Relationships between soil microbial properties and aboveground stand characteristics of conifer forests in Oregon

Eight forest sites representing a large range of climate, vegetation, and productivity were sampled in a transect across Oregon to study the relationships between aboveground stand characteristics and soil microbial properties. These sites had a range in leaf area index of 0.6 to 16 m² m^–2 and net primary productivity of 0.3 to 14 Mg ha^–1 yr^–1.Measurements of soil and forest floor inorganic N concentrations and in situ net N mineralization, nitrification, denitrification, and soil respiration were made monthly for one year. Microbial biomass C and anaerobic N mineralization, an index of N availability, were also measured. Annual mean concentrations of NH ₄ ⁺ ranged from 37 to 96 mg N kg^–1 in the forest floor and from 1.7 to 10.7 mg N kg^–1 in the mineral soil. Concentrations of NO ₃ ^– were low ( < 1 mg N kg^–1) at all sites. Net N mineralization and nitrification, as measured by the buried bag technique, were low on most sites and denitrification was not detected at any site. Available N varied from 17 to 101 mg N kg^–1, microbial biomass C ranged from 190 to 1230 mg Ckg^–1, and soil respiration rates varied from 1.3 to 49 mg C kg^–1 day^–1 across these sites. Seasonal peaks in NH ₄ ⁺ concentrations and soil respiration rates were usually observed in the spring and fall.The soils data were positively correlated with several aboveground variables, including leaf area index and net primary productivity, and the near infrared-to-red reflectance ratio obtained from the airborne simulator of the Thematic Mapper satellite. The data suggest that close relationships between aboveground productivity and soil microbial processes exist in forests approaching semi-equilibrium conditions.Abbreviations IR infrared - LAI leaf area index - k _c proportion of microbial biomass C mineralized to CO₂ - NPP net primary productivity - TM Thematic Mapper     

Measured and modelled effects of temperature,dissolved oxygen and nutrient concentration on sediment-water nutrient exchange

Empirical models of sediment-water fluxes of NH₄ ⁺, NO₃ ^– were and PO₄ ^3– were formed based on published reports. The models were revised and parameters evaluated based on laboratory incubations of sediments collected from Gunston Cove, VA. Observed fluxes ranged from — 18 (sediments uptake) to 276 (sediment release) mg NH₄ ⁺ m^–2 day^–1, –17 to –509 mg NO₃ ^– m^–2 day^–1, and –16.4 to 8.9 mg PO₄ ^3– m^–2 day^–1. The model and observations indicated release of NH₄ ⁺ was enhanced by high temperature and by low DO. Uptake of NO₃ ^– was enhanced primarily by high NO₃ ^– concentration and to a lesser extent by high temperature and by low DO. Direction of PO₄ ^3– flux depended on concentration in the water. Release was enhanced by low DO. No effect of temperature on PO₄ ^3– flux was observed.     

Inorganic nitrogen and microbial biomass dynamics before and during spring snowmelt

Recent work in seasonally snow covered ecosystems has identifiedthawed soil and high levels of heterotrophic activity throughout the winterunder consistent snow cover. We performed measurements during the winter of1994 to determine how the depth and timing of seasonal snow cover affectsoil microbial populations, surface water NO loss during snowmelt, and plant Navailability early in the growing season. Soil under early accumulating,consistent snow cover remained thawed during most of the winter and bothmicrobial biomass and soil inorganic N pools gradually increased under thesnowpack. At the initiation of snowmelt, microbial biomass N pools increasedfrom 3.0 to 5.9 g n m^-2,concurrent with a decrease in soil inorganic N pools. During the latterstages of snowmelt, microbial biomass N pools decreased sharply without aconcurrent increase in inorganic N pools or significant leaching losses. Incontrast, soil under inconsistent snow cover remained frozen during most ofthe winter. During snowmelt, microbial biomass initially increased from 1.7to 3.1 g N m^-2 and thendecreased as sites became snow-free. In contrast to smaller pool sizes,NO export during snowmeltfrom the inconsistent snow cover sites of 1.14 (±0.511) g N m^-2 was significantly greater (p< 0.001) than the 0.27 (±0.16) g N m^-2 exported from sites with consistent snowcover. These data suggest that microbial biomass in consistentlysnow-covered soil provides a significant buffer limiting the export ofinorganic N to surface water during snowmelt. However, this buffer is verysensitive to changes in snowpack regime. Therefore, interannual variabilityin the timing and depth of snowpack accumulation may explain the year toyear variability in inorganic N concentrations in surface water theseecosystems.     

