post-stress. We hypothesized that agricultural grasslands would have different degrees of resistance and resilience to flood and drought stress. We also investigated two alternative hypotheses that the combined flood + drought treatment would either, (A) promote ecosystem resilience through more rapid recovery of soil moisture conditions or (B) exacerbate the impact of the single flood or drought event. Our results showed that flooding had a much greater effect than drought on ecosystem processes and that the grassland was more resistant and resilient to drought than to flood. The immediate impact of flooding on all indicators was negative, especially for those related to production, and climate and water regulation. Flooding stress caused pronounced and persistent shifts in soil microbial and plant communities with large implications for nutrient cycling and long-term ecosystem function. The compound flood + drought treatment failed to show a more severe impact than the single extreme events. Rather, there was an indication of quicker recovery of soil and microbial parameters suggesting greater resilience in line with hypothesis (A). This study clearly reveals that contrasting extreme weather events differentially affect grassland ecosystem function but that concurrent events of a contrasting nature may promote ecosystem resilience to future stress.

Iron-Modified Biochar Strengthens Simazine Adsorption and Decreases Simazine Decomposition in the Soil.

Currently, modified biochar has been successfully used in the remediation of soil polluted with heavy metals. However, the effects of the modified biochar on pesticides (such as simazine) are still unclear. Herein, the environmental fate of simazine, such as decomposition, leaching, and adsorption in unamended soil, in the soil amended with unmodified and modified biochar (biochar + FeCl3, biochar + FeOS, biochar + Fe) were evaluated. In addition, an incubation experiment was also performed to observe the influence of modified biochar on the microbial community and diversity in the soil. The results showed that modified biochar significantly decreased the decomposition of simazine in the soil compared to its counterpart. Modified biochar also reduced the concentration of simazine in the leachate. Compared with the control, soil microbial biomass in the soil amended with unmodified biochar, biochar + FeCl3, biochar + Fe, and biochar + FeOS was decreased by 5.3%, 18.8%, 8.7%, and 18.1%, respectively. Furthermore, modified biochar changed the structure of the microbial community. This shows that modified biochar could increase the soil adsorption capacity for simazine and change the amount and microbial community that regulates the fate of simazine in the soil. This study concludes that iron-modified biochar has positive and negative effects on the soil. Therefore, its advantages and side effects should be considered before applying it to the soil.

Identification and predictability of soil quality indicators from conventional soil and vegetation classifications.

The physical, chemical and biological attributes of a soil combined with abiotic factors (e.g. climate and topography) drive pedogenesis and some of these attributes have been used as proxies to soil quality. Thus, we investigated: (1) whether appropriate soil quality indicators (SQIs) could be identified in soils of Great Britain, (2) whether conventional soil classification or aggregate vegetation classes (AVCs) could predict SQIs and (3) to what extent do soil types and/ or AVCs act as major regulators of SQIs. Factor analysis was used to group 20 soil attributes into six SQI which were named as; soil organic matter (SOM), dissolved organic matter (DOM), soluble N, reduced N, microbial biomass, DOM humification (DOMH). SOM was identified as the most important SQI in the discrimination of both soil types and AVCs. Soil attributes constituting highly to the SOM factor were, microbial quotient and bulk density. The SOM indicator discriminated three soil type groupings and four aggregate vegetation class groupings. Among the soil types, only the peat soils were discriminated from other groups while among the AVCs only the heath and bog classes were isolated from others. However, the peat soil and heath and bog AVC were the only groups that were distinctly discriminated from other groups. All other groups heavily overlapped with one another, making it practically impossible to define reference values for each soil type or AVC. The two-way ANOVA showed that the AVCs were a better regulator of the SQIs than the soil types. We conclude that conventionally classified soil types cannot predict the SQIs defined from large areas with differing climatic and edaphic factors. Localised areas with similar climatic and topoedaphic factors may hold promise for the definition of SQI that may predict the soil types or AVCs.

Substrate control of sulphur utilisation and microbial stoichiometry in soil: Results of 13C, 15N, 14C, and 35S quad labelling.

