Phosphorus Interactions with Iron in Undisturbed and Disturbed Arctic Tundra Ecosystems.

Phosphorus (P) limitation often constrains biological processes in Arctic tundra ecosystems. Although adsorption to soil minerals may limit P bioavailability and export from soils into aquatic systems, the contribution of mineral phases to P retention in Arctic tundra is poorly understood. Our objective was to use X-ray absorption spectroscopy to characterize P speciation and associations with soil minerals along hillslope toposequences and in undisturbed and disturbed low-lying wet sedge tundra on the North Slope, AK. Biogenic mats comprised of short-range ordered iron (Fe) oxyhydroxides were prevalent in undisturbed wet sedge meadows. Upland soils and pond sediments impacted by gravel mining or thermokarst lacked biogenic Fe mats and were comparatively iron poor. Phosphorus was primarily contained in organic compounds in hillslope soils but associated with Fe(III) oxyhydroxides in undisturbed wet sedge meadows and calcium (Ca) in disturbed pond sediments. We infer that phosphate mobilized through organic decomposition binds to Fe(III) oxyhydroxides in wet sedge, but these associations are disrupted by physical disturbance that removes Fe mats. Increasing disturbances of the Arctic tundra may continue to alter the mineralogical composition of soils at terrestrial-aquatic interfaces and binding mechanisms that could inhibit or promote transport of bioavailable P from soils to aquatic ecosystems.

Bacterioplankton dispersal and biogeochemical function across Alaskan Arctic catchments.

In Arctic catchments, bacterioplankton are dispersed through soils and streams, both of which freeze and thaw/flow in phase, seasonally. To characterize this dispersal and its potential impact on biogeochemistry, we collected bacterioplankton and measured stream physicochemistry during snowmelt and after vegetation senescence across multiple stream orders in alpine, tundra, and tundra-dominated-by-lakes catchments. In all catchments, differences in community composition were associated with seasonal thaw, then attachment status (i.e. free floating or sediment associated), and then stream order. Bacterioplankton taxonomic diversity and richness were elevated in sediment-associated fractions and in higher-order reaches during snowmelt. Families Chthonomonadaceae, Pyrinomonadaceae, and Xiphinematobacteraceae were abundantly different across seasons, while Flavobacteriaceae and Microscillaceae were abundantly different between free-floating and sediment-associated fractions. Physicochemical data suggested there was high iron (Fe+ ) production (alpine catchment); Fe+ production and chloride (Cl- ) removal (tundra catchment); and phosphorus (SRP) removal and ammonium (NH4 + ) production (lake catchment). In tundra landscapes, these 'hot spots' of Fe+ production and Cl- removal accompanied shifts in species richness, while SRP promoted the antecedent community. Our findings suggest that freshet increases bacterial dispersal from headwater catchments to receiving catchments, where bacterioplankton-mineral relations stabilized communities in free-flowing reaches, but bacterioplankton-nutrient relations stabilized those punctuated by lakes.

Tundra wildfire triggers sustained lateral nutrient loss in Alaskan Arctic.

Climate change is creating widespread ecosystem disturbance across the permafrost zone, including a rapid increase in the extent and severity of tundra wildfire. The expansion of this previously rare disturbance has unknown consequences for lateral nutrient flux from terrestrial to aquatic environments. Lateral loss of nutrients could reduce carbon uptake and slow recovery of already nutrient-limited tundra ecosystems. To investigate the effects of tundra wildfire on lateral nutrient export, we analyzed water chemistry in and around the 10-year-old  Anaktuvuk River fire scar in northern Alaska. We collected water samples from 21 burned and 21 unburned watersheds during snowmelt, at peak growing season, and after plant senescence in 2017 and 2018. After a decade of ecosystem recovery, aboveground biomass had recovered in burned watersheds, but overall carbon and nitrogen remained ~20% lower, and the active layer remained ~10% deeper. Despite lower organic matter stocks, dissolved organic nutrients were substantially elevated in burned watersheds, with higher flow-weighted concentrations of organic carbon (25% higher), organic nitrogen (59% higher), organic phosphorus (65% higher), and organic sulfur (47% higher). Geochemical proxies indicated greater interaction with mineral soils in watersheds with surface subsidence, but optical analysis and isotopes suggested that recent plant growth, not mineral soil, was the main source of organic nutrients in burned watersheds. Burned and unburned watersheds had similar δ15 N-NO3 - , indicating that exported nitrogen was of preburn origin (i.e., not recently fixed). Lateral nitrogen flux from burned watersheds was 2- to 10-fold higher than rates of background nitrogen fixation and atmospheric deposition estimated in this area. These findings indicate that wildfire in Arctic tundra can destabilize nitrogen, phosphorus, and sulfur previously stored in permafrost via plant uptake and leaching. This plant-mediated nutrient loss could exacerbate terrestrial nutrient limitation after disturbance or serve as an important nutrient release mechanism during succession.

