Biogeochemistry of Nitrogen Across the Everglades Landscape

Compared to phosphorus (P), nitrogen (N) has received little attention across the Everglades landscape. Despite this lack of attention, N plays important roles in many Everglades systems, including being a significant pollutant in Florida Bay and the Gulf of Mexico, the limiting nutrient in highly P-impacted areas, and an important substrate for microbial metabolism. Storage and transport of N throughout the Everglades is dominated by organic forms, including peat soils and dissolved organic N in the water column. In general, N sources are highest in the northern areas; however, atmospheric deposition and active N2 fixation by the periphyton components are a significant N source throughout most systems. Many of the processes involved in the wetland N cycle remain unmeasured for most of the Everglades systems. In particular, the lack of in situ rates for N2 fixation and denitrification prevent the construction of system-level budgets, especially for the Southern mangrove systems where N export into Florida Bay is critical. There is also the potential for several novel N processes (e.g., Anammox) with an as yet undetermined importance for nitrogen cycling and function of the Everglades ecosystem. Phosphorus loading alters the N cycle by stimulating organic N mineralization with resulting flux of ammonium and DON, and at elevated P concentrations, by increasing rates of N2 fixation and N assimilation. Restoration of hydrology has a potential for significantly impacting N cycling in the Everglades both in terms of affecting N transport, but also by altering aerobic-anaerobic transitions at the soil-water interface or in areas with seasonal drawdowns (e.g., marl prairies). Based on the authors’ understanding of N processes, much more research is necessary to adequately predict potential impacts from hydrologic restoration, as well as the function of Everglades systems as sinks, sources, and transformers of N in the South Florida landscape.

Biogeochemical Contributions of Tree Islands to Everglades Wetland Landscape Nitrogen Cycling During Seasonal Inundation

In the Florida Everglades, tree islands are conspicuous heterogeneous elements in the herbaceous wetland landscape. We characterized the biogeochemical role of a seasonally flooded tree island during wet season inundation, specifically examining hydrologically mediated flows of nitrogen (N) and N retention by the tree island. We estimated ecosystem N standing stocks and fluxes, soil and litter N transformation rates, and hydrologic fluxes of N to quantify the net ecosystem N mass flux. Results showed that hydrologic sources of N were dominated by surface water loads of nitrate (NO3) and ammonium (NH4). Nitrate immobilization by soils and surficial leaf litter was an important sink for surface water dissolved inorganic N (DIN). We estimated that the net annual DIN retention by a seasonally flooded tree island was 20.5 ± 5.0 g m−2 during wet season inundation. Based on the estimated tree island surface water DIN loading rate, a seasonally flooded tree island retained 76% of imported DIN. As such, seasonally flooded tree islands have the potential to retain 55% of DIN entering the marsh landscape via upstream canal overland flow in the wet season. By increasing reactive surface area and DOC availability, we suggest that tree islands promote convergence of elements that enhance DIN retention. Tree islands of this region are thus important components of landscape-scale restoration efforts that seek to reduce sources of anthropogenic DIN to downstream estuaries.

The Importance of Dissimilatory Nitrate Reduction to Ammonium (DNRA) in the Nitrogen Cycle of Coastal Ecosystems

Until recently, it was believed that biological assimilation and gaseous nitrogen (N) loss through denitrification were the two major fates of nitrate entering or produced within most coastal ecosystems. Denitrification is often viewed as an important ecosystem service that removes reactive N from the ecosystem. However, there is a competing nitrate reduction process, dissimilatory nitrate reduction to ammonium (DNRA), that conserves N within the ecosystem. The recent application of nitrogen stable isotopes as tracers has generated growing evidence that DNRA is a major nitrogen pathway that cannot be ignored. Measurements comparing the importance of denitrification vs. DNRA in 55 coastal sites found that DNRA accounted for more than 30% of the nitrate reduction at 26 sites. DNRA was the dominant pathway at more than one-third of the sites. Understanding what controls the relative importance of denitrification and DNRA, and how the balance changes with increased nitrogen loading, is of critical importance for predicting eutrophication trajectories. Recent improvements in methods for assessing rates of DNRA have helped refine our understanding of the rates and controls of this process, but accurate measurements in vegetated sediment still remain a challenge.

