Soil organic sulfur dynamics in a coniferous forest

Sulfate microbial immobilization and the mineralization of organic S were measured in vitro in soil horizons (LFH, Ae, Bhf, Bf and C) of the Lake Laflamme watershed (47°17 N, 71°14 O) using ³⁵SO₄. LFH samples immobilized from 23 to 77% of the added ³⁵SO₄ within 2 to 11 days. The ³⁵SO₄ microbial immobilization increased with temperature and reached an asymptote after a few days. The mineral soil generally immobilized less than 20% of the added ³⁵SO₄, and an asymptote was reached after 2 days. An isotopic equilibrium was rapidly reached in mineral horizons. A two-compartment (SO₄ and organic S) model adequately described ³⁵SO₄ microbial immobilization kinetics. The active organic reservoir in the whole soil profile represented less than 1% of the total organic S. The average concentrations of dissolved organic S (DOS) in the soil solutions leaving the LFH, Bhf and Bf horizons were respectively 334, 282 and 143 µgL^–1. Assuming that the DOS decrease with soil depth corresponded to the quantities adsorbed in the B horizons, we estimated that 12 800 kgha^–1 of organic S could have been formed since the last glaciation, which is about 13 times the size of the actual B horizons reservoirs. Our results suggest that the organic S reservoirs present in mineral forest soils are mostly formed by the DOS adsorption resulting from incomplete litter decomposition in the humus layer. The capability of these horizons to immobilize SO₄ from the soil solution would be restricted to a 1% active fraction composed of microorganisms. Despite their refractory nature, these reservoirs can, however, be slowly decomposed by microorganisms and contribute to the S-SO₄ export from the watershed in the long term.     

Patterns of stable S isotopes in a forested catchment as indicators for biological S turnover

Despite intensive biogeochemical research during the last thirty years, the relative importance of biological S turnover for the overall SO ₄ ^2– budget of forested catchments remains uncertain. The objective of the present study was (i) to gain new insight into the S cycle of theLehstenbach catchment (Northeastern Bavaria, Germany) through the analysis of stable isotopes of S and (ii) to differentiate between sites which are hot spots for SO ₄ ^2– reduction and sites where mineralization and adsorption/desorption processes are more important. The ³⁴S values and SO ₄ ^2– concentrations of soil solutions, throughfall and groundwater at four different sites as well as runoff of the catchment were measured. The relatively low variability of ³⁴S in throughfall and bulk precipitation was in contrast to the high temporal and spatial variability of ³⁴S in the soil solution. Sulfate in the soil solution of upland sites was slightly depleted in³⁴S compared to input values. This was most likely due to S mineralization. Sulfate in the soil solution from wetland soils was clearly enriched in³⁴S, indicating dissimilatory SO ₄ ^2– reduction. The observed spatial and temporal patterns of³⁴S turnover and SO ₄ ^2– concentrations might explain the overall balanced S budget of the catchment. At a time of decreasing anthropogenic deposition SO ₄ ^2– is currently released from upland soils. Furthermore, mineralization of organic S may contribute to SO ₄ ^2– release. Wetland soils in the catchment represent a sink for SO ₄ ^2– due to dissimilatory SO ₄ ^2– reduction.     