Global plant sulphur (S) deficiency is increasing because of a reduction in sulphate-based fertiliser application combined with continuous S withdrawal during harvest. Here, we applied 13C, 15N, 14C, and 35S quad labelling of the S-containing amino acids cysteine (Cys) and methionine (Met) to understand S cycling and microbial S transformations in the soil. The soil microorganisms absorbed the applied Cys and Met within minutes and released SO42- within hours. The SO42- was reutilised by the MB within days. The initial microbial utilisation and SO42- release were determined by amino acid structure. Met released 2.5-fold less SO42- than Cys. The microbial biomass retained comparatively more C and S from Met than Cys. The microorganisms decomposed Cys to pyruvate and H2S whereas they converted Met to α-ketobutyrate and S-CH3. The microbial stoichiometries of C, N, and S derived from Cys and Met were balanced after 4 d by Cys-derived SO42- uptake and Met-derived CO2 release. The microbial C:N:S ratio dynamics showed rapid C utilisation and loss, stable N levels, and S accumulation. Thus, short-term organic S utilisation by soil microorganisms is determined by amino acid structure whilst long-term organic S utilisation by soil microorganisms is determined by microbially controlled stoichiometry.

Do plants use root-derived proteases to promote the uptake of soil organic nitrogen?

AIMS: The capacity of plant roots to directly acquire organic nitrogen (N) in the form of oligopeptides and amino acids from soil is well established. However, plants have poor access to protein, the central reservoir of soil organic N. Our question is: do plants actively secrete proteases to enhance the breakdown of soil protein or are they functionally reliant on soil microorganisms to undertake this role? METHODS: Growing maize and wheat under sterile hydroponic conditions with and without inorganic N, we measured protease activity on the root surface (root-bound proteases) or exogenously in the solution (free proteases). We compared root protease activities to the rhizosphere microbial community to estimate the ecological significance of root-derived proteases. RESULTS: We found little evidence for the secretion of free proteases, with almost all protease activity associated with the root surface. Root protease activity was not stimulated under N deficiency. Our findings suggest that cereal roots contribute one-fifth of rhizosphere protease activity. CONCLUSIONS: Our results indicate that plant N uptake is only functionally significant when soil protein is in direct contact with root surfaces. The lack of protease upregulation under N deficiency suggests that root protease activity is unrelated to enhanced soil N capture.

Angiosperm symbioses with non-mycorrhizal fungal partners enhance N acquisition from ancient organic matter in a warming maritime Antarctic.

In contrast to the situation in plants inhabiting most of the world's ecosystems, mycorrhizal fungi are usually absent from roots of the only two native vascular plant species of maritime Antarctica, Deschampsia antarctica and Colobanthus quitensis. Instead, a range of ascomycete fungi, termed dark septate endophytes (DSEs), frequently colonise the roots of these plant species. We demonstrate that colonisation of Antarctic vascular plants by DSEs facilitates not only the acquisition of organic nitrogen as early protein breakdown products, but also as non-proteinaceous d-amino acids and their short peptides, accumulated in slowly-decomposing organic matter, such as moss peat. Our findings suggest that, in a warming maritime Antarctic, this symbiosis has a key role in accelerating the replacement of formerly dominant moss communities by vascular plants, and in increasing the rate at which ancient carbon stores laid down as moss peat over centuries or millennia are returned to the atmosphere as CO2 .

Plant-microbe competition: does injection of isotopes of C and N into the rhizosphere effectively characterise plant use of soil N?

Despite considerable attention over the last 25 yr, the importance of early protein breakdown products to plant nitrogen (N) nutrition remains uncertain. We used rhizosphere injection of 15 N-, 13 C- and 14 C-labelled inorganic N and amino acid (l-alanine), with chase periods from 1 min to 24 h, to investigate the duration of competition for amino acid between roots (Triticum aestivum) and soil microorganisms. We further investigated how microbial modification of l-alanine influenced plant carbon (C) and N recovery. From recovery of C isotopes, intact alanine uptake was 0.2-1.3% of added. Soil microbes appeared to remove alanine from soil solution within 1 min and release enough NH4+ to account for all plant 15 N recovery (over 24 h) within 5 min. Microbially generated inorganic or keto acid C accounted for < 25% of the lowest estimate of intact alanine uptake. Co-location of C and N labels appears a reasonable measure of intact uptake. Potential interference from microbially modified C is probably modest, but may increase with chase period. Similarly, competition for l-alanine is complete within a few minutes in soil, whereas NO3- added at the same rate is available for > 24 h, indicating that long chase periods bias outcomes and fail to accurately simulate soil processes.

Methodological bias associated with soluble protein recovery from soil.

