Differential Response of High-Elevation Planktonic Bacterial Community Structure and Metabolism to Experimental Nutrient Enrichment

Nutrient enrichment of high-elevation freshwater ecosystems by atmospheric deposition is increasing worldwide, and bacteria are a key conduit for the metabolism of organic matter in these oligotrophic environments. We conducted two distinct in situ microcosm experiments in a high-elevation lake (Emerald Lake, Sierra Nevada, California, USA) to evaluate responses in bacterioplankton growth, carbon utilization, and community structure to short-term enrichment by nitrate and phosphate. The first experiment, conducted just following ice-off, employed dark dilution culture to directly assess the impact of nutrients on bacterioplankton growth and consumption of terrigenous dissolved organic matter during snowmelt. The second experiment, conducted in transparent microcosms during autumn overturn, examined how bacterioplankton in unmanipulated microbial communities responded to nutrients concomitant with increasing phytoplankton-derived organic matter. In both experiments, phosphate enrichment (but not nitrate) caused significant increases in bacterioplankton growth, changed particulate organic stoichiometry, and induced shifts in bacterial community composition, including consistent declines in the relative abundance of Actinobacteria. The dark dilution culture showed a significant increase in dissolved organic carbon removal in response to phosphate enrichment. In transparent microcosms nutrient enrichment had no effect on concentrations of chlorophyll, carbon, or the fluorescence characteristics of dissolved organic matter, suggesting that bacterioplankton responses were independent of phytoplankton responses. These results demonstrate that bacterioplankton communities in unproductive high-elevation habitats can rapidly alter their taxonomic composition and metabolism in response to short-term phosphate enrichment. Our results reinforce the key role that phosphorus plays in oligotrophic lake ecosystems, clarify the nature of bacterioplankton nutrient limitation, and emphasize that evaluation of eutrophication in these habitats should incorporate heterotrophic microbial communities and processes

Phosphorus in sediments of high-elevation lakes in the Sierra Nevada (California): implications for internal phosphorus loading

In high-elevation lakes of the Sierra Nevada (California), increases in phosphorus (P) supply have been inferred from changes in phytoplankton growth during summer. To quantify rates of sediment P release to high-elevation Sierran lakes, we performed incubations of sediment cores under ambient and reducing conditions at Emerald Lake and analyzed long-term records of lake chemistry for Emerald and Pear lakes. We also measured concentrations of individual P forms in sediments from 50 Sierra Nevada lakes using a sequential fractionation procedure to examine landscape controls on P forms in sediments. On average, the sediments contained 1,445 µg P g−1, of which 5 % was freely exchangeable, 13 % associated with reducible metal hydroxides, 68 % associated with Al hydroxides, and the remaining 14 % stabilized in recalcitrant pools. Multiple linear regression analysis indicated that sediment P fractions were not well correlated with soluble P concentrations. In general, sediments behaved as net sinks for P even under reducing conditions. Our findings suggest that internal P loading does not explain the increase in P availability observed in high-elevation Sierran lakes. Rather, increased atmospheric P inputs and increased P supply via dissolved organic C leaching from soils may be driving the observed changes in P biogeochemistry

Nutrients cause grassland biomass to outpace herbivory

Human activities are transforming grassland biomass via changing climate, elemental nutrients, and herbivory. Theory predicts that food-limited herbivores will consume any additional biomass stimulated by nutrient inputs ('consumer-controlled'). Alternatively, nutrient supply is predicted to increase biomass where herbivores alter community composition or are limited by factors other than food ('resource-controlled'). Using an experiment replicated in 58 grasslands spanning six continents, we show that nutrient addition and vertebrate herbivore exclusion each caused sustained increases in aboveground live biomass over a decade, but consumer control was weak. However, at sites with high vertebrate grazing intensity or domestic livestock, herbivores consumed the additional fertilization-induced biomass, supporting the consumer-controlled prediction. Herbivores most effectively reduced the additional live biomass at sites with low precipitation or high ambient soil nitrogen. Overall, these experimental results suggest that grassland biomass will outstrip wild herbivore control as human activities increase elemental nutrient supply, with widespread consequences for grazing and fire risk

Pools, transformations, and sources of P in high-elevation soils: Implications for nutrient transfer to Sierra Nevada lakes