Cavity-enhanced non-destructive detection of atoms for an optical lattice clock.

We demonstrate a new method of cavity-enhanced non-destructive detection of atoms for a strontium optical lattice clock. The detection scheme is shown to be linear in atom number up to at least 2×104 atoms, to reject technical noise sources, to achieve signal to noise ratio close to the photon shot noise limit, to provide spatially uniform atom-cavity coupling, and to minimize inhomogeneous ac Stark shifts. These features enable detection of atoms with minimal perturbation to the atomic state, a critical step towards realizing an ultra-high-stability, quantum-enhanced optical lattice clock.

Linking permafrost thaw to shifting biogeochemistry and food web resources in an arctic river.

Rapidly, increasing air temperatures across the Arctic are thawing permafrost and exposing vast quantities of organic carbon, nitrogen, and phosphorus to microbial processing. Shifts in the absolute and relative supplies of these elements will likely alter patterns of ecosystem productivity and change the way carbon and nutrients are delivered from upland areas to surface waters such as rivers and lakes. The ultra-oligotrophic nature of surface waters across the Arctic renders these ecosystems particularly susceptible to changes in productivity and food web dynamics as permafrost thaw alters terrestrial-aquatic linkages. The objectives of this study were to evaluate decadal-scale patterns in surface water chemistry and assess potential implications of changing water chemistry to benthic organic matter and aquatic food webs. Data were collected from the upper Kuparuk River on the North Slope of Alaska by the U.S. National Science Foundation's Long-Term Ecological Research program during 1978-2014. Analyses of these data show increases in stream water alkalinity and cation concentrations consistent with signatures of permafrost thaw. Changes are also documented for discharge-corrected nitrate concentrations (+), discharge-corrected dissolved organic carbon concentrations (-), total phosphorus concentrations (-), and δ13 C isotope values of aquatic invertebrate consumers (-). These changes show that warming temperatures and thawing permafrost in the upland environment are leading to shifts in the supply of carbon and nutrients available to surface waters and consequently changing resources that support aquatic food webs. This demonstrates that physical, geochemical, and biological changes associated with warming permafrost are fundamentally altering linkages between upland and aquatic ecosystems in rapidly changing arctic environments.

Drivers of nitrogen transfer in stream food webs across continents.

Studies of trophic-level material and energy transfers are central to ecology. The use of isotopic tracers has now made it possible to measure trophic transfer efficiencies of important nutrients and to better understand how these materials move through food webs. We analyzed data from thirteen 15 N-ammonium tracer addition experiments to quantify N transfer from basal resources to animals in headwater streams with varying physical, chemical, and biological features. N transfer efficiencies from primary uptake compartments (PUCs; heterotrophic microorganisms and primary producers) to primary consumers was lower (mean 11.5%, range <1% to 43%) than N transfer efficiencies from primary consumers to predators (mean 80%, range 5% to >100%). Total N transferred (as a rate) was greater in streams with open compared to closed canopies and overall N transfer efficiency generally followed a similar pattern, although was not statistically significant. We used principal component analysis to condense a suite of site characteristics into two environmental components. Total N uptake rates among trophic levels were best predicted by the component that was correlated with latitude, DIN:SRP, GPP:ER, and percent canopy cover. N transfer efficiency did not respond consistently to environmental variables. Our results suggest that canopy cover influences N movement through stream food webs because light availability and primary production facilitate N transfer to higher trophic levels.