Proteins play a crucial role in many soil processes, however, standardised methods to extract soluble protein from soil are lacking. The aim of this study was to compare the ability of different extractants to quantify the recovery of soluble proteins from three soil types (Cambisol, Ferralsol and Histosol) with contrasting clay and organic matter contents. Known amounts of plant-derived 14C-labelled soluble proteins were incubated with soil and then extracted with solutions of contrasting pH, concentration and polarity. Protein recovery proved highly solvent and soil dependent (Histosol > Cambisol > Ferralsol) and no single extractant was capable of complete protein recovery. In comparison to deionised water (10-60% of the total protein recovered), maximal recovery was observed with NaOH (0.1 M; 61-80%) and Na-pyrophosphate (0.05 M, pH 7.0; 45-75% recovery). We conclude that the dependence of protein recovery on both extractant and soil type prevents direct comparison of studies using different recovery methods, particularly if no extraction controls are used. We present recommendations for a standard protein extraction protocol.

Stoichiometric constraints on the microbial processing of carbon with soil depth along a riparian hillslope.

Soil organic matter (SOM) content is a key indicator of riparian soil functioning and in the provision of ecosystem services such as water retention, flood alleviation, pollutant attenuation and carbon (C) sequestration for climate change mitigation. Here, we studied the importance of microbial biomass and nutrient availability in regulating SOM turnover rates. C stabilisation in soil is expected to vary both vertically, down the soil profile and laterally across the riparian zone. In this study, we evaluated the influence of five factors on C mineralisation (Cmin): (i) substrate quantity, (ii) substrate quality, (iii) nutrient (C, N and P) stoichiometry, (iv) soil microbial activity with proximity to the river (2 to 75 m) and (v) as a function of soil depth (0-3 m). Substrate quality, quantity and nutrient stoichiometry were evaluated using high and low molecular weight 14C-labelled dissolved organic (DOC) along with different nutrient additions. Differences in soil microbial activity with proximity to the river and soil depth were assessed by comparing initial (immediate) Cmin rates and cumulative C mineralised at the end of the incubation period. Overall, microbial biomass C (MBC), organic matter (OM) and soil moisture content (MC) proved to be the major factors controlling rates of Cmin at depth. Differences in the immediate and medium-term response (42 days) of Cmin suggested that microbial growth increased and carbon use efficiency (CUE) decreased down the soil profile. Inorganic N and/or P availability had little or no effect on Cmin suggesting that microbial community growth and activity is predominantly C limited. Similarly, proximity to the watercourse also had relatively little effect on Cmin. This work challenges current theories suggesting that areas adjacent to watercourse process C differently from upslope areas. In contrast, our results suggest that substrate quality and microbial biomass are more important in regulating C processing rates rather than proximity to a river.

Crop residues exacerbate the negative effects of extreme flooding on soil quality.

Extreme flood events are predicted to have a negative impact on soil quality. Currently, there is a lack of information about the effect of agricultural practices on soil functioning and microbial processes under these events. We hypothesized that the impact of flooding on soil quality will be exacerbated when crop residues are present in the soil as they will induce more extreme anaerobicity. A spring extreme flood event (10 °C, 9 weeks) was simulated in mesocosms containing an arable sandy-loam soil low in nutrients. The main treatments were (1) with and without flooding and (2) with and without maize residue addition (8 Mg ha-1). We monitored changes in soil chemical quality indicators (e.g. pH, salinity, Fe3+, P, C, NH4 +, NO3 - and organic N), greenhouse gas (GHG) emissions (CO2, CH4, N2O) and soil microbial community composition (PLFAs) during a prolonged flood period (9 weeks) and an 8-week "recovery" period after flooding. In comparison to the other treatments, flooding in the presence of crop residues resulted in a dramatic drop in soil redox potential. This was associated with the enhanced release of Fe and C into solution and an increase in CH4 emissions. In contrast, maize residues reduced potential nitrate losses and N2O emissions, possibly due to complete denitrification and microbial N immobilization. Both flooding and maize residues stimulated microbial growth and promoted a shift in microbial community composition. Following floodwater removal, most of the soil quality indicators returned to the levels of the control treatment within 5 weeks. After this short recovery phase, no major impact of flooding could be observed on plant growth (maize pot-grown). Overall, we conclude that both extreme flooding and management regime negatively impact upon a range of soil quality indicators (e.g. redox, GHG emissions); however, the soil showed high resilience and recovered quickly after floodwater removal. Further work is required to investigate the impact of repeated extreme flood events on soil quality and function over longer timescales.

Rewriting yeast central carbon metabolism for industrial isoprenoid production.