The biogeochemistry of sulfur at Hubbard Brook

A synthesis of the biogeochemistry of S was done during 34 yr(1964–1965 to 1997–1998) in reference and human-manipulated forestecosystems of the Hubbard Brook Experimental Forest (HBEF), NH. There have beensignificant declines in concentration (–0.44µmol/liter-yr) and input (–5.44mol/ha-yr)of SO₄ ^2– in atmospheric bulk wet deposition, and inconcentration(–0.64 µmol/liter-yr) an d output (–3.74mol/ha-yr) of SO₄ ^2– in stream water ofthe HBEF since 1964. These changes arestrongly correlated with concurrent decreases in emissions of SO₂from the source area for the HBEF. The concentration and input ofSO₄ ^2– in bulk deposition ranged from a low of 13.1µmol/liter (1983–1984) and 211 mol/ha-yr(1997–1998) to a high of 34.7 µmol/liter(1965–1966) and 479 mol/ha-yr (1967–1968), with along-term mean of 23.9 µmol/liter and 336mol/ha-yr during 1964–1965 to 1997–1998. Despiterecentdeclines in concentrations, SO₄ ^2– is the dominantanion in both bulk deposition and streamwater at HBEF. Dry deposition is difficult to measure, especially inmountainousterrain, but was estimated at 21% of bulk deposition. Thus, average totalatmospheric deposition was 491 and 323 mol/ha-yr during1964–1969 and 1993–1998, respectively. Based on the long-term³⁴S pattern associated with anthropogenic emissions,SO₄ ^2– deposition at HBEF is influenced by numerousSO₂sources, but biogenic sources appear to be small. Annual throughfall plusstemflow in 1993–1994 was estimated at 346 molSO₄ ^2–/ha. Aboveground litterfall, for thewatershed-ecosystemaveraged about 180 mol S/ha-yr, with highest inputs (190 molS/ha-yr) in the lower elevation, more deciduous forest zone. Weatheringrelease was calculated at a maximum of 50 mol S/ha-yr. Theconcentration and output of SO₄ ^2– in stream waterranged from a low of 42.3µmol/liter (1996–1997) and 309 mol/ha-yr(1964–1965), to a high of 66.1 µmol/liter(1970–1971) and 849 mol/ha-yr (1973–1974), with along-term mean of 55.5 µmol/liter and 496mol/ha-yr during the 34 yrs of study. Gross outputs ofSO₄ ^2– in stream water consistently exceeded inputsin bulkdeposition and were positively and significantly related to annualprecipitationand streamflow. The relation between gross SO₄ ^2–output and annual streamflow changed with time asatmospheric inputs declined. In contrast to the pattern for bulk depositionconcentration, there was no seasonal pattern for streamSO₄ ^2– concentration. Nevertheless, stream outputs ofSO₄ ^2– were highly seasonal, peaking during springsnowmelt, andproducing a monthly cross-over pattern where net hydrologic flux (NHF) ispositive during summer and negative during the remainder of the year. Nosignificant elevational pattern in streamwaterSO₄ ^2– concentration was observed. Mean annual,volume-weightedsoil water SO₄ ^2– concentrations were relativelyuniform by soil horizon andacross landscape position. Based upon isotopic evidence, much of theSO₄ ^2– entering HBEF in atmospheric depositioncycles throughvegetation and microbial biomass before being released to the soil solution andstream water. Gaseous emissions of S from watershed-ecosystems at HBEF areunquantified, but estimated to be very small. Organic S (carbon bonded andestersulfates) represents some 89% of the total S in soil at HBEF. Some 6% exists asphosphate extractable SO₄ ^2– (PSO₄).About 73% of the total S in the soilprofile at HBEF occurs in the Bs2 horizon, and some 9% occurs in the forestfloor. The residence time for S in the soil was calculated to be 9 yr, butonly a small portion of the total organic soil pool turns over relativelyquickly. The S content of above- and belowground biomass is about 2885mol/ha, of which some 3–5% is in standing dead trees. Yellowbirch, American beech and sugar maple accounted for 89% of the S in trees, with31% in branches, 27% in roots and 25% in the lightwood of boles. The pool of Sin living biomass increased from 1965 to 1982 due to biomass accretion, andremained relatively constant thereafter. Of current inputs to the availablenutrient compartment of the forest ecosystem, 50% is from atmospheric bulkdeposition, 24% from net soil release, 11% from dry deposition, 11% from rootexudates and 4% is from canopy leaching. Comparing ecosystem processes for Sfrom 1964–1969 to 1993–1998, atmospheric bulk deposition decreasedby 34%, stream output decreased by 10%, net annual biomass storage decreased by92%, and net soil release increased by 184% compared to the 1964–1969values. These changes are correlated with decreased emissions of SO₂from the source area for the HBEF. Average, annual bulk deposition inputsexceeded streamwater outputs by 160.0 ± 75.3 SD molS/ha-yr,but average annual net ecosystem fluxes (NEF) were much smaller, mostlynegativeand highly variable during the 34 yr period (–54.3 ± 72.9 SDmol S/ha-yr; NEF range, +86.8 to –229.5). While severalmechanisms may explain this small discrepancy, the most likely are netdesorption of S and net mineralization of organic S largely associated with theforest floor. Our best estimates indicate that additional S from dry depositionand weathering release is probably small and that desorption accounts for about37% of the NEF imbalance and net mineralization probably accounts for theremainder (60%). Additional inputs from dry deposition would result fromunmeasured inputs of gaseous and particulate deposition directly to the forestfloor. The source of any unmeasured S input has important implications for therecovery of soils and streams in response to decreases in inputs of acidicdeposition. Sulfate is a dominant contributor to acid deposition at HBEF,seriously degrading aquatic and terrestrial ecosystems. Because of the strongrelation between SO₂ emissions and concentrations ofSO₄ ^2– in both atmospheric deposition and streamwater at HBEF,further reductions in SO₂ emissions will be required to allowsignificant ecosystem recovery from the effects of acidic deposition. Thedestruction or removal of vegetation on experimental watershed-ecosystems atHBEF resulted in increased rates of organic matter decomposition andnitrification, a lowering of soil and streamwater pH, enhancedSO₄ ^2– adsorption on mineral soil and smallerconcentrations andlosses of SO₄ ^2– in stream water. With vegetationregrowth, this adsorbedSO₄ ^2– is released from the soil, increasingconcentrations andfluxes of SO₄ ^2– in drainage water. Streamwaterconcentration ofSO₄ ^2– and gross annual output ofSO₄ ^2–/ha are essentially the same throughout theHubbard BrookValley in watersheds varying in size by about 4 orders of magnitude, from 3 to3000 ha.     