Removal of municipal solid waste COD and NH4–N by phyto-reduction: A laboratory–scale comparison of terrestrial and aquatic species at different organic loads

Laboratory scale tests on phytodepuration of raw and pre-treated leachate from municipal sanitary waste were carried out with four vegetable aquatic and terrestrial species at different organic loads. We used the terrestrial species Stenotaphrum secundatum and the free-floating aquatic species Lemna minor, Eichhornia crassipes and Myriophyllum verticellatum to purify leachate from municipal solid waste. The organic load characterized by COD varied from 2–30 g m⁻² day⁻¹. Blanks using tap water served as controls. Duration of the experiments varied from 9–90 days. Maximum concentrations in the experiments were 1600 mg l⁻¹ COD and 300 mg l⁻¹ NH₄–N for S. secundatum. Best results in terms of COD, BOD, and ammonia removal were obtained for raw leachate with COD=2 g m⁻² day⁻¹ in free water surface (FWS) wetlands, and with 2 and 5 g m⁻² day⁻¹ in subsurface flow (SSF) wetlands. Results show that for pretreated leachate (labeled c) low in BOD and NH₄–N, the aquatic species showed low removal and stress even at the lowest load of COD=2 g m⁻² day⁻¹. We cannot say if this is due to the pretreatment itself or the chemical or microbial composition of this leachate. The Stenotaphrum system operated well with this load of leachate c. For untreated leachate (type a and b) the removal and plant growing conditions seemed good at COD=2 g m⁻² day⁻¹. For S. secundatum a load of COD=5 g m⁻² day⁻¹ operated well. All loads above COD=5 g m⁻² day⁻¹ caused low removal and stress, and the green parts of the plants disappeared. Oxygen was, however, consumed throughout the experimental period. For pretreated leachate (type c), the removal of COD were low (−24 to 17%) but good for NH₄–N (52–91%). This leachate also experienced high ammonia removal from the beginning of the experiments, probably due to existing consortia of nitrifying bacteria in it. Statistical analysis shows that the S. secundatum and L. minor systems maintained higher oxygen levels than the M. verticellatum and E. crassipes systems, when operated with tap water. For Lemna minor, this may be due to a better capacity for transporting oxygen into the water. With leachate all S. secundatum systems have higher oxygen levels than the aquatic systems, basically because the water content of the soil has been kept well below saturation. S. secundatum shows a significantly lower removal of COD than did the aquatic systems at a loading of COD=2 g m⁻² day⁻¹ of raw leachate. There is no significant difference between the systems in the removal of NH₄–N at a loading of COD=2 g m⁻² day⁻¹ of both types of leachate. E. crassipes has a lower removal of NH₄–N than M. verticellatum and S. secundatum at a loading of 5 g m⁻² day⁻¹ of COD of both types of leachate. In our experiments, it appears that the amount of free ammonia explains the toxicity of the leachate to the plants. This, however, does not exclude other possible toxic factors.     

Microbial biomass and nitrogen cycling responses to fertilization and litter removal in young northern hardwood forests

The influence of site fertility on soil microbial biomass and activity is not well understood but is likely to be complex because of interactions with plant responses to nutrient availability. We examined the effects of long-term (8 yr) fertilization and litter removal on forest floor microbial biomass and N and C transformations to test the hypothesis that higher soil resource availability stimulates microbial activity. Microbial biomass and respiration decreased by 20–30 % in response to fertilization. Microbial C averaged 3.8 mg C/g soil in fertilized, 5.8 mg C/g in control, and 5.5 mg C/g in litter removal plots. Microbial respiration was 200 µg CO₂-C g^–1 d^–1 in fertilized plots, compared to 270 µg CO₂-C g^–1 d^–1 in controls. Gross N mineralization and N immobilization did not differ among treatments, despite higher litter nutrient concentrations in fertilized plots and the removal of substantial quantities of C and N in litter removal plots. Net N mineralization was significantly reduced by fertilization. Gross nitrification and NO₃ ^– immobilization both were increased by fertilization. Nitrate thus became a more important part of microbial N cycling in fertilized plots even though NH₄ ⁺ availability was not stimulated by fertilization.Soil microorganisms did not mineralize more C or N in response to fertilization and higher litter quality; instead, results suggest a difference in the physiological status of microbial biomass in fertilized plots that influenced N transformations. Respiration quotients (qCO₂, respiration per unit biomass) were higher in fertilized plots (56 µg CO₂-C mg C^–1 d^–1) than control (48 µg CO₂-C mg C^–1 d ^–1) or litter removal (45 µg CO₂-C mg C^–1 d^–1), corresponding to higher microbial growth efficiency, higher proportions of gross mineralization immobilized, and lower net N mineralization in fertilized plots. While microbial biomass is an important labile nutrient pool, patterns of microbial growth and turnover were distinct from this pool and were more important to microbial function in nitrogen cycling.     