Half-degree gridded nitrogen and phosphorus fertilizer use for global agriculture production during 1900-2013

In addition to enhance agricultural productivity, synthetic nitrogen (N) and phosphorous (P) fertilizer application in croplands dramatically altered global nutrient budget, water quality, greenhouse gas balance, and their feedbacks to the climate system. However, due to the lack of geospatial fertilizer input data, current Earth system/land surface modeling studies have to ignore or use over-simplified data (e.g., static, spatially uniform fertilizer use) to characterize agricultural N and P input over decadal or century-long period. We therefore develop a global time-series gridded data of annual synthetic N and P fertilizer use rate in croplands, matched with HYDE 3,2 historical land use maps, at a resolution of 0.5º latitude by longitude during 1900-2013. Our data indicate N and P fertilizer use rates increased by approximately 8 times and 3 times, respectively, since the year 1961, when IFA (International Fertilizer Industry Association) and FAO (Food and Agricultural Organization) survey of country-level fertilizer input were available. Considering cropland expansion, increase of total fertilizer consumption amount is even larger. Hotspots of agricultural N fertilizer use shifted from the U.S. and Western Europe in the 1960s to East Asia in the early 21st century. P fertilizer input show the similar pattern with additional hotspot in Brazil. We find a global increase of fertilizer N/P ratio by 0.8 g N/g P per decade (p< 0.05) during 1961-2013, which may have important global implication of human impacts on agroecosystem functions in the long run. Our data can serve as one of critical input drivers for regional and global assessment on agricultural productivity, crop yield, agriculture-derived greenhouse gas balance, global nutrient budget, land-to-aquatic nutrient loss, and ecosystem feedback to the climate system.

Comparison of the stable carbon and nitrogen isotopic values of gill and white muscle tissue of fish

The potential use of stable carbon and nitrogen isotope ratios (d13C, d15N) of fish gills for studies on fish feeding ecology was evaluated by comparing the d13C and d15N of gill tissue with the more commonly used white muscle tissue. To account for the effect of lipid content on the d13C signatures, a study-specific lipid correction model based on C:N ratios was developed and applied to the bulk d13C data. For the majority of species in the study, we found no significant difference in d13C values between gill and muscle tissue after correction, but several species showed a small (0.3-1.4 per mil) depletion in 13C in white muscle compared to gill tissue. The average species difference in d15N between muscle and gill tissue ranged from -0.2 to 1.6 per mil for the different fish species with muscle tissue generally more enriched in 15N. The d13C values of muscle and gill were strongly linearly correlated (R**2 = 0.85) over a large isotopic range (13 per mil), suggesting that both tissues can be used to determine long-term feeding or migratory habits of fish. Muscle and gill tissue bulk d15N values were also strongly positively correlated (R**2= 0.76) but with a small difference between muscle and gill tissue. This difference indicates that the bulk d15N of the two tissue types may be influenced by different isotopic turnover rates or a different composition of amino acids.

Data compilation on the biological response to ocean acidification: environmental and experimental context of data sets and related literature

The exponential growth of studies on the biological response to ocean acidification over the last few decades has generated a large amount of data. To facilitate data comparison, a data compilation hosted at the data publisher PANGAEA was initiated in 2008 and is updated on a regular basis (doi:10.1594/PANGAEA.149999). By January 2015, a total of 581 data sets (over 4 000 000 data points) from 539 papers had been archived. Here we present the developments of this data compilation five years since its first description by Nisumaa et al. (2010). Most of study sites from which data archived are still in the Northern Hemisphere and the number of archived data from studies from the Southern Hemisphere and polar oceans are still relatively low. Data from 60 studies that investigated the response of a mix of organisms or natural communities were all added after 2010, indicating a welcomed shift from the study of individual organisms to communities and ecosystems. The initial imbalance of considerably more data archived on calcification and primary production than on other processes has improved. There is also a clear tendency towards more data archived from multifactorial studies after 2010. For easier and more effective access to ocean acidification data, the ocean acidification community is strongly encouraged to contribute to the data archiving effort, and help develop standard vocabularies describing the variables and define best practices for archiving ocean acidification data.

