Soil temperature and moisture as key controls of phosphorus export in mountain watersheds.

Oligotrophic mountain lakes act as sensitive indicators of landscape-scale changes in mountain regions due to their low nutrient concentration and remote, relatively undisturbed watersheds. Recent research shows that phosphorus (P) concentrations are increasing in mountain lakes around the world, creating more mesotrophic states and altering lake ecosystem structure and function. The relative importance of atmospheric deposition and climate-driven changes to local biogeochemistry in driving these shifts is not well established. In this study, we test whether increasing temperatures in watershed soils may be contributing to the observed increases in mountain lake P loading. Specifically, we test whether higher soil temperatures increase P mobilization from mountain soils by accelerating the rate of geochemical weathering and soil organic matter decomposition. We used paired soil incubation (lab) and soil transplant (field) experiments with mountain soils from around the western United States to test the effects of warming on rain-leachable P concentration, soil P mobilization, and soil respiration. Our results show that while higher temperature can increase soil P mobilization, low soil moisture can limit the effects of warming in some situations. Soils with lower bulk densities, higher pH, lower aluminum oxide contents, and lower ratios of carbon to nitrogen had much higher rain-leachable P concentration across all sites and experimental treatments. Together, these results suggest that mountain watersheds with high-P soils and relatively high soil moisture could have the largest increases in P mobilization with warming. Consequently, lakes and streams in such watersheds could become especially susceptible to soil-driven eutrophication as temperatures rise.

Evaluation of nutrient assimilative capacity in waterfowl impoundments: The role of environmental stressors.

The effectiveness of wetlands in sequestering nutrients and improving water quality relies on a suite of abiotic and biotic conditions. To more fully understand the restraints on nutrient removal, especially salinity and plant cover, we created field-scale mesocosms and monitored nutrient sequestration with nutrient additions and isotopic pool dilutions over 2 years in two wetlands near the Great Salt Lake in Utah. Surprisingly, we found no differences in nutrient removal with plant removal, increased salinity, and altered ambient nutrient concentrations, suggesting functional redundancy in associated primary producers. When submerged aquatic vegetation was removed, chlorophyll α concentration (0.1-9.0 µg/L) increased while nitrogen (N) and phosphorus (P) assimilation remained the same as phytoplankton occupied the open niche space. We did find ammonium concentrations to be inversely related to nitrate assimilation-as the ammonium concentration increased, nitrate assimilation decreased, suggesting preferential uptake of ammonium. Last, in our high N and P treatment mesocosms, the nitrate dramatically declined from 43.9 mg/L to background levels (<0.1 mg/L) within 1 week, showing a high potential for N remediation in these wetlands.

Carbon control on terrestrial ecosystem function across contrasting site productivities: the carbon connection revisited.

Understanding how altered soil organic carbon (SOC) availability affects microbial communities and their function is imperative in predicting impacts of global change on soil carbon (C) storage and ecosystem function. However, the response of soil microbial communities and their function to depleted C availability in situ is unclear. We evaluated the role of soil C inputs in controlling microbial biomass, community composition, physiology, and function by (1) experimentally excluding plant C inputs in situ for 9 yr in four temperate forest ecosystems along a productivity gradient in Oregon, USA; and (2) integrating these findings with published data from similar C-exclusion studies into a global meta-analysis. Excluding plant C inputs for 9 yr resulted in a 13% decrease in SOC across the four Oregon sites and an overall shift in the microbial community composition, with a 45% decrease in the fungal : bacterial ratio and a 13% increase in Gram-positive : Gram-negative bacterial ratio. Although gross N mineralization decreased under C exclusion, decreases in gross N immobilization were greater, resulting in increased net N mineralization rates in all but the lowest-productivity site. Microbial biomass showed a variable response to C exclusion that was method dependent; however, we detected a 29% decrease in C-use efficiency across the sites, with greater declines occurring in less-productive sites. Although extracellular enzyme activity increased with C exclusion, C exclusion resulted in a 31% decrease in microbial respiration across all sites. Our meta-analyses of published data with similar C-exclusion treatments were largely consistent with our experimental results, showing decreased SOC, fungal : bacterial ratios, and microbial respiration, and increased Gram-positive : Gram-negative bacterial ratio following exclusion of C inputs to soil. Effect sizes of SOC and respiration correlated negatively with the duration of C exclusion; however, there were immediate effects of C exclusion on microbial community composition and biomass that were unaltered by duration of treatment. Our field-based experimental results and analyses demonstrate unequivocally the dominant control of C availability on soil microbial biomass, community composition, and function, and provide additional insight into the mechanisms for these effects in forest ecosystems.