Winter microbial activity in Lake Tuusulanjärvi

Microbial heterotrophic activity, dark CO₂ assimilation, primary productivity and microbial ATP were measured monthly in the extremely eutrophic Lake Tuusulanjärvi during the winter of 1979–1980. Because of continuous water circulation caused by low temperature and artificial aeration of the lake, no winter stratification developed. Very low summertime ³H-glucose turnover times of 5 h increased to a level of 10–20 h from August to January. Winter maximum of 110 h was measured in March, and turnover times returned to 10–20 h in April, before the vernal bloom of algae occured. Oxygen saturation remained over 46% during the winter.High primary productivity was observed in November (400–500 mg C m^–3 day^–1), and measurable productivity was detected under ice in January (80 mg C m^–3 day^–1). Dark CO₂ assimilation increased to 14% of primary productivity in March. No correlation was found between ³H-glucose turnover rate and dark CO₂ assimilation. ATP correlated slightly better with primary productivity than with turnover rate. The single concentration method proved to be sensitive for winter heterotrophic activity measurement.     

Effects of mild winter freezing on soil nitrogen and carbon dynamics in a northern hardwood forest

Overwinter and snowmelt processes are thought to be critical to controllersof nitrogen (N) cycling and retention in northern forests. However, therehave been few measurements of basic N cycle processes (e.g.mineralization, nitrification, denitrification) during winter and littleanalysis of the influence of winter climate on growing season N dynamics.In this study, we manipulated snow cover to assess the effects of soilfreezing on in situ rates of N mineralization, nitrification and soilrespiration, denitrification (intact core, C₂H₂ – based method),microbial biomass C and N content and potential net N mineralization andnitrification in two sugar maple and two yellow birch stands with referenceand snow manipulation treatment plots over a two year period at theHubbard Brook Experimental Forest, New Hampshire, U.S.A. The snowmanipulation treatment, which simulated the late development of snowpackas may occur in a warmer climate, induced mild (temperatures >–5 °C) soil freezing that lasted until snowmelt. The treatmentcaused significant increases in soil nitrate (NO₃ ^–)concentrations in sugar maple stands, but did not affect mineralization,nitrification, denitrification or microbial biomass, and had no significanteffects in yellow birch stands. Annual N mineralization and nitrificationrates varied significantly from year to year. Net mineralization increasedfrom 12.0 g N m^–2 y^–1 in 1998 to 22 g N m^–2 y^–1 in 1999 and nitrification increased from 8 g N m^–2 y^–1 in 1998 to 13 g N m^–2 y^–1 in 1999.Denitrification rates ranged from 0 to 0.65 g N m^–2 y^–1. Ourresults suggest that mild soil freezing must increase soil NO₃ ^– levels by physical disruption of the soil ecosystem and not by direct stimulation of mineralization and nitrification. Physical disruption canincrease fine root mortality, reduce plant N uptake and reduce competitionfor inorganic N, allowing soil NO₃ ^– levels to increase evenwith no increase in net mineralization or nitrification.     

Interactions between sediment and water in a shallow and hypertrophic lake: a study on phytoplankton collapses in Lake Søbygård,Denmark

