Nutrition vs association: plant defenses are altered by arbuscular mycorrhizal fungi association not by nutritional provisioning alone.

BACKGROUND: While it is known that arbuscular mycorrhizal fungi (AMF) can improve nutrient acquisition and herbivore resistance in crops, the mechanisms by which AMF influence plant defense remain unknown. Plants respond to herbivory with a cascade of gene expression and phytochemical biosynthesis. Given that the production of defensive phytochemicals requires nutrients, a commonly invoked hypothesis is that the improvement to plant defense when grown with AMF is simply due to an increased availability of nutrients. An alternative hypothesis is that the AMF effect on herbivory is due to changes in plant defense gene expression that are not simply due to nutrient availability. In this study, we tested whether changes in plant defenses are regulated by nutritional provisioning alone or the response of plant to AMF associations. Maize plants grown with or without AMF and with one of three fertilizer treatments (standard, 2 × nitrogen, or 2 × phosphorous) were infested with fall armyworm (Spodoptera frugiperda; FAW) for 72 h. We measured general plant characteristics (e.g. height, number of leaves), relative gene expression (rtPCR) of three defensive genes (lox3, mpi, and pr5), total plant N and P nutrient content, and change in FAW mass per plant. RESULTS: We found that AMF drove the defense response of maize by increasing the expression of mpi and pr5. Furthermore, while AMF increased the total phosphorous content of maize it had no impact on maize nitrogen. Fertilization alone did not alter upregulation of any of the 3 induced defense genes tested, suggesting the mechanism through which AMF upregulate defenses is not solely via increased N or P plant nutrition. CONCLUSION: This work supports that maize defense may be optimized by AMF associations alone, reducing the need for artificial inputs when managing FAW.

Cover crop functional types differentially alter the content and composition of soil organic carbon in particulate and mineral-associated fractions.

Cover crops (CCs) can increase soil organic carbon (SOC) sequestration by providing additional OC residues, recruiting beneficial soil microbiota, and improving soil aggregation and structure. The various CC species that belong to distinct plant functional types (PFTs) may differentially impact SOC formation and stabilization. Biogeochemical theory suggests that selection of PFTs with distinct litter quality (C:N ratio) should influence the pathways and magnitude of SOC sequestration. Yet, we lack knowledge on the effect of CCs from different PFTs on the quantity and composition of physiochemical pools of SOC. We sampled soils under monocultures of three CC PFTs (legume [crimson clover]; grass [triticale]; and brassica [canola]) and a mixture of these three species, from a long-term CC experiment in Pennsylvania, USA. We measured C content in bulk soil and C content and composition in contrasting physical fractions: particulate organic matter, POM; and mineral-associated organic matter, MAOM. The bulk SOC content was higher in all CC treatments compared to the fallow. Compared to the legume, monocultures of grass and brassica with lower litter quality (wider C:N) had higher proportion of plant-derived C in POM, indicating selective preservation of complex structural plant compounds. In contrast, soils under legumes had greater accumulation of microbial-derived C in MAOM. Our results for the first time, revealed that the mixture contributed to a higher concentration of plant-derived compounds in POM relative to the legume, and a greater accumulation of microbial-derived C in MAOM compared to monocultures of grass and brassica. Mixtures with all three PFTs can thus increase the short- and long-term SOC persistence balancing the contrasting effects on the chemistries in POM and MAOM imposed by monoculture CC PFTs. Thus, despite different cumulative C inputs in CC treatments from different PFTs, the total SOC stocks did not vary between CC PFTs, rather PFTs impacted whether C accumulated in POM or MAOM fractions. This highlights that CCs of different PFTs may shift the dominant SOC formation pathways (POM vs. MAOM), subsequently impacting short- and long-term SOC stabilization and stocks. Our work provides a strong applied field test of biogeochemical theory linking litter quality to pathways of C accrual in soil.

Organic fertility inputs synergistically increase denitrification-derived nitrous oxide emissions in agroecosystems.