Four years continuous record of CH4-exchange between the atmosphere and untreated and limed soil of a N-saturated spruce and beech forest ecosystem in Germany

In order to gain information about seasonal and interannual variations of CH₄-fluxes at a spruce control site, a limed spruce site and a beech site of the Höglwald Forest, Bavaria, Germany, complete annual cycles of CH₄-exchange between the soil and the atmosphere with 2-hourly resolution were followed for 4 consecutive years. The ranges of CH₄ fluxes observed for the different sites were: +12.4 to –69.4 g CH₄ m^–2 h^–1 (spruce control site), +11.7 to –51.4 g CH₄ m^–2 h^–1 (limed spruce site), and –4.4 to –167.3 g CH₄ m^–2 h^–1 (beech site). Lowest rates of atmospheric CH₄-uptake or even a weak net-emission of CH₄ by the soils were observed during winter/spring times, whereas highest rates of CH₄-uptake were always found in summer/spring. Over the entire observation period of 4 years, mean CH₄-uptake rates were –1.82 kg CH₄-C ha^–1 yr^–1 at the spruce control site, –1.31 kg CH₄-C ha^–1 yr^–1 at the limed spruce site, and –4.84 kg CH₄-C ha^–1 yr^–1 at the beech site. The results obtained in this study demonstrate that in view of the huge interannual variations in CH₄-fluxes of approx. 1 kg CH₄-C ha^–1 yr^–1, multiple year measurements of CH₄-fluxes are necessary to accurately characterize the sink strength of temperate forest for atmospheric CH₄. By comparison of CH₄-fluxes measured at the spruce control site and the limed spruce site, a significant negative effect of forest floor liming on CH₄-uptake could be demonstrated. Compared to the spruce stand, the beech stand showed on average approx. 3 times higher rates of atmospheric CH₄-uptake, most likely due to pronounced differences between both sites with regard to the organic layer structure and bulk density of the mineral soil. Regression analysis between CH₄-fluxes and environmental parameters revealed that at all sites the dominating factors regulating temporal variations of CH₄ fluxes were soil moisture and soil temperature. Field measurements of CH₄ concentrations in the soil profile and laboratory measurements of CH₄-oxidation and CH₄-production activity on soil samples taken from different soil depths showed that the CH₄-flux at the Höglwald Forest sites is the net-result of simultaneous occurring production and consumption of CH₄ within the soil. Highest CH₄-oxidation activity was found in the uppermost centimeters of the mineral soil, whereas highest potential CH₄-production activity was found in the organic layer.     

The roles of malate and aspartate in C4 photosynthetic metabolism of Flaveria bidentis (L.)

In C₄ grasses belonging to the NADP-malic enzyme-type subgroup, malate is considered to be the predominant C₄ acid metabolized during C₄ photosynthesis, and the bundle sheath cell chloroplasts contain very little photosystem-II (PSII) activity. The present studies showed that Flaveria bidentis (L.), an NADP-malic enzyme-type C₄ dicotyledon, had substantial PSII activity in bundle sheath cells and that malate and aspartate apparently contributed about equally to the transfer of CO₂ to bundle sheath cells. Preparations of bundle sheath cells and chloroplasts isolated from these cells evolved O₂ at rates between 1.5 and 2 mol · min^–1 · mg^–1 chlorophyll (Chl) in the light in response to adding either 3-phosphoglycerate plus HCO ₃ ^– or aspartate plus 2-oxoglutarate. Rates of more than 2 mol O₂ · min^–1 · mg^–1 Chl were recorded for cells provided with both sets of these substrates. With bundle sheath cell preparations the maximum rates of light-dependent CO₂ fixation and malate decarboxylation to pyruvate recorded were about 1.7 mol · min^–1 · mg^–1 Chl. Compared with NADP-malic enzyme-type grass species, F. bidentis bundle sheath cells contained much higher activities of NADP-malate dehydrogenase and of aspartate and alanine aminotransferases. Time-course and pulse-chase studies following the kinetics of radiolabelling of the C-4 carboxyl of C₄ acids from ¹⁴CO₂ indicated that the photosynthetically active pool of malate was about twice the size of the aspartate pool. However, there was strong evidence for a rapid flux of carbon through both these pools. Possible routes of aspartate metabolism and the relationship between this metabolism and PSII activity in bundle sheath cells are considered.Abbreviations DHAP dihydroxyacetone phosphate - NADP-ME(-type) NADP-malic enzyme (type) - NADP-MDH NADP-malate dehydrogenase - OAA oxaloacetic acid - 2-OG 2-oxoglutarate - PEP phosphoenolpyruvate - PGA 3-phosphoglycerate - Pi orthophosphate - Ru5P ribulose 5-phosphate     