Short-term changes in phytoplankton and zooplankton biomass have occurred 1–3 times every summer for the past 5 years in the shallow and hypertrophic Lake Søbygård, Denmark. These changes markedly affected lake water characteristics as well as the sediment/water interaction. Thus during a collapse of the phytoplankton biomass in 1985, lasting for about 2 weeks, the lake water became almost anoxic, followed by rapid increase in nitrogen and phosphorus at rates of 100–400 mg N M^–2 day^–1 and 100–200 mg P m^–1 day^–1. Average external loading during this period was about 350 mg N m^–2 day^–1 and 5 mg P m^–2 day^–1, respectively.Due to high phytoplankton biomass and subsequently a high sedimentation and recycling of nutrients, gross release rates of phosphorus and nitrogen were several times higher than net release rates. The net summer sediment release of phosphorus was usually about 40 mg P m^–2 day^–1, corresponding to a 2–3 fold increase in the net phosphorus release during the collapse. The nitrogen and phosphorus increase during the collapse is considered to be due primarily to a decreased sedimentation because of low algal biomass. The nutrient interactions between sediment and lake water during phytoplankton collapse, therefore, were changed from being dominated by both a large input and a large sedimentation of nutrients to a dominance of only a large input. Nitrogen was derived from both the inlet and sediment, whereas phosphorus was preferentially derived from the sediment. Different temperature levels may be a main reason for the different release rates from year to year.     

Winter fluxes of methane from Minnesota peatlands

Winter fluxes of methane were investigated in northern Minnesota during 1988–89 and 1989–90. Two bogs and a fen emitted methane throughout the snow-covered season (November through March). Fluxes decreased to a low level of 3–16 mg CH₄ m^–2 d^–1 in late March, reflecting decreasing peat temperatures and (in 1989–90) increasing depth of frost in the peat. Winter fluxes calculated by integration for an open poor fen, an open bog, a forested bog hollow, and a hummock site in the forested bog averaged 49, 12, 13, and 5 mg m^–2 d^–1, respectively, in 1989–1990 (the year most measurements were made). These comprised 11%, 4%, 15%, and 21% of total annual flux.     

Nitrogen dynamics in a eutrophic lake sediment

Nitrogen flux from sediment of a shallow lake and subsequent utilization by water hyacinth (Eichhornia crassipes [Mart] Solms) present in the water column were evaluated using an outdoor microcosm sediment-water column. Sediment N was enriched with ¹⁵N to quantitatively determine the movement of NH₄-N from the sediment to the overlying water column. During the first 30 days. 48% of the total N uptake by water hyacinth was derived from sediment ¹⁵NH₄-N. This had decreased to 14% after 183 days. Mass balance of N indicates that about 25% sediment NH₄-N was released into the overlying water, but only 17% was assimilated by water hyacinth. NH₄-N levels in the water column were very low, with very little or no concentration gradients. NH₄-N levels in the interstitial water of the sediment were in the range of 30–35 mg L^–1 for the lower depths (> 35 cm), while in the surface 5 cm of depth NH₄-N levels decreased to 3.2 mg L^–1. Simulated results also showed similar trends for the interstitial NH₄-N concentration of the sediment. The overall estimated NH₄-N flux from the sediment to the overlying water was 4.8 µg cm^–2 day^–1, and the soluble organic N flux was 5.8 µg N cm^–2 day^–1. Total N flux was 10.6 µg N cm^–2 day^–1.     

Winter production of CO2 and N2O from alpine tundra: environmental controls and relationship to inter-system C and N fluxes

Fluxes of CO₂ and N₂O were measured from both natural and experimentally augmented snowpacks during the winters of 1993 and 1994 on Niwot Ridge in the Colorado Front Range. Consistent snow cover insulated the soil surface from extreme air temperatures and allowed heterotrophic activity to continue through much of the winter. In contrast, soil remained frozen at sites with inconsistent snow cover and production did not begin until snowmelt. Fluxes were measured when soil temperatures under the snow ranged from –5°C to 0°C, but there was no significant relationship between flux for either gas and temperature within this range. While early developing snowpacks resulted in warmer minimum soil temperatures allowing production to continue for most of the winter, the highest CO₂ fluxes were recorded at sites which experienced a hard freeze before a consistent snowpack developed. Consequently, the seasonal flux of CO₂ –C from snow covered soils was related both to the severity of freeze and the duration of snow cover. Over-winter CO₂ –C loss ranged from 0.3 g C m⁻² season⁻¹ at sites characterized by inconsistent snow cover to 25.7 g C m⁻² season⁻¹ at sites that experienced a hard freeze followed by an extended period of snow cover. In contrast to the pattern observed with C loss, a hard freeze early in the winter did not result in greater N₂O–N loss. Both mean daily N₂O fluxes and the total over-winter N₂O–N loss were related to the length of time soils were covered by a consistent snowpack. Over-winter N₂O–N loss ranged from less 0.23 mg N m⁻² from the latest developing, short duration snowpacks to 16.90 mg N m⁻² from sites with early snow cover. These data suggest that over-winter heterotrophic activity in snow-covered soil has the potential to mineralize from less than 1% to greater than 25% of the carbon fixed in ANPP, while over-winter N₂O fluxes range from less than half to an order of magnitude higher than growing season fluxes. The variability in these fluxes suggests that small changes in climate which affect the timing of seasonal snow cover may have a large effect on C and N cycling in these environments. Received: 5 April 1996 / Accepted: 25 November 1996     