The invasive annual cheatgrass releases more nitrogen than crested wheatgrass through root exudation and senescence.

Plant-soil feedbacks are an important aspect of invasive species success. One type of feedback is alteration of soil nutrient cycling. Cheatgrass invasion in the western USA is associated with increases in plant-available nitrogen (N), but the mechanism for this has not been elucidated. We labeled cheatgrass and crested wheatgrass, a common perennial grass in western rangelands, with (15)N-urea to determine if differences in root exudates and turnover could be a mechanism for increases in soil N. Mesocosms containing plants were either kept moist, or dried out during the final 10 days to determine the role of senescence in root N release. Soil N transformation rates were determined using (15)N pool dilution. After 75 days of growth, cheatgrass accumulated 30 % more total soil N and organic carbon than crested wheatgrass. Cheatgrass roots released twice as much N as crested wheatgrass roots (0.11 vs. 0.05 mg N kg(-1) soil day(-1)) in both soil moisture treatments. This occurred despite lower root abundance (7.0 vs. 17.3 g dry root kg(-1) soil) and N concentration (6.0 vs. 7.6 g N kg(-1) root) in cheatgrass vs. crested wheatgrass. We propose that increases in soil N pool sizes and transformation rates under cheatgrass are caused by higher rates of root exudation or release of organic matter containing relatively large amounts of labile N. Our results provide the first evidence for the underlying mechanism by which the invasive annual cheatgrass increases N availability and establishes positive plant-soil feedbacks that promote its success in western rangelands.

The invasive annual cheatgrass increases nitrogen availability in 24-year-old replicated field plots.

Previous studies comparing invaded and non-invaded sites suggest that cheatgrass (Bromus tectorum L.) causes soil N cycling to increase. Unfortunately, these correlative studies fail to distinguish whether cheatgrass caused the differences in N cycling, or if cheatgrass simply invaded sites where N availability was greater. We measured soil C and N concentrations and net and gross N-cycling rates on 24-year-old replicated field plots in a sagebrush-steppe ecosystem that had been plowed, fumigated, and seeded to different plant communities in 1984. Laboratory assays of soil collected throughout the soil profiles (0-60 cm) showed that soil NO3 (-), organic C and N, and net N mineralization, net nitrification, and soil respiration rates were all greater beneath cheatgrass than in sagebrush-perennial grass plots. In surface soils (0-10 cm), field and lab assays on five sampling dates during 2 years showed gross N mineralization, net N mineralization, and net nitrification rates were all faster beneath cheatgrass than in sagebrush-perennial grass plots. Modeling analyses based on soil respiration and gross N-cycling rates suggest that cheatgrass provides soil microbes with lower C:N substrates and that this could explain the faster N-cycling rates beneath cheatgrass. This is the first long-term replicated field study to conclusively show that cheatgrass created greater soil organic N pool sizes and stimulated N-cycling rates compared to similar-aged stands of sagebrush and native perennial grasses. Increased N-cycling rates may represent a positive plant-soil feedback that promotes continued dominance by cheatgrass, even in the absence of soil disturbance or fire.

Sagebrush carrying out hydraulic lift enhances surface soil nitrogen cycling and nitrogen uptake into inflorescences.

Plant roots serve as conduits for water flow not only from soil to leaves but also from wetter to drier soil. This hydraulic redistribution through root systems occurs in soils worldwide and can enhance stomatal opening, transpiration, and plant carbon gain. For decades, upward hydraulic lift (HL) of deep water through roots into dry, litter-rich, surface soil also has been hypothesized to enhance nutrient availability to plants by stimulating microbially controlled nutrient cycling. This link has not been demonstrated in the field. Working in sagebrush-steppe, where water and nitrogen limit plant growth and reproduction and where HL occurs naturally during summer drought, we slightly augmented deep soil water availability to 14 HL+ treatment plants throughout the summer growing season. The HL+ sagebrush lifted greater amounts of water than control plants and had slightly less negative predawn and midday leaf water potentials. Soil respiration was also augmented under HL+ plants. At summer's end, application of a gas-based (15)N isotopic labeling technique revealed increased rates of nitrogen cycling in surface soil layers around HL+ plants and increased uptake of nitrogen into HL+ plants' inflorescences as sagebrush set seed. These treatment effects persisted even though unexpected monsoon rainstorms arrived during assays and increased surface soil moisture around all plants. Simulation models from ecosystem to global scales have just begun to include effects of hydraulic redistribution on water and surface energy fluxes. Results from this field study indicate that plants carrying out HL can also substantially enhance decomposition and nitrogen cycling in surface soils.