Effects of long-term nutrient additions on Arctic tundra, stream, and lake ecosystems: beyond NPP.

Primary producers form the base of food webs but also affect other ecosystem characteristics, such as habitat structure, light availability, and microclimate. Here, we examine changes caused by 5-30+ years of nutrient addition and resulting increases in net primary productivity (NPP) in tundra, streams, and lakes in northern Alaska. The Arctic provides an important opportunity to examine how ecosystems characterized by low diversity and low productivity respond to release from nutrient limitation. We review how responses of algae and plants affect light availability, perennial biotic structures available for consumers, oxygen levels, and temperature. Sometimes, responses were similar across all three ecosystems; e.g., increased NPP significantly reduced light to the substrate following fertilization. Perennial biotic structures increased in tundra and streams but not in lakes, and provided important new habitat niches for consumers as well as other producers. Oxygen and temperature responses also differed. Life history traits (e.g., longevity) of the primary producers along with the fate of detritus drove the responses and recovery. As global change persists and nutrients become more available in the Arctic and elsewhere, incorporating these factors as response variables will enable better prediction of ecosystem changes and feedbacks in this biome and others.

Microbial iron oxidation in the Arctic tundra and its implications for biogeochemical cycling.

The role that neutrophilic iron-oxidizing bacteria play in the Arctic tundra is unknown. This study surveyed chemosynthetic iron-oxidizing communities at the North Slope of Alaska near Toolik Field Station (TFS) at Toolik Lake (lat 68.63, long -149.60). Microbial iron mats were common in submerged habitats with stationary or slowly flowing water, and their greatest areal extent is in coating plant stems and sediments in wet sedge meadows. Some Fe-oxidizing bacteria (FeOB) produce easily recognized sheath or stalk morphotypes that were present and dominant in all the mats we observed. The cool water temperatures (9 to 11°C) and reduced pH (5.0 to 6.6) at all sites kinetically favor microbial iron oxidation. A microbial survey of five sites based on 16S rRNA genes found a predominance of Proteobacteria, with Betaproteobacteria and members of the family Comamonadaceae being the most prevalent operational taxonomic units (OTUs). In relative abundance, clades of lithotrophic FeOB composed 5 to 10% of the communities. OTUs related to cyanobacteria and chloroplasts accounted for 3 to 25% of the communities. Oxygen profiles showed evidence for oxygenic photosynthesis at the surface of some mats, indicating the coexistence of photosynthetic and FeOB populations. The relative abundance of OTUs belonging to putative Fe-reducing bacteria (FeRB) averaged around 11% in the sampled iron mats. Mats incubated anaerobically with 10 mM acetate rapidly initiated Fe reduction, indicating that active iron cycling is likely. The prevalence of iron mats on the tundra might impact the carbon cycle through lithoautotrophic chemosynthesis, anaerobic respiration of organic carbon coupled to iron reduction, and the suppression of methanogenesis, and it potentially influences phosphorus dynamics through the adsorption of phosphorus to iron oxides.

High-throughput analysis of single hematopoietic stem cell proliferation in microfluidic cell culture arrays.

Heterogeneity in cell populations poses a major obstacle to understanding complex biological processes. Here we present a microfluidic platform containing thousands of nanoliter-scale chambers suitable for live-cell imaging studies of clonal cultures of nonadherent cells with precise control of the conditions, capabilities for in situ immunostaining and recovery of viable cells. We show that this platform mimics conventional cultures in reproducing the responses of various types of primitive mouse hematopoietic cells with retention of their functional properties, as demonstrated by subsequent in vitro and in vivo (transplantation) assays of recovered cells. The automated medium exchange of this system made it possible to define when Steel factor stimulation is first required by adult hematopoietic stem cells in vitro as the point of exit from quiescence. This technology will offer many new avenues to interrogate otherwise inaccessible mechanisms governing mammalian cell growth and fate decisions.

Anomalous behavior of dark states in quantum gases of (6)Li.