Impact of NH4NO3 on microbial biomass C and N and extractable DOM in raised bog peat beneath Sphagnum capillifolium and S. recurvum

Regular bi-weekly additions of NH₄NO₃, equivalent to a rate of 3 g N m^–2 yr^–1, were applied to cores of Sphagnum capillifolium, inhabiting hummocks and S. recurvum a pool and hollow colonizer, in a raisedbog in north east Scotland. Microbial biomass C and N,both measured by chloroform extraction, showed similarseasonal patterns and, for most depths, the effects ofadded N on microbial biomass C and N changed withtime. The addition of inorganic N had greatest effectduring October when the water table had risen to thesurface and microbial C and N in the untreated coreshad decreased. Microbial C and N were maintained at75 g C m^–2 and 8.3 g N m^–2 above the values in the untreated cores and far exceeded the amounts of N that had been added up to that date (1 g N m^–2) as NH₄NO₃. This increased microbial biomass was interpreted as leaching of carbonaceous material from the NH₄NO₃ treated moss resulting in greater resistance of the microbialbiomass to changes induced by the rising water table.Treatment with N also caused significant reductions inextractable dissolved organic N (DON) at 10–15 cmdepth, beneath the surface of the moss, but at lowerdepths to 25 cm no changes were observed. Extracteddissolved organic carbon (DOC) was not affected by Ntreatment and showed less seasonal variation than DON,such that the C:N ratio of dissolved organic matter(DOM) in all depths increased from approximately 4 inJuly to around 30 in December.     

A comparison of oxygen,nitrate, and sulfate respiration in coastal marine sediments

Aerobic respiration with oxygen and anaerobic respiration with nitrate (denitrification) and sulfate (sulfate reduction) were measured during winter and summer in two coastal marine sediments (Denmark). Both aerobic respiration and denitrification took place in the oxidized surface layer, whereas sulfate reduction was most significant in the deeper, reduced sediment. The low availability of nitrate apparently limited the activity of denitrification during summer to less than 0.2 mmoles NO ₃ ^– m^–2 day^–1, whereas activities of 1.0–3.0 mmoles NO ₃ ^– m^–2 day^–1 were measured during winter. Sulfate reduction, on the contrary, increased from 2.6–7.6 mmoles SO ₄ ^2– m^–2 day^–1 during winter to 9.8–15.1 mmoles SO ₄ ^2– m^–2 day^–1 during summer. The aerobic respiration was high during summer, 135–140 mmoles O₂ m^–2 day^–1, as compared to estimated winter activities of about 30 mmoles O₂ m^–2 day^–1. The little importance of denitrification relative to aerobic respiration and sulfate reduction is discussed in relation to the availability and distribution of oxygen, nitrate, and sulfate in the sediments and to the detritus mineralization.     

Nitrogen dynamics in two antarctic streams

The many glacier meltwater streams of southern Victoria Land flow through catchments where life forms are almost entirely microbial. Allochthonous inputs of nitrogen from two study streams near McMurdo Sound were derived mostly from the melting glaciers (ca. 100–200 mg N m^–3) with some originating from N₂-fixation by heterocystous cyanobacteria (max. 939 mg N m^–2 year^–1). Thirty to fifty per cent of the glacier derived N was dissolved organic N and a major proportion of this was identified as urea N which was utilised by the rich algal and cyanobacterial mats in the streams. A nutrient budget for Fryxell Stream was estimated, quantifying uptake of urea-N and dissolved inorganic N and the release of dissolved organic (non urea) and particulate N by the stream communities. An index of in-stream nitrogen processing, the Net Uptake Length Constant in these streams was compared with that from temperate climates and was found to be similar. Despite the influence of low temperatures on microbial activity (mean daily water temperature = 5 °C) nutrient removal rates from these antarctic streams are high because of the large standing stock of microbial biomass there.     