Soil fertility in organic agriculture relies on microbial cycling of nutrient inputs from legume cover crops and animal manure. However, large quantities of labile carbon (C) and nitrogen (N) in these amendments may promote the production and emission of nitrous oxide (N2 O) from soils. Better ecological understanding of the N2 O emission controls may lead to new management strategies to reduce these emissions. We measured soil N2 O emission for two growing seasons in four corn-soybean-winter grain rotations with tillage, cover crop, and manure management variations typical of organic agriculture in temperate and humid North America. To identify N2 O production pathways and mitigation opportunities, we supplemented N2 O flux measurements with determinations of N2 O isotopomer composition and microbiological genomic DNA abundances in microplots where we manipulated cover crop and manure additions. The N input from legume-rich cover crops and manure prior to corn planting made the corn phase the main source of N2 O emissions, averaging 9.8 kg/ha of N2 O-N and representing 80% of the 3-yr rotations' total emissions. Nitrous oxide emissions increased sharply when legume cover crop and manure inputs exceeded 1.8 and 4 Mg/ha (dry matter), respectively. Removing the legume aboveground biomass before corn planting to prevent co-location of fresh biomass and manure decreased N2 O emissions by 60% during the corn phase. The co-occurrence of peak N2 O emission and high carbon dioxide emission suggests that oxygen (O2 ) consumption likely caused hypoxia and bacterial denitrification. This interpretation is supported by the N2 O site preference values trending towards denitrification during peak emissions with limited N2 O reduction, as revealed by the N2 O δ15 N and δ18 O and the decrease in clade I nosZ gene abundance following incorporation of cover crops and manure. Thus, accelerated microbial O2 consumption seems to be a critical control of N2 O emissions in systems with large additions of decomposable C and N substrates. Because many agricultural systems rely on combined fertility inputs from legumes and manures, our research suggests that controlling the rate and timing of organic input additions, as well as preventing the co-location of legume cover crops and manure, could mitigate N2 O emissions.

COSORE: A community database for continuous soil respiration and other soil-atmosphere greenhouse gas flux data.

Globally, soils store two to three times as much carbon as currently resides in the atmosphere, and it is critical to understand how soil greenhouse gas (GHG) emissions and uptake will respond to ongoing climate change. In particular, the soil-to-atmosphere CO2 flux, commonly though imprecisely termed soil respiration (RS ), is one of the largest carbon fluxes in the Earth system. An increasing number of high-frequency RS measurements (typically, from an automated system with hourly sampling) have been made over the last two decades; an increasing number of methane measurements are being made with such systems as well. Such high frequency data are an invaluable resource for understanding GHG fluxes, but lack a central database or repository. Here we describe the lightweight, open-source COSORE (COntinuous SOil REspiration) database and software, that focuses on automated, continuous and long-term GHG flux datasets, and is intended to serve as a community resource for earth sciences, climate change syntheses and model evaluation. Contributed datasets are mapped to a single, consistent standard, with metadata on contributors, geographic location, measurement conditions and ancillary data. The design emphasizes the importance of reproducibility, scientific transparency and open access to data. While being oriented towards continuously measured RS , the database design accommodates other soil-atmosphere measurements (e.g. ecosystem respiration, chamber-measured net ecosystem exchange, methane fluxes) as well as experimental treatments (heterotrophic only, etc.). We give brief examples of the types of analyses possible using this new community resource and describe its accompanying R software package.

Slow carbon and nutrient accumulation in trees established following fire exclusion in the southwestern United States.

Increasing tree density that followed fire exclusion after the 1880s in the southwestern United States may have also altered nutrient cycles and led to a carbon (C) sink that constitutes a significant component of the U.S. C budget. Yet, empirical data quantifying century-scale changes in C or nutrients due to fire exclusion are rare. We used tree-ring reconstructions of stand structure from five ponderosa pine-dominated sites from across northern Arizona to compare live tree C, nitrogen (N), and phosphorus (P) storage between the 1880s and 1990s. Live tree biomass in the 1990s contained up to three times more C, N, and P than in 1880s. However, the increase in C storage was smaller than values used in recent U.S. C budgets. Furthermore, trees that had established prior to the 1880s accounted for a large fraction (28-66%) of the C, N, and P stored in contemporary stands. Overall, our century-scale analysis revealed that forests of the 1880s were on a trajectory to accumulate C and nutrients in trees even in the absence of fire exclusion, either because growing conditions became more favorable after the 1880s or because forests in the 1880s included age or size cohorts poised for accelerated growth. These results may lead to a reduction in the C sink attributed to fire exclusion, and they refine our understanding of reference conditions for restoration management of fire-prone forests.