We create atom-molecule dark states in a degenerate Fermi gas of ^{6}Li in both weakly and strongly interacting regimes using two-photon Raman scattering to couple fermion pairs to bound molecular states in the ground singlet and triplet potential. Near the unitarity point in the BEC-BCS crossover regime, the atom number revival height associated with the dark state abruptly and unexpectedly decreases and remains low for magnetic fields below the Feshbach resonance center at 832.2 G. With a weakly interacting Fermi gas at 0 G, we perform precision dark-state spectroscopy of the least-bound vibrational levels of the lowest singlet and triplet potentials. From these spectra, we obtain binding energies of the v^{''}=9, N^{''}=0 level of the a(1^{3}Σ_{u}^{+}) potential and the v^{''}=38, N^{''}=0 level of the X(1^{1}Σ_{g}^{+}) potential with absolute uncertainty as low as 20 kHz. For the triplet potential, we resolve the molecular hyperfine structure.

Superlinear scaling of riverine biogeochemical function with watershed size.

River networks regulate carbon and nutrient exchange between continents, atmosphere, and oceans. However, contributions of riverine processing are poorly constrained at continental scales. Scaling relationships of cumulative biogeochemical function with watershed size (allometric scaling) provide an approach for quantifying the contributions of fluvial networks in the Earth system. Here we show that allometric scaling of cumulative riverine function with watershed area ranges from linear to superlinear, with scaling exponents constrained by network shape, hydrological conditions, and biogeochemical process rates. Allometric scaling is superlinear for processes that are largely independent of substrate concentration (e.g., gross primary production) due to superlinear scaling of river network surface area with watershed area. Allometric scaling for typically substrate-limited processes (e.g., denitrification) is linear in river networks with high biogeochemical activity or low river discharge but becomes increasingly superlinear under lower biogeochemical activity or high discharge, conditions that are widely prevalent in river networks. The frequent occurrence of superlinear scaling indicates that biogeochemical activity in large rivers contributes disproportionately to the function of river networks in the Earth system.

A scalable arbitrary waveform generator for atomic physics experiments based on field-programmable gate array technology.

We present a field-programmable gate array (FPGA) based control system that has been implemented to control a strontium optical lattice clock at the National Physical Laboratory, UK. Bespoke printed circuit boards have been designed and manufactured, including an 8-channel, 16-bit digital to analog converter board with a 2 µs update rate and a 4-channel direct-digital synthesis board clocked at 1 GHz. Each board includes its own FPGA with 28 digital output lines available alongside the specialized analog or radio frequency outputs. The system is scalable to a large number of control lines by stacking the individual boards in a master-slave arrangement. The timing of the digital and analog outputs is based on the FPGA clock and is thus very predictable and exhibits low jitter. A particular advantage of our hardware is its large data buffers that, when combined with a pseudoclock structure, allow complex waveforms to be created. A high reliability of the system has been demonstrated during extended atomic clock frequency comparisons.

Revealing biogeochemical signatures of Arctic landscapes with river chemistry.

Riverine fluxes of carbon and inorganic nutrients are increasing in virtually all large permafrost-affected rivers, indicating major shifts in Arctic landscapes. However, it is currently difficult to identify what is causing these changes in nutrient processing and flux because most long-term records of Arctic river chemistry are from small, headwater catchments draining <200 km2 or from large rivers draining >100,000 km2. The interactions of nutrient sources and sinks across these scales are what ultimately control solute flux to the Arctic Ocean. In this context, we performed spatially-distributed sampling of 120 subcatchments nested within three Arctic watersheds spanning alpine, tundra, and glacial-lake landscapes in Alaska. We found that the dominant spatial scales controlling organic carbon and major nutrient concentrations was 3-30 km2, indicating a continuum of diffuse and discrete sourcing and processing dynamics. These patterns were consistent seasonally, suggesting that relatively fine-scale landscape patches drive solute generation in this region of the Arctic. These network-scale empirical frameworks could guide and benchmark future Earth system models seeking to represent lateral and longitudinal solute transport in rapidly changing Arctic landscapes.

A pyramid MOT with integrated optical cavities as a cold atom platform for an optical lattice clock.