The humus layer determines SO<Subscript>4</Subscript><Superscript>2−</Superscript> isotope values in the mineral soil

Biocycling of sulfur (S) has been proposed to play an important role in the recovery of ecosystems following anthropogenic S deposition. Here, we investigated the importance of the humus layer in the biocycling of S in three forested catchments in the Gårdsjön area of southwestern Sweden with differing S inputs and S isotope signature values. These experimental sites consisted of two reference catchments and the Gårdsjön roof experiment catchment (G1), where anthropogenic deposition was intercepted from 1991 until May 2002 by a roof placed over the entire catchment area. Under the roof, controlled levels of deposition were applied, using a sprinkler system, and the only form of S added was marine SO₄²⁻ with a δ of +19.5‰.We installed ion exchange resin bags at the interface between the humus layer and mineral soil at each of the catchments to collect SO₄²⁻ passing through the humus. The resin bags were installed on four occasions, in 1999 and 2000, covering two summer and two winter periods. The ions collected by each bag during these sampling periods were then eluted and their δ values and SO₄²⁻ concentrations determined. The most striking result is that the average δ value in the resin bags was more than 12‰ lower compared to that of the sprinkler water in the G1 roof catchment. There was no increasing trend in the isotope value in the resin bag SO₄²⁻ despite that the roof treatment has been on-going for almost 10 years; the average value for all resin bags was +7.1‰. The highest δ values found in the G1 roof catchment were between +11‰ and +12‰. However, these values were all obtained from resin bags installed at a single sampling location. Throughfall and resin bag δ values were more similar in the two reference catchments: about +7.5‰ in both cases. There was, however, an increase in resin bag δ values during the first winter period, from about +7‰ to +9‰. The resin bag δ value was linearly and positively related (r² = 0.26, p < 0.001) to the amount of SO₄²⁻ extracted from the resin bags, if relatively high amounts (>50 mmol m⁻²) were excluded. High amounts of resin bag SO₄²⁻ seemed to be related to groundwater inputs, as indicated by the δ value. Our results suggest that rapid immobilization of SO₄²⁻ into a large organic S pool may alter the S isotope value and affect the δ values measured in the mineral soil and runoff.     

Seasonal N2 fixation by cool-season pulses based on several15N methods

Summary Accurate estimates of N₂ fixation by legumes are requisite to determine their net contribution of fixed N₂ to the soil N pool. However, estimates of N₂ fixation derived with the traditional¹⁵N methods of isotope dilution and A_N value are costly.Field experiments utilizing¹⁵N-enriched (NH₄)₂SO₄ were conducted to evaluate a modified difference method for determining N₂ fixation by fababean, lentil, Alaska pea, Austrian winter pea, blue lupin and chickpea, and to quantify their net contribution of fixed N₂ to the soil N pool. Spring wheat and non-nodulated chickpea, each fertilized with two N rates, were utilized as non-fixing controls.Estimates of N₂ fixation based on the two control crops were similar. Increasing the N rate to the controls reduced A_N values 32, 18 and 43% respectively in 1981, 1982 and 1983 resulting in greater N₂ fixation estimates. Mean seasonal N₂ fixation by fababean, lentil and Austrian winter pea was near 80 kg N ha^–1, pea and blue lupin near 60 kg N ha^–1, and chickpea less than 10 kg N ha^–1. The net effects of the legume crops on the soil N pool ranged from a 70 kg N ha^–1 input by lentil in 1982, to a removal of 48 kg N ha^–1 by chickpea in 1983.Estimates of N₂ fixation obtained by the proposed modified difference method approximate those derived by the isotope dilution technique, are determined with less cost, and are more reliable than the total plant N procedure.Scientific paper No. 6605. College of Agriculture and Home Economics Research Center, Washington State University, Pullman, WA 99164, U.S.A.     