Effects of introduced moose (Alces alces) on vegetation composition, nutrient dynamics, and decomposition rates in boreal forest ecosystems in Newfoundland, Canada

Globally, consumers affect ecosystem processes including nutrient dynamics. Herbivores have been known to slow nutrient flow in boreal forest ecosystems. I examined the effects of introduced moose on disturbed forests of Newfoundland, Canada by conducting a field experiment during August - November 2014 in 20 paired moose exclosure-control plots. I tested whether moose browsing directly and indirectly affected forests by measuring plant species composition, litter quality and quantity, soil quality, and decomposition rates in areas moose exclosure-control plots. I analyzed moose effects using linear mixed effects models and found evidence indicating that moose reduce plant height and litter biomass affecting the availability of carbon, nitrogen, and phosphorus. However, plant diversity, soil quality, and litter decomposition did not differ between moose exclosures and controls. Moose in Newfoundland directly influence plant regeneration and litter biomass while indirect effects on soil ecosystems may be limited by time, disturbance, and climate.

Gut content, fatty acid composition and metabolic rates of the amphipod Anonyx sarsi during POLARSTERN cruise ARK-XIX/1, ice station PS64/070-6

During a winter expedition to the western Barents Sea in March 2003, benthic amphipods of the species Anonyx sarsi were observed directly below pack ice. Only males and juveniles [16.0-37.0 mm long, 16.2-120.8 mg dry mass (DM)] were collected. Guts contained macroalgal fibres, fish eggs and flesh from large carrion. Amphipods had very low levels of total lipids (2.7-17.2% DM). Analysis of lipid biomarkers showed that some of the specimens had preyed on pelagic copepods. Individual respiration rates ranged over 0.4-1.7 ml O2/day (mean: 1.2 ml, SD: 0.5 ml). Individual ammonia excretion rates varied between 7.8 µg and 49.3 µg N/day (mean: 30.7 µg, SD: 15.2 µg). The atomic O:N ratio ranged over 35 to 71 (mean: 55, SD: 14), indicating lipid-dominated metabolism. Mass-specific respiration ranged over 9.8-16.6 ml O2/day/g DM (mean: 13.1 ml, SD: 2.2 ml). The metabolic rates of A. sarsi were twice as high as those of the truly sympagic amphipod Gammarus wilkitzkii, which is better adapted to the under-ice habitat by its energy-saving attached lifestyle. It is concluded that males and juveniles of A. sarsi were actively searching for food in the water column and at the ice underside, but that the nutritional status of the amphipods in late Arctic winter was generally very poor.

Lithological and chemical composition, stages of sediment accumulation, and sedimentation rates in Core PSh24-2537, West Gotland Basin, Baltic Sea

We studied the grain-size, mineral and chemical compositions, physical properties, radiocarbon age, spore-pollen spectra, and diatom composition in sediments from Core PSh24-2537 sampled in the West Gotland Basin. Four lithological-stratigraphic units were distinguished: varved clays of the Baltic Ice Lake, black and black-gray (sulfide) clays of the Yoldian Sea, gray clays of Ancylus Lake, and greenish-gray sapropel-like littorine and post-littorine silts of the marine stage of Holocene. These units differ from each other both in their matter composition and in plant remains. In the littorine silts organic carbon concentra¬tion reached from 1.5 to 10.35%. Conditions of sediment accumulation and the stages of evolution of the West Gotland Basin over the post-glacial time are characterized.

Leaf Traits, Neighbors, and Abiotic Factors: Ways That Context Can Mediate the Impact of Invasive Species on Nitrogen Cycling

Species invasions are more prevalent than ever before. While the addition of a species can dramatically change critical ecosystem processes, factors that mediate the direction and magnitude of those impacts have received less attention. A better understanding of the factors that mediate invasion impacts on ecosystem functioning is needed in order to target which exotic species will be most harmful and which systems are most vulnerable. The role of invasion on nitrogen (N) cycling is particularly important since N cycling controls ecosystem services that provision human health, e.g. nutrient retention and water quality.