In high-elevation lakes of the Sierra Nevada (California), increases in P supply have been inferred from shifts in P to N limitation. To examine factors possibly leading to changes in P supply, we measured pools and transformations in soil P, and developed a long-term mass balance to estimate the contribution of parent material weathering to soil P stocks. Common Sierra Nevada soils were found to not be P-deficient and to be retentive of P due to the influence of Fe- and Al-oxides. Total P averaged 867μgPg-1 in the top 10cm of soil (O and A horizons) and 597μgPg-1 in the 10-60cm depth (B horizons), of which 70% in A horizons and 60% in B horizons was freely exchangeable or associated with Fe and Al. Weathering of parent material explained 69% of the P found in soils and lost from the catchment since deglaciation, implying that long-term atmospheric P deposition (0.02kgha-1yr-1) represented the balance of P inputs (31%) during the past 10,000years of soil development. During spring snowmelt ~27% of the total soil P was transferred between organic and inorganic pools; average inorganic P pools decreased by 232μgPg-1, while organic P pools increased by 242μgPg-1. Microbial biomass P was highest during winter and decreased six-fold to a minimum in the fall. Interactions between hydrology and biological processes strongly influence the rate of P transfer from catchment soils to lakes. © 2013 Elsevier B.V

Pools, transformations, and sources of P in high-elevation soils: Implications for nutrient transfer to Sierra Nevada lakes

In high-elevation lakes of the Sierra Nevada (California), increases in P supply have been inferred from shifts in P to N limitation. To examine factors possibly leading to changes in P supply, we measured pools and transformations in soil P, and developed a long-term mass balance to estimate the contribution of parent material weathering to soil P stocks. Common Sierra Nevada soils were found to not be P-deficient and to be retentive of P due to the influence of Fe- and Al-oxides. Total P averaged 867μgPg-1 in the top 10cm of soil (O and A horizons) and 597μgPg-1 in the 10-60cm depth (B horizons), of which 70% in A horizons and 60% in B horizons was freely exchangeable or associated with Fe and Al. Weathering of parent material explained 69% of the P found in soils and lost from the catchment since deglaciation, implying that long-term atmospheric P deposition (0.02kgha-1yr-1) represented the balance of P inputs (31%) during the past 10,000years of soil development. During spring snowmelt ~27% of the total soil P was transferred between organic and inorganic pools; average inorganic P pools decreased by 232μgPg-1, while organic P pools increased by 242μgPg-1. Microbial biomass P was highest during winter and decreased six-fold to a minimum in the fall. Interactions between hydrology and biological processes strongly influence the rate of P transfer from catchment soils to lakes. © 2013 Elsevier B.V

Phosphorus in sediments of high-elevation lakes in the Sierra Nevada (California): implications for internal phosphorus loading

In high-elevation lakes of the Sierra Nevada (California), increases in phosphorus (P) supply have been inferred from changes in phytoplankton growth during summer. To quantify rates of sediment P release to high-elevation Sierran lakes, we performed incubations of sediment cores under ambient and reducing conditions at Emerald Lake and analyzed long-term records of lake chemistry for Emerald and Pear lakes. We also measured concentrations of individual P forms in sediments from 50 Sierra Nevada lakes using a sequential fractionation procedure to examine landscape controls on P forms in sediments. On average, the sediments contained 1,445 µg P g−1, of which 5 % was freely exchangeable, 13 % associated with reducible metal hydroxides, 68 % associated with Al hydroxides, and the remaining 14 % stabilized in recalcitrant pools. Multiple linear regression analysis indicated that sediment P fractions were not well correlated with soluble P concentrations. In general, sediments behaved as net sinks for P even under reducing conditions. Our findings suggest that internal P loading does not explain the increase in P availability observed in high-elevation Sierran lakes. Rather, increased atmospheric P inputs and increased P supply via dissolved organic C leaching from soils may be driving the observed changes in P biogeochemistry

A two-component hydrograph separation for three high-elevation catchments in the Sierra Nevada, California

Two-component hydrograph separations were performed for three, nested, snowmelt-dominated catchments in Sequoia National Park. The purpose of the hydrograph separations was to: (i) differentiate between the old and new water contributions to discharge during snowmelt using δ18O signatures; (ii) identify the fraction of snowmelt that travelled through the subsurface (reactive) compartment during the snowmelt period using silica or sodium; and (iii) investigate the impact of changing end-member signatures on the separations. 'Old' water refers to water that was stored in the watershed during the previous year, whereas 'new' water is current snowmelt. Hydrograph separations were performed for both a high-accumulation (1998, annual precipitation 2·4 m) and an average year (1999, 1·3 m). The proportion of old water contribution to discharge during the rising limb of the hydrograph was 10-20%, with 80-100% of snowmelt being reactive, i.e. passing through soil and talus. Estimates of old and new soil water and direct snowmelt entering the stream varied among the catchments in 1999. Differences between these components were minimal in 1998, regardless of varying topography and differing proportions of soil, rock and talus. Using time-dependent rather than constant δ18O meltwater and silica soil-water signatures made a meaningful impact on both new and old water, and reactive and unreactive, estimates. © 2004 John Wiley and Sons, Ltd