Drivers of nitrogen dynamics in ecologically based agriculture revealed by long-term, high-frequency field measurements.

Nitrogen (N) loss from agriculture impacts ecosystems worldwide. One strategy to mitigate these losses, ecologically based nutrient management (ENM), seeks to recouple carbon (C) and N cycles to reduce environmental losses and supply N to cash crops. However, our capacity to apply ENM is limited by a lack of field-based high-resolution data on N dynamics in actual production contexts. We used data from a five-year study of organic cropping systems to investigate soil inorganic N (SIN) variability and nitrate (NO3-) leaching in ENM. Four production systems initiated in 2007 and 2008 in central Pennsylvania varied in crop rotation, timing and intensity of tillage, inclusion of fallow periods, and N inputs. Extractable SIN was measured fortnightly from March through November throughout the experiment, and NO3- N concentration below the rooting zone was sampled with lysimeters during the first year of the 2008 start. We used recursive partitioning models to assess the importance of management and environmental factors to SIN variability and NO3- leaching and identify interactions between influential variables. Air temperature and tillage were the most important drivers of SIN across systems. The highest SIN concentrations occurred when the average air temperature three weeks prior to measurement was above 21 degrees C. Above this temperature and within 109 days of moldboard plowing, average SIN concentrations were 22.1 mg N/kg soil; 109 days or more past plowing average SIN dropped to 7.7 mg N/kg soil. Other drivers of SIN dynamics were N available from manure and cover crops. Highest average leachate NO3- N concentrations (15.2 ppm) occurred in fall and winter when SIN was above 4.9 mg/kg six weeks prior to leachate collection. Late season tillage operations leading to elevated SIN and leachate NO3- N concentrations were a strategy to reduce weeds while meeting consumer demand for organic products. Thus, while tillage that incorporates organic N inputs preceding cash crops can promote synchrony of N mineralization and crop demand, late or post-season tillage promotes NO3 leaching by stimulating SIN pulses that are asynchronous with plant uptake.

Stand-replacing wildfires increase nitrification for decades in southwestern ponderosa pine forests.

Stand-replacing wildfires are a novel disturbance within ponderosa pine (Pinus ponderosa) forests of the southwestern United States, and they can convert forests to grasslands or shrublands for decades. While most research shows that soil inorganic N pools and fluxes return to pre-fire levels within a few years, we wondered if vegetation conversion (ponderosa pine to bunchgrass) following stand-replacing fires might be accompanied by a long-term shift in N cycling processes. Using a 34-year stand-replacing wildfire chronosequence with paired, adjacent unburned patches, we examined the long-term dynamics of net and gross nitrogen (N) transformations. We hypothesized that N availability in burned patches would become more similar to those in unburned patches over time after fire as these areas become re-vegetated. Burned patches had higher net and gross nitrification rates than unburned patches (P < 0.01 for both), and nitrification accounted for a greater proportion of N mineralization in burned patches for both net (P < 0.01) and gross (P < 0.04) N transformation measurements. However, trends with time-after-fire were not observed for any other variables. Our findings contrast with previous work, which suggested that high nitrification rates are a short-term response to disturbance. Furthermore, high nitrification rates at our site were not simply correlated with the presence of herbaceous vegetation. Instead, we suggest that stand-replacing wildfire triggers a shift in N cycling that is maintained for at least three decades by various factors, including a shift from a woody to an herbaceous ecosystem and the presence of fire-deposited charcoal.

Effects of dietary protein concentration on ammonia volatilization, nitrate leaching, and plant nitrogen uptake from dairy manure applied to lysimeters.