Long- and short-term changes in sulfate deposition: Effects of the 1990 Clean Air Act Amendments

Annual, volume-weighted concentrations ofSO₄ ^2– in bulk precipitation have declinedsteadily (–0.44 mol/liter-yr) since 1965 atthe Hubbard Brook Experimental Forest (HBEF), NH inresponse to decreases in regional SO₂ emissions(r ² = 0.74). Similar declines in concentrationshave occurred in wet-only precipitation at HBEF and atnearby sites since 1978. However, decreases inSO₄ ^2– concentrations following passage ofthe U.S. Clean Air Act Amendments in 1990, were notunusual from the perspective of long-term data fromthe HBEF. Statistically significant declines (–5.6mol/ha-yr) in bulk deposition of SO₄ ^2– also have occurred since 1965 in relation to decreases inSO₂ emissions (r ² = 0.58), but annualvariations in deposition also are strongly related toamount of precipitation and other factors.     

Cycling of inorganic and organic sulfur in peat from Big Run Bog,West Virginia

Total S concentration in the top 35 cm of Big Run Bog peat averaged 9.7 mol·g — wet mass^–1 (123 mol·g dry mass^–1). Of that total, an average of 80.8% was carbon bonded S, 10.4% was ester sulfate S, 4.5% was FeS₂­S, 2.7% was FeS­S, 1.2% was elemental S, and 0.4% was SO₄ ^2–­S. In peat collected in March 1986, injected with³⁵S­SO₄ ^2– and incubated at 4 °C, mean rates of dissimilatory sulfate reduction (formation of H₂S + S⁰ + FeS + FeS₂), carbon bonded S formation, and ester sulfate S formation averaged 3.22, 0.53, and 0.36 nmol·g wet mass^–1·h^–1, respectively. Measured rates of sulfide oxidation were comparable to rates of sulfate reduction. Although dissolved SO₄ ^2– concentrations in Big Run Bog interstitial water (< 200 µM) are low enough to theoretically limit sulfate reducing bacteria, rates of sulfate reduction integrated throughout the top 30–35 cm of peat of 9 and 34 mmol·m^–2·d^–1 (at 4 °C are greater than or comparable to rates in coastal marine sediments. We suggest that sulfate reduction was supported by a rapid turnover of the dissolved SO₄ ^2– pool (average turnover time of 1.1 days). Although over 90% of the total S in Big Run Bog peat was organic S, cycling of S was dominated by fluxes through the inorganic S pools.     

Nutrient losses in runoff from grasslandand shrubland habitats in southern New Mexico: II. Field plots

Losses of dissolved nutrients (N, P, K, Ca, Mg, Na,Cl, and SO₄) in runoff were measured on grasslandand shrubland plots in the Chihuahuan desert ofsouthern New Mexico. Runoff began at a lowerthreshold of rainfall in shrublands than ingrasslands, and the runoff coefficient averaged 18.6%in shrubland plots over a 7-year period. In contrast,grassland plots lost 5.0 to 6.3% of incidentprecipitation in runoff during a 5.5-year period. Nutrient losses from shrubland plots were greater thanfrom grassland plots, with nitrogen losses averaging0.33 kg ha^–1 yr^–1 vs0.15 kg ha^–1 yr^–1, respectively, during a 3-year period. Thegreater nutrient losses in shrublands were due tohigher runoff, rather than higher nutrientconcentrations in runoff. In spite of these nutrientlosses in runoff, all plots showed net accumulationsof most elements due to inputs from atmosphericdeposition. Therefore, loss of soil nutrients byhillslope runoff cannot, by itself, account for thedepletion of soil fertility associated withdesertification in the Chihuahuan desert.     

Exchange of N-gases at the Höglwald Forest – A summary

During 4 years continuous measurements of N-trace gas exchange were carried out at the forest floor-atmosphere interface at the Höglwald Forest that is highly affected by atmospheric N-deposition. The measurements included spruce control, spruce limed and beech sites. Based on these field measurements and on intensive laboratory measurements of N₂-emissions from the soils of the beech and spruce control sites, a total balance of N-gas emissions was calculated. NO₂-deposition was in a range of –1.6 –2.9 kg N ha^–1 yr^–1 and no huge differences between the different sites could be demonstrated. In contrast to NO₂-deposition, NO- and N₂O-emissions showed a huge variability among the different sites. NO emissions were highest at the spruce control site (6.4–9.1 kg N ha^–1 yr^–1), lowest at the beech site (2.3–3.5 kg N ha^–1 yr^–1) and intermediate at the limed spruce site (3.4–5.4 kg N ha^–1 yr^–1). With regard to N₂O-emissions, the following ranking between the sites was found: beech (1.6–6.6 kg N ha^–1 yr^–1) >> spruce limed (0.7–4.0 kg N ha^–1 yr^–1) > spruce control (0.4–3.1 kg N ha^–1 yr^–1). Average N-trace gas emissions (NO, NO₂, N₂O) for the years 1994–1997 were 6.8 kg N ha^–1 yr^–1 at the spruce control site, 3.6 kg N ha^–1 yr^–1 at the limed spruce site and 4.5 kg N ha^–1 yr^–1 at the beech site. Considering N₂-losses, which were significantly higher at the beech (12.4 kg N ha^–1 yr^–1) than at the spruce control site (7.2 kg N ha^–1 yr^–1), the magnitude of total gaseous N losses, i.e. N₂-N + NO-N + NO₂-N + N₂O-N, could be calculated for the first time for a forest ecosystem. Total gaseous N-losses were 14.0 kg N ha^–1 yr^–1 at the spruce control site and 15.5 kg N ha^–1 yr^–1 at the beech site, respectively. In view of the huge interannual variability of N-trace gas fluxes and the pronounced site differences in N-gas emissions it is concluded that more research is needed in order to fully understand patterns of microbial N-cycling and N-gas production/emission in forest ecosystems and mechanisms of reactions of forest ecosystems to the ecological stress factor of atmospheric N-input.     