A bio-based economy has the potential to provide sustainable substitutes for petroleum-based products and new chemical building blocks for advanced materials. We previously engineered Saccharomyces cerevisiae for industrial production of the isoprenoid artemisinic acid for use in antimalarial treatments. Adapting these strains for biosynthesis of other isoprenoids such as ß-farnesene (C15H24), a plant sesquiterpene with versatile industrial applications, is straightforward. However, S. cerevisiae uses a chemically inefficient pathway for isoprenoid biosynthesis, resulting in yield and productivity limitations incompatible with commodity-scale production. Here we use four non-native metabolic reactions to rewire central carbon metabolism in S. cerevisiae, enabling biosynthesis of cytosolic acetyl coenzyme A (acetyl-CoA, the two-carbon isoprenoid precursor) with a reduced ATP requirement, reduced loss of carbon to CO2-emitting reactions, and improved pathway redox balance. We show that strains with rewired central metabolism can devote an identical quantity of sugar to farnesene production as control strains, yet produce 25% more farnesene with that sugar while requiring 75% less oxygen. These changes lower feedstock costs and dramatically increase productivity in industrial fermentations which are by necessity oxygen-constrained. Despite altering key regulatory nodes, engineered strains grow robustly under taxing industrial conditions, maintaining stable yield for two weeks in broth that reaches >15% farnesene by volume. This illustrates that rewiring yeast central metabolism is a viable strategy for cost-effective, large-scale production of acetyl-CoA-derived molecules.

Challenging the paradigm of nitrogen cycling: no evidence of in situ resource partitioning by coexisting plant species in grasslands of contrasting fertility.

In monoculture, certain plant species are able to preferentially utilize different nitrogen (N) forms, both inorganic and organic, including amino acids and peptides, thus forming fundamental niches based on the chemical form of N. Results from field studies, however, are inconsistent: Some showing that coexisting plant species predominantly utilize inorganic N, while others reveal distinct interspecies preferences for different N forms. As a result, the extent to which hypothetical niches are realized in nature remains unclear. Here, we used in situ stable isotope tracer techniques to test the idea, in temperate grassland, that niche partitioning of N based on chemical form is related to plant productivity and the relative availability of organic and inorganic N. We also tested in situ whether grassland plants vary in their ability to compete for, and utilize peptides, which have recently been shown to act as an N source for plants in strongly N-limited ecosystems. We hypothesized that plants would preferentially use NO3 (-)-N and NH4 (+)-N over dissolved organic N in high-productivity grassland where inorganic N availability is high. On the other hand, in low-productivity grasslands, where the availability of dissolved inorganic N is low, and soil availability of dissolved organic N is greater, we predicted that plants would preferentially use N from amino acids and peptides, prior to microbial mineralization. Turves from two well-characterized grasslands of contrasting productivity and soil N availability were injected, in situ, with mixtures of (15)N-labeled inorganic N (NO3 (-) and NH4 (+)) and (13)C(15)N labeled amino acid (l-alanine) and peptide (l-tri-alanine). In order to measure rapid assimilation of these N forms by soil microbes and plants, the uptake of these substrates was traced within 2.5 hours into the shoots of the most abundant plant species, as well as roots and the soil microbial biomass. We found that, contrary to our hypothesis, the majority of plant species across both grasslands took up most N in the form of NH4 (+), suggesting that inorganic N is their predominant N source. However, we did find that organic N was a source of N which could be utilized by plant species at both sites, and in the low-productivity grassland, plants were able to capture some tri-alanine-N directly. Although our findings did not support the hypothesis that differences in the availability of inorganic and organic N facilitate resource partitioning in grassland, they do support the emerging view that peptides represent a significant, but until now neglected, component of the terrestrial N cycle.

Living roots magnify the response of soil organic carbon decomposition to temperature in temperate grassland.

Increasing atmospheric carbon dioxide (CO2 ) concentration is both a strong driver of primary productivity and widely believed to be the principal cause of recent increases in global temperature. Soils are the largest store of the world's terrestrial C. Consequently, many investigations have attempted to mechanistically understand how microbial mineralisation of soil organic carbon (SOC) to CO2 will be affected by projected increases in temperature. Most have attempted this in the absence of plants as the flux of CO2 from root and rhizomicrobial respiration in intact plant-soil systems confounds interpretation of measurements. We compared the effect of a small increase in temperature on respiration from soils without recent plant C with the effect on intact grass swards. We found that for 48 weeks, before acclimation occurred, an experimental 3 °C increase in sward temperature gave rise to a 50% increase in below ground respiration (ca. 0.4 kg C m(-2) ; Q10  = 3.5), whereas mineralisation of older SOC without plants increased with a Q10 of only 1.7 when subject to increases in ambient soil temperature. Subsequent (14) C dating of respired CO2 indicated that the presence of plants in swards more than doubled the effect of warming on the rate of mineralisation of SOC with an estimated mean C age of ca. 8 years or older relative to incubated soils without recent plant inputs. These results not only illustrate the formidable complexity of mechanisms controlling C fluxes in soils but also suggest that the dual biological and physical effects of CO2 on primary productivity and global temperature have the potential to synergistically increase the mineralisation of existing soil C.