Comparison of soil nitrogen mineralization and nitrification in a mixed grassland and forested ecosystem in central Taiwan

We used the buried bag incubation method to study temporal patterns of net N mineralization and net nitrification in soils at Ta-Ta-Chia forest in central Taiwan. The site included a grassland zone, (dominant vegetation consists of Yushania niitakayamensis and Miscanthus transmorrisonensis Hayata) and a forest zone (Tsuga chinensis var. formosana and Yushania niitakamensis). In the grassland, soil concentration NH₄ ⁺ in the organic horizon (0.1–0.2 m) ranged from 1.0 to 12.4 mg N kg^–1 soil and that of NO₃ ^– varied from 0.2 to 2.1 mg N kg^–1 soil. In the forest zone, NH₄ ⁺ concentration was between 2.8 and 25.0 mg N kg^–1 soil and NO₃ ^–varied from 0.2 to 1.3 mg N kg^–1 soil. There were lower soil NH₄ ⁺ concentrations during the summer than other seasons. Net N mineralization was higher during the summer while net nitrification rates did not show a distinct seasonal pattern. In the grassland, net N mineralization and net nitrification rates were between –0.1 and 0.24 and from –0.04 to 0.04 mg N kg^–1 soil day^–1, respectively. In the forest zone, net N mineralization rates were between –0.03 and 0.45 mg N kg^–1 soil day^–1 and net nitrification rates were between –0.01 and 0.03 mg N kg^–1 soil day^–1. These differences likely result from differing vegetation communities (C₃ versus C₄ plant type) and soil characteristics.     

Concentration and flux of solutes from snow and forest floor during snowmelt in the West-Central Adirondack region of New York

Decreases in pH and increases in the concentration of Al and NO ₃ ^– have been observed in surface waters draining acid-sensitive regions in the northeastern U.S. during spring snowmelt. To assess the source of this acidity, we evaluated solute concentrations in snowpack, and in meltwater collected from snow and forest floor lysimeters in the west-central Adirondack Mountains of New York during the spring snowmelt period, 29 March through 15 April 1984.During the initial phase of snowmelt, ions were preferentially leached from the snowpack resulting in elevated concentrations in snowmelt water (e.g. H⁺ = 140 eq.l^–1; NO ₄ ^2– = 123 eq.l^–1; SO ₃ ^– = 160 eq.l^–1). Solute concentrations decreased dramatically within a few days of the initial melt (< 50 eq.l^–1). The concentrations of SO ₄ ^2– and NO ₃ ^– in snowpack and snowmelt water were similar, whereas NO^– ₃ in the forest floor leachate was at least two times the concentration of SO ₄ ^2– .Study results suggest that the forest floor was a sink for snowmelt inputs of alkalinity, and a net source of H⁺, NO ₃ ^– , dissolved organic carbon, K⁺ and Al inputs to the mineral soil. The forest floor was relatively conservative with respect to snowmelt inputs of Ca²⁺, SO ₄ ^2– and Cl^–. These results indicate that mineralization of N, followed by nitrification in the forest floor may be an important process contributing to elevated concentrations of H⁺ and NO ₃ ^– in streams during the snowmelt period.     

Primary production by benthic microalgae in nearshore marine sediments of Signy Island,Antarctica

Summary During the austral summer of 1987/1988, three 24 h in situ primary productivity measurements were made at a nearshore sublittoral site on the east coast of Signy Island, Antarctica. The first experiment in December, coincided with the peak of the benthic algal bloom as shown by benthic chlorophyll measurements and a primary productivity rate of 700.9 mg carbon m^–2 day^–1. In January, the experiment was undertaken during the peak of the phytoplankton bloom when light intensities reaching the benthos were greatly reduced. A rate of 313.4 mg carbon m^–2 day^–1 was measured, half that of the previous month. In March the phytoplankton bloom had died off, benthic light intensities had increased and production was 391.8 mg m^–2 day^–1. The experiments indicate changes in benthic microalgal activity during the summer, linked to changes in the benthic light climate. Compared with previous measurements of phytoplanktonic activity at Signy, the microphytobenthos seems to be an important source of primary production. A production estimate of 100.9 mg carbon m^–2, for the ice-free summer period, lies within the range of values of results from other polar studies.     