This lysimeter experiment was designed to investigate the effects of dietary crude protein (CP) concentration on nitrate-N (NO-N) and ammonia (NH) losses from dairy manure applied to soil and manure N used for plant growth. Lactating dairy cows were fed diets with 16.7% CP (HighCP) or 14.8% CP (LowCP) content. Feces and urine were labeled with N by ruminal pulse-doses of NHCl. Unlabeled and N-labeled feces and urine were used to produce manure for a study with 21 lysimeters in a greenhouse. Manure application rate was 277 kg N ha. Ammonia emissions were measured at 3, 8, 23, 28, 54, and 100 h after manure application. Manure was incorporated into the soil, and a leaching event was simulated. Spring barley was planted (387 plants per m) 7 d after the leaching event and harvested at senescence. Ammonia emission rates and the contribution of urinary N to NO-N were on average about 100% greater for HighCP vs. LowCP manures. With both LowCP and HighCP manures, a greater proportion of urinary vs. fecal N was recovered in leachate NO-N. There was no difference in whole-crop barley N yields between LowCP and HighCP manures, but barley kernel N yield tended to be greater ( = 0.09) for lysimeters treated with HighCP manures. Using a unique labeling approach, this lysimeter experiment demonstrated that when applied at equal soil N application rates, manure from cows fed the HighCP diet resulted in markedly greater NH emissions and urinary N losses with leachate NO-N than manure from cows fed the LowCP diet.

Simulation of Nitrous Oxide Emissions and Estimation of Global Warming Potential in Turfgrass Systems Using the DAYCENT Model.

Nitrous oxide (NO) emissions are an important component of the greenhouse gas budget for turfgrasses. To estimate NO emissions and global warming potential, the DAYCENT ecosystem model was parameterized and applied to turfgrass ecosystems. The annual cumulative NO emissions predicted by the DAYCENT model were close to the measured emission rates of Kentucky bluegrass ( L.) sites in Colorado (within 16% of the observed values). For the perennial ryegrass ( L.) site in Kansas, the DAYCENT model initially overestimated the NO emissions for all treatments (urea and ammonium sulfate at 250 kg N ha yr and urea at 50 kg N ha yr) by about 200%. After including the effect of biological nitrification inhibition in the root exudate of perennial ryegrass, the DAYCENT model correctly simulated the NO emissions for all treatments (within 8% of the observed values). After calibration and validation, the DAYCENT model was used to simulate NO emissions and carbon sequestration of a Kentucky bluegrass lawn under a series of management regimes. The model simulation suggested that gradually reducing fertilization as the lawn ages from 0 to 50 yr would significantly reduce long-term NO emissions by approximately 40% when compared with applying N at a constant rate of 150 kg N ha yr. Our simulation indicates that a Kentucky bluegrass lawn in Colorado could change from a sink to a weak source of greenhouse gas emissions 20 to 30 yr after establishment.

Eliminating Artificial Boundary Conditions in Time-Dependent Density Functional Theory Using Fourier Contour Deformation.

We present an efficient method for propagating the time-dependent Kohn-Sham equations in free space, based on the recently introduced Fourier contour deformation (FCD) approach. For potentials which are constant outside a bounded domain, FCD yields a high-order accurate numerical solution of the time-dependent Schrödinger equation directly in free space, without the need for artificial boundary conditions. Of the many existing artificial boundary condition schemes, FCD is most similar to an exact nonlocal transparent boundary condition, but it works directly on Cartesian grids in any dimension, and runs on top of the fast Fourier transform rather than fast algorithms for the application of nonlocal history integral operators. We adapt FCD to time-dependent density functional theory (TDDFT), and describe a simple algorithm to smoothly and automatically truncate long-range Coulomb-like potentials to a time-dependent constant outside of a bounded domain of interest, so that FCD can be used. This approach eliminates errors originating from the use of artificial boundary conditions, leaving only the error of the potential truncation, which is controlled and can be systematically reduced. The method enables accurate simulations of ultrastrong nonlinear electronic processes in molecular complexes in which the interference between bound and continuum states is of paramount importance. We demonstrate results for many-electron TDDFT calculations of absorption and strong field photoelectron spectra for one and two-dimensional models, and observe a significant reduction in the size of the computational domain required to achieve high quality results, as compared with the popular method of complex absorbing potentials.

Using fixed-potential electrodes to quantify iron and manganese redox cycling in upland soils.