Evaluation of dissolved organic carbon as a soil quality indicator in national monitoring schemes.

BACKGROUND: Monitoring the properties of dissolved organic carbon (DOC) in soil water is frequently used to evaluate changes in soil quality and to explain shifts in freshwater ecosystem functioning. METHODS: Using >700 individual soils (0-15 cm) collected from a 209,331 km(2) area we evaluated the relationship between soil classification (7 major soil types) or vegetation cover (8 dominant classes, e.g. cropland, grassland, forest) and the absorbance properties (254 and 400 nm), DOC quantity and quality (SUVA, total soluble phenolics) of soil water. RESULTS: Overall, a good correlation (r(2)= 0.58) was apparent between soil water absorbance and DOC concentration across the diverse range of soil types tested. In contrast, both DOC and the absorbance properties of soil water provided a poor predictor of SUVA or soluble phenolics which we used as a measure of humic substance concentration. Significant overlap in the measured ranges for UV absorbance, DOC, phenolic content and especially SUVA of soil water were apparent between the 8 vegetation and 7 soil classes. A number of significant differences, however, were apparent within these populations with total soluble phenolics giving the greatest statistical separation between both soil and vegetation groups. CONCLUSIONS: We conclude that the quality of DOC rather than its quantity provides a more useful measure of soil quality in large scale surveys.

How significant to plant N nutrition is the direct consumption of soil microbes by roots?

The high degree to which plant roots compete with soil microbes for organic forms of nitrogen (N) is becoming increasingly apparent. This has culminated in the finding that plants may consume soil microbes as a source of N, but the functional significance of this process remains unknown. We used (15) N- and (14) C-labelled cultures of soil bacteria to measure rates of acquisition of microbes by sterile wheat roots and plants growing in soil. We compared these rates with acquisition of (15) N delivered as nitrate, amino acid monomer (l-alanine) and short peptide (l-tetraalanine), and the rate of decomposition of [(14) C] microbes by indigenous soil microbiota. Acquisition of microbe (15) N by both sterile roots and roots growing in soil was one to two orders of magnitude slower than acquisition of all other forms of (15) N. Decomposition of microbes was fast enough to account for all (15) N recovered, but approximately equal recovery of microbe (14) C suggests that microbes entered roots intact. Uptake of soil microbes by wheat (Triticum aestivum) roots appears to take place in soil. If wheat is typical, the importance of this process to terrestrial N cycling is probably minor in comparison with fluxes of other forms of soil inorganic and organic N.

Acquisition and assimilation of nitrogen as peptide-bound and D-enantiomers of amino acids by wheat.

Nitrogen is a key regulator of primary productivity in many terrestrial ecosystems. Historically, only inorganic N (NH(4)(+) and NO(3)(-)) and L-amino acids have been considered to be important to the N nutrition of terrestrial plants. However, amino acids are also present in soil as small peptides and in D-enantiomeric form. We compared the uptake and assimilation of N as free amino acid and short homopeptide in both L- and D-enantiomeric forms. Sterile roots of wheat (Triticum aestivum L.) plants were exposed to solutions containing either (14)C-labelled L-alanine, D-alanine, L-trialanine or D-trialanine at a concentration likely to be found in soil solution (10 µM). Over 5 h, plants took up L-alanine, D-alanine and L-trialanine at rates of 0.9±0.3, 0.3±0.06 and 0.3±0.04 µmol g(-1) root DW h(-1), respectively. The rate of N uptake as L-trialanine was the same as that as L-alanine. Plants lost ca.60% of amino acid C taken up in respiration, regardless of the enantiomeric form, but more (ca.80%) of the L-trialanine C than amino acid C was respired. When supplied in solutions of mixed N form, N uptake as D-alanine was ca.5-fold faster than as NO(3)(-), but slower than as L-alanine, L-trialanine and NH(4)(+). Plants showed a limited capacity to take up D-trialanine (0.04±0.03 µmol g(-1) root DW h(-1)), but did not appear to be able to metabolise it. We conclude that wheat is able to utilise L-peptide and D-amino acid N at rates comparable to those of N forms of acknowledged importance, namely L-amino acids and inorganic N. This is true even when solutes are supplied at realistic soil concentrations and when other forms of N are available. We suggest that it may be necessary to reconsider which forms of soil N are important in the terrestrial N cycle.