Sulfur cycling in forests

Sulfur is essential for the production of certain amino acids in plants. As amino acid sulfur is the major form of sulfur in trees, there is a strong relationship between organic S and organic N in tree tissue. Sulfur deficiencies occur in parts of southeastern Australia and northwestern North America, remote from pollutant inputs. Since bilogical S requirements of forests are modest (< 5 kg · ha^–1 yr^–1 for net vegetative increment), however, atmospheric S inputs in polluted regions (10–80 kg · ha^–1 yr^–1 ) often exceed not only the forest ecosystem S requirement but also its ability to biologically accumulate S. There is some increase in the SO^2– ₄–S content of forest vegetation in response to elevated atmospheric S inputs, but this capacity is apparently easily saturated. Soil SO^2–2 ₄adsorption is often the dominant feature of S cycling in polluted ecosystems and often accounts for net ecosytem S accumulations.Contribution from a symposium on the role of sulfur in ecosystem processes held August 10, 1983, at the annual meeting of the A.I.B.S., Grand Forks, ND; Myron Mitchell, convenor.     

Losses of inorganic carbon and nitrous oxide from a temperate freshwater wetland in relation to nitrate loading

We studied the export of inorganic carbon and nitrous oxide (N₂O) from a Danish freshwater wetland. The wetland is situated in an agricultural catchment area and is recharged by groundwater enriched with nitrate (NO₃ ^–) (1000 M). NO₃ ^– in recharging groundwater was reduced (57.5 mol NO₃ ^– m^–2 yr^–) within a narrow zone of the wetland. Congruently, the annual efflux of carbon dioxide (CO₂) from the sediment was 19.1 mol C m^–2 when estimated from monthly in situ measurements. In comparison the CO₂ efflux was 4.8 mol C m^–2 yr^–1 further out in the wetland, where no NO₃ ^– reduction occurred. Annual exports of inorganic carbon in groundwater and surface water was 78.4 mol C m^–2 and 6.1 mol C m^–2 at the two sites, respectively. N₂O efflux from the sedimenst was detectable on five out of twelve sampling dates and was significantly (P < 0.0001) higher in the NO₃ ^– reduction zone (0.35–9.40 mol m^–2 h^–1, range of monthly means) than in the zone without NO₃ ^– reduction (0.21–0.41 mol m^–2 h^–1). No loss of dissolved N₂O could be measured. Total annual export of N₂O was not estimated. The reduction of oxygen (O₂) in groundwater was minor throughout the wetland and did not exceed 0.2 mol 0₂ m^–2yr^–1. Sulfate (SO₄ ^––) was reduced in groundwater (2.1 mol SO₄ ^–– m^–2 yr^–1) in the zone without NO₃ ^– reduction. Although the NO₃ ^– in our wetland can be reduced along several pathways our results strongly suggest that NO₃ ^– loading of freshwater wetlands disturb the carbon balance of such areas, resulting in an accelerated loss of inorganic carbon in gaseous and dissolved forms.     

Effect of urea fertilizer and environmental factors on CH4 emissions from a Louisiana,USA rice field

Methane emissions from a flooded Louisiana, USA, rice field were measured over the first cropgrowing season. Microplots contained the semidwarf Lemont rice cultivar drill seeded into a Crowley silt loam soil (Typic Albaqualfs). Urea fertilizer was applied preflood at rates of 0, 100, 200 and 300 kg N ha^–1. Emissions of CH₄ from the plots to the atmosphere were measured over a 86-d sampling period until harvest. Methane samples were collected in the morning hours (0730–0930) using a closed-chamber technique. Emissions of CH₄ were highly variable over the first cropping season and a significant urea fertilizer effect was observed. Two peak CH₄ emission periods were observed and occurred about 11 d after panicle differentiation and during the ripening stages. Maximum CH₄ emmissions from the 0, 100, 200 and 300 urea-N treatments were 6.0, 8.9, 9.8 and 11.2 kg CH₄ ha^–1 d^–1, respectively. These flux measurements corresponded to approximately 210, 300, 310 and 360 kg CH₄ evolved ha^–1 over the 86-d sampling period for the 4 treatments.     