We conducted a meta-analysis and in-depth studies focused on the invasive grass species, Microstegium vimineum, to better understand how (i) plant characteristics, (ii) invader abundance and neighbor identity, and (iii) environmental conditions mediate the impacts of invasion on N pools and fluxes. The results of our global meta-analysis support the concept that invasive species and reference community traits such as leaf %N and leaf C:N are useful for understanding invasion impacts on soil N cycling, but that trait dissimilarities between invaded and reference communities are most informative. Regarding the in-depth studies of Microstegium, we did not find evidence to suggest that invasion increases net nitrification as other studies have shown. Instead, we found that an interaction between its abundance and the neighboring plant identify were important for determining soil nitrate concentrations and net nitrification rates in the greenhouse. In field, we found that variability in environmental conditions mediated the impact of Microstegium invasion on soil N pools and fluxes, primarily net ammonification, between sites through direct, indirect, and interactive pathways. Notably, we detected a scenario in which forest openness has a negative direct effect and indirect positive effect on ammonification in sites with high soil moisture and organic matter. Collectively, our findings suggest that dissimilarity in plant community traits, neighbor identity, and environmental conditions can be important drivers of invasion impacts on ecosystem N cycling and should be considered when evaluating the ecosystem impacts of invasive species across heterogeneous landscapes.

Ocean acidification slows nitrogen fixation and growth in the dominant diazotroph Trichodesmium under low-iron conditions

Dissolution of anthropogenic CO(2) increases the partial pressure of CO(2) (pCO(2)) and decreases the pH of seawater. The rate of Fe uptake by the dominant N(2)-fixing cyanobacterium Trichodesmium declines as pH decreases in metal-buffered medium. The slower Fe-uptake rate at low pH results from changes in Fe chemistry and not from a physiological response of the organism. Contrary to previous observations in nutrient-replete media, increasing pCO(2)/decreasing pH causes a decrease in the rates of N(2) fixation and growth in Trichodesmium under low-Fe conditions. This result was obtained even though the bioavailability of Fe was maintained at a constant level by increasing the total Fe concentration at low pH. Short-term experiments in which pCO(2) and pH were varied independently showed that the decrease in N(2) fixation is caused by decreasing pH rather than by increasing pCO(2) and corresponds to a lower efficiency of the nitrogenase enzyme. To compensate partially for the loss of N(2) fixation efficiency at low pH, Trichodesmium synthesizes additional nitrogenase. This increase comes partly at the cost of down-regulation of Fe-containing photosynthetic proteins. Our results show that although increasing pCO(2) often is beneficial to photosynthetic marine organisms, the concurrent decreasing pH can affect primary producers negatively. Such negative effects can occur both through chemical mechanisms, such as the bioavailability of key nutrients like Fe, and through biological mechanisms, as shown by the decrease in N(2) fixation in Fe-limited Trichodesmium.

Interactive effects of ocean acidification and nitrogen limitation on the diatom Phaeodactylum tricornutum

Climate change is expected to bring about alterations in the marine physical and chemical environment that will induce changes in the concentration of dissolved CO2 and in nutrient availability. These in turn are expected to affect the physiological performance of phytoplankton. In order to learn how phytoplankton respond to the predicted scenario of increased CO2 and decreased nitrogen in the surface mixed layer, we investigated the diatom Phaeodactylum tricornutum as a model organism. The cells were cultured in both low CO2 (390 µatm) and high CO2 (1000 µatm) conditions at limiting (10 µmol/L) or enriched (110 µmol/L) nitrate concentrations. Our study shows that nitrogen limitation resulted in significant decreases in cell size, pigmentation, growth rate and effective quantum yield of Phaeodactylum tricornutum, but these parameters were not affected by enhanced dissolved CO2 and lowered pH. However, increased CO2 concentration induced higher rETRmax and higher dark respiration rates and decreased the CO2 or dissolved inorganic carbon (DIC) affinity for electron transfer (shown by higher values for K1/2 DIC or K1/2 CO2). Furthermore, the elemental stoichiometry (carbon to nitrogen ratio) was raised under high CO2 conditions in both nitrogen limited and nitrogen replete conditions, with the ratio in the high CO2 and low nitrate grown cells being higher by 45% compared to that in the low CO2 and nitrate replete grown ones. Our results suggest that while nitrogen limitation had a greater effect than ocean acidification, the combined effects of both factors could act synergistically to affect marine diatoms and related biogeochemical cycles in future oceans.