Regulation and measurement of nitrification in terrestrial systems.

Understanding nitrification rates and their regulation continues as a key area of research for assessing human's increasing impact on the terrestrial N cycle. We review the organisms and processes responsible for nitrification in terrestrial systems. The control of nitrification by substrate availability is discussed with particular attention to the factors affecting ammonia/ammonium availability. The effects on nitrification rates of environmental controls including oxygen, water potential, temperature and pH are described. With this general understanding of the factors affecting nitrification rates as a basis, we present an in depth analysis of methods used to measure nitrification in terrestrial systems. Net, gross and potential nitrification rate measurements are explained including the use of isotopes and inhibitors to measure rates in soils. Methods for the estimation of nitrification kinetics and modeling are briefly described. Future challenges will require understanding the factors controlling nitrification across spatial scales from ecosystems to soil microsites if we are to sustainably manage reactive nitrogen in terrestrial environments.

Nitrification exhibits Haldane kinetics in an agricultural soil treated with ammonium sulfate or dairy-waste compost.

An agricultural soil was treated with dairy-waste compost, ammonium-sulfate fertilizer or no added nitrogen (control) and planted to silage corn for 6 years. The kinetics of nitrification were determined in laboratory-shaken slurry assays with a range of substrate concentrations (0-20 mM NH(4)(+)) over a 24-h period for soils from the three treatments. Determined concentrations of substrate and product were fit to Michaelis-Menten and Haldane models. For all the treatments, the Haldane model was a better fit, suggesting that significant nitrification inhibition may occur in soils under high ammonium conditions similar to those found immediately after fertilization or waste applications. The maximum rate of nitrification (V(max)) was significantly higher for the fertilized and compost-treated soils (1.74 and 1.50 mmol N kg(-1) soil day(-1)) vs. control soil (0.98 mmol kg(-1) soil day(-1)). The K(m) and K(i) values were not significantly different, with average values of 0.02 and 27 mM NH(4)(+), respectively. Our results suggest that both N sources increased nitrifier community size, but did not shift the nitrifier community structure in ways that influenced enzyme affinity or sensitivity to ammonium. The K(m) values are comparable to those determined directly in other soils, but are substantially lower than those from most pure cultures of ammonia-oxidizing bacteria.

Microbial dynamics and carbon and nitrogen cycling following re-wetting of soils beneath two semi-arid plant species.

Sporadic summer rainfall in semi-arid ecosystems can provide enough soil moisture to drastically increase CO(2) efflux and rates of soil N cycling. The magnitudes of C and N pulses are highly variable, however, and the factors regulating these pulses are poorly understood. We examined changes in soil respiration, bacterial, fungal and microfaunal populations, and gross rates of N mineralization, nitrification, and NH(4) (+) and NO(3) (-) immobilization during the 10 days following wetting of dry soils collected from stands of big sagebrush (Artemisia tridentata) and cheatgrass (Bromus tectorum) in central Utah. Soil CO(2) production increased more than tenfold during the 17 h immediately following wetting. The labile organic C pool released by wetting was almost completely respired within 2-3 days, and was nearly three times as large in sagebrush soil as in cheatgrass. In spite of larger labile C pools beneath sagebrush, microbial and microfaunal populations were nearly equal in the two soils. Bacterial and fungal growth coincided with depletion of labile C, and populations peaked in both soils 2 days after wetting. Protozoan populations, whose biomass was nearly 3,000-fold lower than bacteria and fungi, peaked after 2-4 days. Gross N mineralization and nitrification rates were both faster in cheatgrass soil than in sagebrush, and caused greater nitrate accumulation in cheatgrass soil. Grazing of bacteria and fungi by protozoans and nematodes could explain neither temporal trends in N mineralization rates nor differences between soil types. However, a mass balance model indicated that the initial N pulse was associated with degradation of microbial substrates that were rich in N (C:N <8.3), and that microbes had shifted to substrates with lower N contents (C:N =15-25) by day 7 of the incubation. The model also suggested that the labile organic matter in cheatgrass soil had a lower C:N ratio than in sagebrush, and this promoted faster N cycling rates and greater N availability. This study provides evidence that the high N availability often associated with wetting of cheatgrass soils is a result of cheatgrass supplying substrates to microbes that are of high decomposability and N content.