We realize a two-stage, hexagonal pyramid magneto-optical trap (MOT) with strontium, and demonstrate loading of cold atoms into cavity-enhanced 1D and 2D optical lattice traps, all within a single compact assembly of in-vacuum optics. We show that the device is suitable for high-performance quantum technologies, focusing especially on its intended application as a strontium optical lattice clock. We prepare 2 × 104 spin-polarized atoms of 87Sr in the optical lattice within 500 ms; we observe a vacuum-limited lifetime of atoms in the lattice of 27 s; and we measure a background DC electric field of 12 V m-1 from stray charges, corresponding to a fractional frequency shift of (-1.2 ± 0.8) × 10-18 to the strontium clock transition. When used in combination with careful management of the blackbody radiation environment, the device shows potential as a platform for realizing a compact, robust, transportable optical lattice clock with systematic uncertainty at the 10-18 level.

An adaptable dual species effusive source and Zeeman slower design demonstrated with Rb and Li.

We present a dual-species effusive source and Zeeman slower designed to produce slow atomic beams of two elements with a large mass difference and with very different oven temperature requirements. We demonstrate this design for the case of (6)Li and (85)Rb and achieve magneto-optical trap (MOT) loading rates equivalent to that reported in prior work on dual species (Rb+Li) Zeeman slowers operating at the same oven temperatures. Key design choices, including thermally separating the effusive sources and using a segmented coil design to enable computer control of the magnetic field profile, ensure that the apparatus can be easily modified to slow other atomic species. By performing the final slowing using the quadrupole magnetic field of the MOT, we are able to shorten our Zeeman slower length making for a more compact system without compromising performance. We outline the construction and analyze the emission properties of our effusive sources. We also verify the performance of the source and slower, and we observe sequential loading rates of 12 × 10(8) atoms/s for a Rb oven temperature of 140 °C and 1.1 × 10(8) atoms/s for a Li reservoir at 460 °C, corresponding to reservoir lifetimes for continuous operation of 10 and 4 years, respectively.

Microbial biogeography of arctic streams: exploring influences of lithology and habitat.

Terminal restriction fragment length polymorphism and 16S rRNA gene sequencing were used to explore the community composition of bacterial communities in biofilms on sediments (epipssamon) and rocks (epilithon) in stream reaches that drain watersheds with contrasting lithologies in the Noatak National Preserve, Alaska. Bacterial community composition varied primarily by stream habitat and secondarily by lithology. Positive correlations were detected between bacterial community structure and nutrients, base cations, and dissolved organic carbon. Our results showed significant differences at the stream habitat, between epipssamon and epilithon bacterial communities, which we expected. Our results also showed significant differences at the landscape scale that could be related to different lithologies and associated stream biogeochemistry. These results provide insight into the bacterial community composition of little known and pristine arctic stream ecosystems and illustrate how differences in the lithology, soils, and vegetation community of the terrestrial environment interact to influence stream bacterial taxonomic richness and composition.

Carbon and nitrogen stoichiometry and nitrogen cycling rates in streams.

Stoichiometric analyses can be used to investigate the linkages between N and C cycles and how these linkages influence biogeochemistry at many scales, from components of individual ecosystems up to the biosphere. N-specific NH4+ uptake rates were measured in eight streams using short-term 15N tracer additions, and C to N ratios (C:N) were determined from living and non-living organic matter collected from ten streams. These data were also compared to previously published data compiled from studies of lakes, ponds, wetlands, forests, and tundra. There was a significant negative relationship between C:N and N-specific uptake rate; C:N could account for 41% of the variance in N-specific uptake rate across all streams, and the relationship held in five of eight streams. Most of the variation in N-specific uptake rate was contributed by detrital and primary producer compartments with large values of C:N and small values for N-specific uptake rate. In streams, particulate materials are not as likely to move downstream as dissolved N, so if N is cycling in a particulate compartment, N retention is likely to be greater. Together, these data suggest that N retention may depend in part on C:N of living and non-living organic matter in streams. Factors that alter C:N of stream ecosystem compartments, such as removal of riparian vegetation or N fertilization, may influence the amount of retention attributed to these ecosystem compartments by causing shifts in stoichiometry. Our analysis suggests that C:N of ecosystem compartments can be used to link N-cycling models across streams.