Although metal redox reactions in soils can strongly affect carbon mineralization and other important soil processes, little is known about temporal variations in this redox cycling. Recently, potentiostatically poised electrodes (fixed-potential electrodes) have shown promise for measuring the rate of oxidation and reduction at a specific reduction potential in situ in riparian soils. Here for the first time, we used these electrodes in unsaturated soils to explore the fine-scale temporal redox fluctuations of both iron and manganese in response to environmental conditions. We used three-electrode systems with working electrodes fixed at 100 mV (vs. SHE) and 400 mV at 50 cm and 70 cm in the valley floor soil of a headwater watershed. Electrodes fixed at 100 mV to mimic iron oxides and at 400 mV to mimic manganese oxides allowed real-time reduction and oxidation rates to be calculated from temporal variations in the electric current. Electrode measurements were compared to soil porewater chemistry, pCO2, pO2, groundwater level, resistivity measurements, and precipitation. The fixed-potential electrodes recorded fluctuations over timescales from minutes to weeks. A consistently negative current was observed at 100 mV (interpreted as oxidation of Fe), while the 400-mV electrode fluctuated between negative and positive currents (Mn oxidation and reduction). When the water table rose above the electrodes, reduction was promoted, but above the water table, rainfall only stimulated oxidation. Precipitation frequency thus drove the multi-day reduction or oxidation events (return interval of 5-10 days). These measurements represent the first direct detections of frequency, period, and amplitude of oxidation and reduction events in unsaturated soils. Fixed-potential electrodes hold promise for accurately exploring the fast-changing biogeochemical impacts of metal redox cycling in soils and represent a significant advance for reactions that have been difficult to quantify.

Leveraging microbiome rediversification for the ecological rescue of soil function.

BACKGROUND: Global biodiversity losses threaten ecosystem services and can impact important functional insurance in a changing world. Microbial diversity and function can become depleted in agricultural systems and attempts to rediversify agricultural soils rely on either targeted microbial introductions or retaining natural lands as biodiversity reservoirs. As many soil functions are provided by a combination of microbial taxa, rather than outsized impacts by single taxa, such functions may benefit more from diverse microbiome additions than additions of individual commercial strains. In this study, we measured the impact of soil microbial diversity loss and rediversification (i.e. rescue) on nitrification by quantifying ammonium and nitrate pools. We manipulated microbial assemblages in two distinct soil types, an agricultural and a forest soil, with a dilution-to-extinction approach and performed a microbiome rediversification experiment by re-introducing microorganisms lost from the dilution. A microbiome water control was included to act as a reference point. We assessed disruption and potential restoration of (1) nitrification, (2) bacterial and fungal composition through 16S rRNA gene and fungal ITS amplicon sequencing and (3) functional genes through shotgun metagenomic sequencing on a subset of samples. RESULTS: Disruption of nitrification corresponded with diversity loss, but nitrification was successfully rescued in the rediversification experiment when high diversity inocula were introduced. Bacterial composition clustered into groups based on high and low diversity inocula. Metagenomic data showed that genes responsible for the conversion of nitrite to nitrate and taxa associated with nitrogen metabolism were absent in the low diversity inocula microcosms but were rescued with high diversity introductions. CONCLUSIONS: In contrast to some previous work, our data suggest that soil functions can be rescued by diverse microbiome additions, but that the concentration of the microbial inoculum is important. By understanding how microbial rediversification impacts soil microbiome performance, we can further our toolkit for microbial management in human-controlled systems in order to restore depleted microbial functions.

Dryland irrigation increases accumulation rates of pedogenic carbonate and releases soil abiotic CO2.