Nutrient fluxes in a snow-dominated, semi-arid forest: Spatial and temporal patterns

We tested five hypotheses regarding the potential effects of precipitation change on spatial and temporal patterns of water flux, ion flux, and ion concentration in a semiarid, snowmelt-dominated forest in Little Valley, Nevada. Variations in data collected from 1995 to 1999 were used to examine the potential effects of snowpack amount and duration on ion concentrations and fluxes. Soil solution NO₃ ^–, NH₄ ⁺, and ortho-phosphate concentrations and fluxes were uniformly low, and the variations in concentration bore no relationship to snowmelt water flux inputs of these ions. Weathering and cation exchange largely controlled the concentrations and fluxes of base cations from soils in these systems; however, soil solution base cation concentrations were affected by cation concentrations during snowmelt episodes. Soil solution Cl^– and SO₄ ^2– concentrations closely followed the patterns in snowmelt water, suggesting minimal buffering of either ion by soils. In contrast to other studies, the highest concentration and the majority of ion flux from the snowpack in Little Valley occurred in the later phases of snowmelt. Possible reasons for this include sublimation of the snowpack and dry deposition of organic matter during the later stages of snowmelt. Our comparison of interannual and spatial patterns revealed that variation in ion concentration rather than water flux is the most important driver of variation in ion flux. Thus, it is not safe to assume that changes in total precipitation amount will cause concomitant changes in ion inputs to this system.     

Winter CO2 flux from soil and snow surfaces in a cool-temperate deciduous forest, Japan

We measured diurnal and wintertime changes in CO₂ fluxes from soil and snow surfaces in a Japanese cool-temperate Quercus/Betula forest between December 1994 and May 1995. To evaluate the relationship between these winter fluxes and temperature, flux measurements were made with the open-flow infrared gas analyzer (IRGA) method rather than with the more commonly used closed chamber method or the snow CO₂ profile method. The open-flow IRGA method proved to be more successful in measurements of winter CO₂ fluxes than the two standard methods. Despite colder air temperatures, soil temperature profiles were greater than 0°C because of the thermal insulation effect of deep snowpack. This reveals that soil temperature is satisfactory for microbial respiration throughout the winter. Unfrozen soils under the snowpack showed neither diurnal nor wintertime trends in CO₂ fluxes or in soil surface temperature, although there was a daily snow surface CO₂ flux of 0.18–0.32 g m^–2. By combining this with other reference data, Japanese cool-temperate forest soils in snowy regions can be estimated to emit < 100 g m^–2 carbon over an entire winter, and this value accounts for < 15% of the annual emission. In the present study, when data for all winter fluxes were taken together, fluxes were most highly correlated with deep soil temperatures rather than the soil surface temperature. Such a high correlation can be attributed to the relatively increased respiration of the deep soil where the temperature was higher than the soil surface temperature. Thus, deeper soil temperature is a better predictor of winter CO₂ fluxes in cold and snowy ecosystems.     

Process-level controls on CO<Subscript>2</Subscript> fluxes from a seasonally snow-covered subalpine meadow soil,Niwot Ridge,Colorado

Fluxes of CO₂ during the snow-covered season contribute to annual carbon budgets, but our understanding of the mechanisms controlling the seasonal pattern and magnitude of carbon emissions in seasonally snow-covered areas is still developing. In a subalpine meadow on Niwot Ridge, Colorado, soil CO₂ fluxes were quantified with the gradient method through the snowpack in winter 2006 and 2007 and with chamber measurements during summer 2007. The CO₂ fluxes of 0.71 μmol m⁻² s⁻¹ in 2006 and 0.86 μmol m⁻² s⁻¹ in 2007 are among the highest reported for snow-covered ecosystems in the literature. These fluxes resulted in 156 and 189 g C m⁻² emitted over the winter, ~30% of the annual soil CO₂ efflux at this site. In general, the CO₂ flux increased during the winter as soil moisture increased. A conceptual model was developed with distinct snow cover zones to describe this as well as the three other reported temporal patterns in CO₂ flux from seasonally snow-covered soils. As snow depth and duration increase, the factor controlling the CO₂ flux shifts from freeze–thaw cycles (zone I) to soil temperature (zone II) to soil moisture (zone III) to carbon availability (zone IV). The temporal pattern in CO₂ flux in each zone changes from periodic pulses of CO₂ during thaw events (zone I), to CO₂ fluxes reaching a minimum when soil temperatures are lowest in mid-winter (zone II), to CO₂ fluxes increasing gradually as soil moisture increases (zone III), to CO₂ fluxes decreasing as available carbon is consumed. This model predicts that interannual variability in snow cover or directional shifts in climate may result in dramatically different seasonal patterns of CO₂ flux from seasonally snow-covered soils.