Water pulses and biogeochemical cycles in arid and semiarid ecosystems.

The episodic nature of water availability in arid and semiarid ecosystems has significant consequences on belowground carbon and nutrient cycling. Pulsed water events directly control belowground processes through soil wet-dry cycles. Rapid soil microbial response to incident moisture availability often results in almost instantaneous C and N mineralization, followed by shifts in C/N of microbially available substrate, and an offset in the balance between nutrient immobilization and mineralization. Nitrogen inputs from biological soil crusts are also highly sensitive to pulsed rain events, and nitrogen losses, particularly gaseous losses due to denitrification and nitrate leaching, are tightly linked to pulses of water availability. The magnitude of the effect of water pulses on carbon and nutrient pools, however, depends on the distribution of resource availability and soil organisms, both of which are strongly affected by the spatial and temporal heterogeneity of vegetation cover, topographic position and soil texture. The 'inverse texture hypothesis' for net primary production in water-limited ecosystems suggests that coarse-textured soils have higher NPP than fine-textured soils in very arid zones due to reduced evaporative losses, while NPP is greater in fine-textured soils in higher rainfall ecosystems due to increased water-holding capacity. With respect to belowground processes, fine-textured soils tend to have higher water-holding capacity and labile C and N pools than coarse-textured soils, and often show a much greater flush of N mineralization. The result of the interaction of texture and pulsed rainfall events suggests a corollary hypothesis for nutrient turnover in arid and semiarid ecosystems with a linear increase of N mineralization in coarse-textured soils, but a saturating response for fine-textured soils due to the importance of soil C and N pools. Seasonal distribution of water pulses can lead to the accumulation of mineral N in the dry season, decoupling resource supply and microbial and plant demand, and resulting in increased losses via other pathways and reduction in overall soil nutrient pools. The asynchrony of resource availability, particularly nitrogen versus water due to pulsed water events, may be central to understanding the consequences for ecosystem nutrient retention and long-term effects on carbon and nutrient pools. Finally, global change effects due to changes in the nature and size of pulsed water events and increased asynchrony of water availability and growing season will likely have impacts on biogeochemical cycling in water-limited ecosystems.

Root responses and nitrogen acquisition by Artemisia tridentata and Agropyron desertorum following small summer rainfall events.

Resources in the Great Basin of western North America often occur in pulses, and plant species must rapidly respond to temporary increases in water and nutrients during the growing season. A field study was conducted to evaluate below ground responses of Artemisia tridentata and Agropyron desertorum, common Great Basin shrub and grass species, respectively, to simulated 5-mm (typical summer rain) and 15-mm (large summer rain) summer rainfall events. The simulated rainfall was labeled with K(15)NO(3) so that timing of plant nitrogen uptake could be monitored. In addition, soil NH(4)(+) and NO(3)(-) concentrations and physiological uptake capacities for NO(3)(-) and NH(4)(+) were determined before and after the rainfall events. Root growth in the top 15 cm of soil was monitored using a minirhizotron system. Surprisingly, there was no difference in the amount of labeled N acquired in response to the two rainfall amounts by either species during the 7-day sample period. However, there were differences between species in the timing of labeled N uptake. The N label was detected in above ground tissue of Agropyron within 1 h of the simulated rainfall events, but not until 24 h after the rainfall in Artemisia. For both Agropyron and Artemisia, root uptake capacity was similarly affected by the 5-mm and 15-mm rainfall. There was, however, a greater increase in uptake capacity for NH(4)(+) than for NO(3)(-), and the 15-mm event resulted in a longer response. No root growth occurred in either species in response to either rainfall event during this 8-day period. The results of this study indicate that these species are capable of utilizing nitrogen pulses following even small summer rainfall events during the most stressful period of the summer and further emphasize the importance of small precipitation events in arid systems.