Rate of accumulation and emission of N2, N2O and CH4 from a flooded rice soil

In a field experiment using microplots, a flooded Crowley silt loam (Typic Albaqualfs) rice soil was fertilized with ¹⁵N labelled (60–74 atom %) urea and KNO₃. Emission of N₂, N₂O and CH₄ and accumulation in soil were measured for 21 d after fertilizer application.Emission of ¹⁵N₂-N measured from the urea and KNO₃ treated plots ranged from <15 to 570 and from 330 to 3,420 g ha^–1 d^–1, respectively. Entrapped ¹⁵N₂-N in the urea treated microplots was significantly lower (<15 g to 2.1 kg ha^–1) on all sampling dates compared to the ¹⁵N₂-N gas accumulation in the KNO₃ treated plots (6.4 to 31.5 kg ha^–1). Emissions of N₂O-N were low and did not exceed 4 g ha^–1 d^–1. Fluxes of CH₄ from the fertilizer and control plots were low and never exceeded 33 g ha^–1 d^–1. Maximum accumulation of CH₄ in the flooded soil measured 460 and 195 g ha^–1 for the urea and KNO₃ treatments, respectively.     

Nitrogen nutrition of rice plants under salinity

Two rice (Oryza sativa L.) cultivars, Koshihikari and Pokkali, were grown in solution culture at three concentrations of NaCl or Na₂SO₄ [0 (S0), 50 (S1), and 100 (S2) mmol dm^–3] and three N contents [0.7 (N1), 7 (N2) and 14 (N3) mmol dm^–3]. Salinity significantly decreased dry matter of both cultivars. Pokkali had better growth than Koshihikari under both saline and non-saline conditions. Applications of N enhanced development of shoot dry mass under S0 and S1 treatments up to N2. Under S2, N application had no effect on shoot dry mass of both cultivars. Root dry mass of both cultivars decreased with increasing N application at S1 and S2. Shoot and root NO₃-N content in both rice cultivars increased with increasing N concentration in the nutrient solutions. The absorption of NO₃-N was less in Koshihikari than Pokkali plants, and also was much less in Cl^– than SO₄ ^2– salinity suggesting the antagonism between Cl^– and NO₃ ^–. In addition a significant negative correlation between concentrations of NO₃-N and Cl^– in the shoots or roots was observed in both cultivars     

The effect of carbon dioxide on the growth kinetics of fructose-limited chemostat cultures of Acetobacterium woodii DSM 1030

The growth of the anaerobic acetogenic bacterium Acetobacterium woodii DSM 1030 was investigated in fructose-limited chemostat cultures. A defined medium was developed which contained fructose, mineral salts, cysteine · HCl and Ca pantothenate (1 mg · 1^–1) supplied in a vitamin supplement. Growth at high dilution rates was dependent on the presence of CO₂ in the gas phase. The ^max was found to be 0.16 h^–1 and the fructose maintenance requirement was 0.1 to 0.13 mmol fructose · (g dry wt)^–1 · h^–1. A growth yield of 61 g dry wt · (mol fructose)^–1, corrected for the cell maintenance requirement and for incorporation of fructose carbon into cell biomass, was determined from the fructose consumption. A corresponding growth yield of 69 g dry wt · (mol fructose)^–1 was calculated from the acetate production assuming that fructose fermentation was homoacetogenic. A Y_ATP of 12.2 to 13.8 g dry wt · (mol ATP)^–1 was calculated from these growth yields using a value of 5 mol ATP · (mol fructose)^–1 as an estimate of the amount of ATP synthesised from fructose fermentation. The addition of yeast extract (0.5 g · 1^–1) to the medium did not influence the ^max or cell yield. After prolonged growth under fructose-limited conditions the requirement of the culture for CO₂ in the gas phase was reduced.Abbreviations YE yeast extract - IC inorganic carbon - D fermenter dilution rate : h^–1 - M_X maintenance requirement for X: mmol X · (g dry wt)^–1 · h^–1 - X may be fructose (Fruct), fructose consumed in energy metabolism (Fruct [E]), acetate (Ac) - ATP CO₂, NH _inf4 ^sup+ or Pi - qX specific rate of utilisation or consumption of X: mmol X · (g dry wt)^–1 · h^–1 - V fermenter volume: litre - rC · Cell, fermenter cell carbon production: mmol C · h^–1 - Y_X yield of cells on X: g dry wt · (mol X)^–1 - Y _infx ^supmax the yield corrected for cell maintenance: g dry wt · (mol X)^–1 - S_ATP stoichiometry of ATP synthesis from fructose: mol ATP · (mol frucose)^–1 - x cell concentration: g dry wt · 1^–1 - specific growth rate : h^–1 - ^max maximum specific growth rate: h^–1     