Agricultural fields in drylands are challenged globally by limited freshwater resources for irrigation and also by elevated soil salinity and sodicity. It is well known that pedogenic carbonate is less soluble than evaporate salts and commonly forms in natural drylands. However, few studies have evaluated how irrigation loads dissolved calcium and bicarbonate to agricultural fields, accelerating formation rates of secondary calcite and simultaneously releasing abiotic CO2 to the atmosphere. This study reports one of the first geochemical and isotopic studies of such "anthropogenic" pedogenic carbonates and CO2 from irrigated drylands of southwestern United States. A pecan orchard and an alfalfa field, where flood-irrigation using the Rio Grande river is a common practice, were compared to a nearby natural dryland site. Strontium and carbon isotope ratios show that bulk pedogenic carbonates in irrigated soils at the pecan orchard primarily formed due to flood-irrigation, and that approximately 20-50% of soil CO2 in these irrigated soils is calcite-derived abiotic CO2 instead of soil-respired or atmospheric origins. Multiple variables that control the salt buildup in this region are identified and impact the crop production and soil sustainability regionally and globally. Irrigation intensity and water chemistry (irrigation water quantity and quality) dictate salt loading, and soil texture governs water infiltration and salt leaching. In the study area, agricultural soils have accumulated up to 10 wt% of calcite after just about 100 years of cultivation. These rates will likely increase in the future due to the combined effects of climate variability (reduced rainfall and more intense evaporation), use of more brackish groundwater for irrigation, and reduced porosity in soils. The enhanced accumulation rates of pedogenic carbonate are accompanied by release of large amounts of abiotic CO2 from irrigated drylands to atmosphere. Extensive field studies and modelling approaches are needed to further quantify these effluxes at local, regional and global scales.

Farm-scale differentiation of active microbial colonizers.

Microbial movement is important for replenishing lost soil microbial biodiversity and driving plant root colonization, particularly in managed agricultural soils, where microbial diversity and composition can be disrupted. Despite abundant survey-type microbiome data in soils, which are obscured by legacy DNA and microbial dormancy, we do not know how active microbial pools are shaped by local soil properties, agricultural management, and at differing spatial scales. To determine how active microbial colonizers are shaped by spatial scale and environmental conditions, we deployed microbial traps (i.e. sterile soil enclosed by small pore membranes) containing two distinct soil types (forest; agricultural), in three neighboring locations, assessing colonization through 16S rRNA gene and fungal ITS amplicon sequencing. Location had a greater impact on fungal colonizers (R2 = 0.31 vs. 0.26), while the soil type within the microbial traps influenced bacterial colonizers more (R2 = 0.09 vs. 0.02). Bacterial colonizers showed greater colonization consistency (within-group similarity) among replicate communities. Relative to bacterial colonizers, fungal colonizers shared a greater compositional overlap to sequences from the surrounding local bulk soil (R2 = 0.08 vs. 0.29), suggesting that these groups respond to distinct environmental constraints and that their in-field management may differ. Understanding how environmental constraints and spatial scales impact microbial recolonization dynamics and community assembly are essential for identifying how soil management can be used to shape agricultural microbiomes.

Ecosystem response to nutrient enrichment across an urban airshed in the Sonoran Desert.

Rates of nitrogen (N) deposition have increased in arid and semiarid ecosystems, but few studies have examined the impacts of long-term N enrichment on ecological processes in deserts. We conducted a multiyear, nutrient-addition study within 15 Sonoran Desert sites across the rapidly growing metropolitan area of Phoenix, Arizona (USA). We hypothesized that desert plants and soils would be sensitive to N enrichment, but that these effects would vary among functional groups that differ in terms of physiological responsiveness, proximity to surface N sources, and magnitude of carbon (C) or water limitation. Inorganic N additions augmented net potential nitrification in soils, moreso than net potential N mineralization, highlighting the important role of nitrifying microorganisms in the nitrate economy of drylands. Winter annual plants were also responsive to nutrient additions, exhibiting a climate-driven cascade of resource limitation, from little to no production in seasons of low rainfall (winter 2006 and 2007), to moderate N limitation with average precipitation (winter 2009), to limitation by both N and P in a season of above-normal rainfall (winter 2008). Herbaceous production is a potentially important mechanism of N retention in arid ecosystems, capable of immobilizing an amount equal to or greater than that deposited annually to soils in this urban airshed. However, interannual variability in precipitation and abiotic processes that limit the incorporation of detrital organic matter into soil pools may limit this role over the long term. In contrast, despite large experimental additions of N and P over four years, growth of Larrea tridentata, the dominant perennial plant of the Sonoran Desert, was unresponsive to nutrient enrichment, even during wet years. Finally, there did not appear to be strong ecological interactions between nutrient addition and location relative to the city, despite the nearby activity of nearly four million people, perhaps due to loss or transfer pathways that limit long-term N enrichment of ecosystems by the urban atmosphere.

Mitigating nitrogen pollution with under-sown legume-grass cover crop mixtures in winter cereals.

Nitrogen (N) pollution from N inputs to agricultural soils contributes to widespread eutrophication and global climate change. One period susceptible to N losses is between winter grain harvest in summer and corn planting in spring in a corn (Zea mays L.)-soybean [Glycine max (L.) Merr.]-winter grain rotation. Cover crops used to immobilize N during this period often depend on tillage, which can exacerbate N losses. Therefore, we evaluated whether reduced-till cover crops could decrease nitrate (NO3 - ) leaching and nitrous oxide (N2 O) emissions during this period. We tested this strategy in a cropping systems experiment on a 4-ha plot in central Pennsylvania over 2 yr. This experiment compared a clover (Trifolium pratense L.)-timothy (Phleum pratense L.) cover crop no-till underseeded into a standing spelt crop with a vetch (Vicia villosa Roth)-triticale (× Triticosecale Wittm. ex A. Camus) cover crop established with tillage after spelt harvest. These systems were compared based on fortnightly N2 O emissions using static chambers (n = 4 per six sample dates) and potential NO3 - leaching using anion resin bags (n = 4 per system per year). Reduced-till cover crops minimized peak N2 O emissions during the fall compared with tilled cover crops. However, reduced-till cover crops did not decrease potentially leachable NO3 - relative to tilled cover crops despite decreases in soil inorganic N. Cover crop N isotopes revealed that clover N may have mineralized and leached over the winter. Our results suggest that reduced-till cover crops can decrease N2 O emissions to mitigate the climate impact of agriculture but that winter-hardy cover crops should be chosen to mitigate leaching.

Cover Crop Soil Legacies Alter Phytochemistry and Resistance to Fall Armyworm (Lepidoptera: Noctuidae) in Maize.

Plant-soil feedbacks can mediate aboveground plant-herbivore interactions by impacting plant chemistry. Given that soil legacies and agricultural practices are closely tied, a better understanding of soil legacy cascades and their application in pest management are needed. We tested how cover crop legacies alter resistance to fall armyworm (Spodoptera frugiperda Smith, Lepidoptera: Noctuidae) in maize (Zea mays L., Poales: Poaceae). We compared herbivore performance and behavior of fall armyworm larvae on maize grown after four cover crop treatments: a leguminous mycorrhizal cover crop (pea: Pisum sativum L., Fabales: Fabaceae), a nonleguminous mycorrhizal cover crop (triticale: x Triticosecale Wittm. Ex A. Camus, Poales: Poaceae), a nonleguminous nonmycorrhizal cover crop (radish: Raphanus sativus L., Brassicales: Brassicaceae), and no cover crops (fallow). Soil inorganic N was highest in pea treatments and lowest in triticale treatments, while maize AMF colonization was greatest when grown after mycorrhizal cover crops compared to nonmycorrhizal or no cover crops. Cover crop legacies altered the emission of maize volatiles and fall armyworm larvae oriented toward odors emitted by maize grown after radish more frequently than triticale in olfactometer assays. Additionally, larvae performed better and consumed more leaf tissue when feeding on maize grown after radish and poorest on plants grown after triticale. When damaged by fall armyworm, maize grown after triticale expressed higher levels of lipoxygenase-3 (lox3), while plants grown after radish upregulated maize proteinase inhibitor (mpi) gene expression. Our results highlight the importance of appropriate cover crop selection and suggest that triticale could strengthen maize resistance to fall armyworm.

Plant and soil carbon accumulation following fire in Mediterranean woodlands in Spain.