Influence of elevated CO2 and nitrogen nutrition on rice plant growth,soil microbial biomass,dissolved organic carbon and dissolved CH4

Rice (Oryza sativa) was grown in six sunlit, semi-closed growth chambers for two seasons at 350 L L^–1 (ambient) and 650 L L^–1 (elevated) CO₂ and different levels of nitrogen (N) supplement. The objective of this research was to study the influence of CO₂ enrichment and N nutrition on rice plant growth, soil microbial biomass, dissolved organic carbon (DOC) and dissolved CH₄. Elevated CO₂ concentration ([CO₂]) demonstrated a wide range of enhancement to both above- and below-ground plant biomass, in particular to stems and roots (for roots when N was not limiting) in the mid-season (80 days after transplanting) and stems/ears at the final harvest, depending on season and the level of N supplement. Elevated [CO₂] significantly increased microbial biomass carbon in the surface 5 cm soil when N (90 kg ha^–1) was in sufficient supply. Low N supplement (30 kg ha^–1) limited the enhancement of root growth by elevated [CO₂], leading consequently to diminished response of soil microbial biomass carbon to CO₂ enrichment. The concentration of dissolved CH₄ (as well as soil DOC, but to a lesser degree) was observed to be positively related to elevated [CO₂], especially at high rate of N application (120 kg ha^–1) or at 10 cm depth (versus 5 cm depth) in the later half of the growing season (at 80 kg N ha^–1). Root senescence in the late season complicated the assessment of the effect of elevated [CO₂] on root growth and soil organic carbon turnover and thus caution should be taken when interpreting respective high CO₂ results.     

Nitrate and chloride ions have different permeation pathways in skeletal muscle fibers ofRana pipiens

Summary The effects of pH on the permeability and conductance of the membranes to nitrate and to chloride of semitendinosus and lumbricalis muscle fibers were examined.Membrane potential responses to quick solution changes were recorded in semitendinosus fibers initially equilibrated in isotonic, high K₂SO₄ solutions. External solutions were first changed to ones in which either Rb⁺ or Cs⁺ replaced K⁺ and then to solutions containing either NO ₃ ^– or Cl^– to replace SO ₄ ^2– . The hyperpolarizations produced by Cl^– depend on external pH, being smaller in acid than in alkaline solutions. By contrast, hyperpolarizations produced by NO ₃ ^– were independent of external pH over a pH range from 5.5 to 9.0.In addition, voltage-clamp measurements were made on short lumbricalis muscle fibers. Initially they were equilibrated in isotonic solutions containing mainly K₂SO₄ plus Na₂SO₄. KCl or KNO₃ were added to the sulfate solutions and the fibers were equilibrated in these new solutions. When finally equilibrated the fibers had the same volume they had in the sulfate solutions before the additions. Constant hyperpolarizing voltage pulses of 0.6-sec duration were applied when all external K⁺ was replaced by TEA⁺. For these conditions, inward currents flowing during the voltage pulses were largely carried by Cl^– or NO ₃ ^– depending on the final equilibrating solution. Cl^– currents during voltage pulses were both external pH and time dependent. By contrast, NO ₃ ^– currents were independent of both external pH and time.The voltage dependence of NO ₃ ^– currents could be fit by constant field equations with aP _{NO 3} of 3.7·10^–6 cm/sec. The voltage dependence of the initial or instantaneous Cl^– currents at pH 7.5 and 9.0 could also be fit by constant field equations with P_Cl of 5.8·10^–6 and 7.9·10^–6 cm/sec, respectively. At pH 5.0, no measurable instantaneous Cl^– currents were found.From these results we conclude that NO ₃ ^– does not pass through the pH, time-dependent Cl^– channels but rather passes through a distinct set of channels. Furthermore, Cl^– ions do not appear to pass through the channels which allow NO ₃ ^– through. Consequently, the measured ratio ofP _Cl/P _{NO 3} based on membrane potential changes to ionic changes made on intact skeletal muscle fibers is not a measure of the selectivity of a single anion channel but rather is a measure of the relative amounts of different channel types.