We measured plant and soil carbon (C) storage following canopy-replacing wildfires in woodlands of northeastern Spain that include an understory of shrubs dominated by Quercus coccifera and an overstory of Pinus halepensis trees. Established plant succession models predict rapid shrub recovery in these ecosystems, and we build on this model by contrasting shrub succession with long-term C storage in soils, trees, and the whole ecosystem. We used chronosequence and repeated sampling approaches to detect change over time. Aboveground plant C increased from <100 to ~3,000 g C m(-2) over 30 years following fire, which is substantially less than the 5,942 ± 487 g C m(-2) (mean ±1 standard error) in unburned sites. As expected, shrubs accumulated C rapidly, but the capacity for C storage in shrubs was <600 g C m(-2). Pines were the largest plant C pool in sites >20 years post fire, and accounted for all of the difference in plant C between older burned sites and unburned sites. In contrast, soil C was initially higher in burned sites (~4,500 g C m(-2)) than in unburned sites (3,264 ± 261 g C m(-2)) but burned site C declined to unburned levels within 10 years after fire. Combining these results with prior research suggests two states for C storage. When pine regeneration is successful, ~9,200 g C m(-2) accumulate in woodlands but when tree regeneration fails (due to microclimatic stress or short fire return intervals), ecosystem C storage of ~4,000 g C m(-2) will occur in the resulting shrublands.

Short-term soil inorganic N pulse after experimental fire alters invasive and native annual plant production in a Mojave Desert shrubland.

Post-fire changes in desert vegetation patterns are known, but the mechanisms are poorly understood. Theory suggests that pulse dynamics of resource availability confer advantages to invasive annual species, and that pulse timing can influence survival and competition among species. Precipitation patterns in the American Southwest are predicted to shift toward a drier climate, potentially altering post-fire resource availability and consequent vegetation dynamics. We quantified post-fire inorganic N dynamics and determined how annual plants respond to soil inorganic nitrogen variability following experimental fires in a Mojave Desert shrub community. Soil inorganic N, soil net N mineralization, and production of annual plants were measured beneath shrubs and in interspaces during 6 months following fire. Soil inorganic N pools in burned plots were up to 1 g m(-2) greater than unburned plots for several weeks and increased under shrubs (0.5-1.0 g m(-2)) more than interspaces (0.1-0.2 g m(-2)). Soil NO(3) (-)-N (nitrate-N) increased more and persisted longer than soil NH(4) (+)-N (ammonium-N). Laboratory incubations simulating low soil moisture conditions, and consistent with field moisture during the study, suggest that soil net ammonification and net nitrification were low and mostly unaffected by shrub canopy or burning. After late season rains, and where soil inorganic N pools were elevated after fire, productivity of the predominant invasive Schismus spp. increased and native annuals declined. Results suggest that increased N availability following wildfire can favor invasive annuals over natives. Whether the short-term success of invasive species following fire will direct long-term species composition changes remains to be seen, yet predicted changes in precipitation variability will likely interact with N cycling to affect invasive annual plant dominance following wildfire.

Cover crop mixture expression is influenced by nitrogen availability and growing degree days.

Cover crop mixtures can provide multiple ecosystem services but provisioning of these services is contingent upon the expression of component species in the mixture. From the same seed mixture, cover crop mixture expression varied greatly across farms and we hypothesized that this variation was correlated with soil inorganic nitrogen (N) concentrations and growing degree days. We measured fall and spring biomass of a standard five-species mixture of canola (Brassica napus L.), Austrian winter pea (Pisum sativum L), triticale (x Triticosecale Wittm.), red clover (Trifolium pratense L.) and crimson clover (Trifolium incarnatum L.) seeded at a research station and on 8 farms across Pennsylvania and New York in two consecutive years. At the research station, soil inorganic N (soil iN) availablity and cumulative fall growing degree days (GDD) were experimentally manipulated through fertilizer additions and planting date. Farmers seeded the standard mixture and a "farm-tuned" mixture of the same five species with component seeding rates adjusted to achieve farmer-desired services. We used Structural Equation Modeling to parse out the effects of soil iN and GDD on cover crop mixture expression. When soil iN and fall GDD were high, canola dominated the mixture, especially in the fall. Low soil iN favored legume species while a shorter growing season favored triticale. Changes in seeding rates influenced mixture composition in fall and spring but interacted with GDD to determine the final expression of the mixture. Our results show that when soil iN availability is high at the time of cover crop planting, highly competitive species can dominate mixtures which could potentially decrease services provided by other species, especially legumes. Early planting dates can exacerbate the dominance of aggressive species. Managers should choose cover crop species and seeding rates according to their soil iN and GDD to ensure the provision of desired services.