Unidirectional fluxes in saturated single-file pores of biological and artificial membranes. II. Asymptotic behavior at high degrees of saturation

A rate-theory analysis of isotope fluxes in filled single-file pores is given. It is assumed that the relation between the activities in solution and the fluxes can be described by the asymptotic laws for high activities. Only pore states containing no more than two vacancies need be taken into account. The flux-ratio exponent n, which is obtained by representing the ratio of unidirectional fluxes as a power of the electrochemical activity ratio, is found to provide information about m, the number of sites per pore. The general relation between n and m, for m ≥ 4, is m ? 3 ≤ n ≤ m ?1. For m = 2 or 3 the minimum value is 1. The theoretical results are discussed with respect to the internal structure of gramicidin pores in artificial bilayer membranes.     

Brownian dynamics study of flux ratios in sodium channels

Measurements of unidirectional fluxes in ion channels provide one of the experimental methods for studying the steps involved in ion permeation in biological pores. Conventionally, the number of ions in the pore is inferred by fitting the ratio of inward and outward currents to an exponential function with an adjustable parameter known as the flux ratio exponent. Here we investigate the relationship between the number of ions in the pore and the flux ratio exponent in a model sodium channel under a range of conditions. Brownian dynamics simulations enable us to count the precise number of ions in the channel and at the same time measure the currents flowing across the pore in both directions. We show here that the values of the flux ratio exponent n′ ranges between 1 and 3 and is highly dependent on the ionic concentrations in which measurements are made. This is a consequence of the fact that both inward and outward currents are susceptible to saturation with increasing concentration. These results indicate that measurements of the flux ratio exponent cannot be directly related to the number of ions in the pore and that interpretation of such experimental measurements requires careful consideration of the conditions in which the study is made.     

The effect of ethanol on ion transport in frog skin

Frog skin has been used as a model epithelial sodium-transporting system to study the effect of ethanol on ion transport. Treatment of the outside of frog skin with ethanol decreased the net sodium transport due to inhibition of ²²Na⁺ influx. Ethanol did not alter sodium outflux when bathing the outside of the skin. The inhibition was in proportion to the concentration of ethanol, 0.25 M resulting in 50% inhibition. The chloride permeability of the skin was increased several-fold when the skin was exposed to ethanol in either bathing solution. With 0.4 M ethanol in the inner bathing solution, all the unidirectional fluxes of Na⁺ and Cl^? were increased. The movement of Cl^? was evaluated by comparison of Cl^? flux with urea flux, since urea is thought to move passively across frog skin via an extracellular (shunt) pathway. Chloride flux was increased to a greater extent than urea flux. These experiments indicate that ethanol affects chloride permeability beyond an increase in extracellular ion flow and independent of its effect on Na⁺ transport.     

Nitrous oxide and methane fluxes from soils of the Orinoco savanna under different land uses

The study investigates the effect of land‐use change on nitrous oxide (N₂O) and methane (CH₄) fluxes from soil, in savanna ecosystems of the Orinoco region (Venezuela). Gas fluxes were measured by closed static chambers, in the wet and dry season, in representative systems of land management of the region: a cultivated pasture, an herbaceous savanna, a tree savanna and a woodland (control site). Higher N₂O emissions were observed in the cultivated pasture and in the herbaceous savanna compared with the tree savanna and the woodland, and differences were mainly related to fine soil particle content and soil volumetric water content measured in the studied sites. Overall N₂O emissions were quite low in all sites (0–1.58 mg N₂O‐N m^?2 day^?1). The cultivated pasture and the woodland savanna were on average weak CH₄ sinks (?0.05±0.07 and ?0.08±0.05 mg CH₄ m^?2 day^?1, respectively), whereas the herbaceous savanna and the tree savanna showed net CH₄ production (0.23±0.05 and 0.19±0.05 mg CH₄ m^?2 day^?1, respectively). Variations of CH₄ fluxes were mainly driven by variation of soil water‐filled pore space (WFPS), and a shift from net CH₄ consumption to net CH₄ production was observed at around 30% WFPS. Overall, the data suggest that conversion of woodland savanna to managed landscape could alter both CH₄ and N₂O fluxes; however, the magnitude of such variation depends on the soil characteristics and on the type of land management before conversion.     

Carbon storage and fluxes in ponderosa pine forests at different developmental stages

We compared carbon storage and fluxes in young and old ponderosa pine stands in Oregon, including plant and soil storage, net primary productivity, respiration fluxes, eddy flux estimates of net ecosystem exchange (NEE), and Biome‐BGC simulations of fluxes. The young forest (Y site) was previously an old‐growth ponderosa pine forest that had been clearcut in 1978, and the old forest (O site), which has never been logged, consists of two primary age classes (50 and 250 years old). Total ecosystem carbon content (vegetation, detritus and soil) of the O forest was about twice that of the Y site (21 vs. 10 kg C m^?2 ground), and significantly more of the total is stored in living vegetation at the O site (61% vs. 15%). Ecosystem respiration (R_e) was higher at the O site (1014 vs. 835 g C m^?2 year^?1), and it was largely from soils at both sites (77% of R_e). The biological data show that above‐ground net primary productivity (ANPP), NPP and net ecosystem production (NEP) were greater at the O site than the Y site. Monte Carlo estimates of NEP show that the young site is a source of CO₂ to the atmosphere, and is significantly lower than NEP(O) by c. 100 g C m^?2 year^?1. Eddy covariance measurements also show that the O site was a stronger sink for CO₂ than the Y site. Across a 15‐km swath in the region, ANPP ranged from 76 g C m^?2 year^?1 at the Y site to 236 g C m^?2 year^?1 (overall mean 158 ± 14 g C m^?2 year^?1). The lowest ANPP values were for the youngest and oldest stands, but there was a large range of ANPP for mature stands. Carbon, water and nitrogen cycle simulations with the Biome‐BGC model suggest that disturbance type and frequency, time since disturbance, age‐dependent changes in below‐ground allocation, and increasing atmospheric concentration of CO₂ all exert significant control on the net ecosystem exchange of carbon at the two sites. Model estimates of major carbon flux components agree with budget‐based observations to within ± 20%, with larger differences for NEP and for several storage terms. Simulations showed the period of regrowth required to replace carbon lost during and after a stand‐replacing fire (O) or a clearcut (Y) to be between 50 and 100 years. In both cases, simulations showed a shift from net carbon source to net sink (on an annual basis) 10–20 years after disturbance. These results suggest that the net ecosystem production of young stands may be low because heterotrophic respiration, particularly from soils, is higher than the NPP of the regrowth. The amount of carbon stored in long‐term pools (biomass and soils) in addition to short‐term fluxes has important implications for management of forests in the Pacific North‐west for carbon sequestration.     

Unidirectional fluxes and blood flow

Unidirectional fluxes determined in vivo (in experiments with intact blood supply) on the basis of the permeation rate of labeled molecules are functions of the absorption site blood flow rate. As the permeability of the substances increases, the “blood flow-independent” unidirectional flux changes to a “blood flow-limited” flux. By contrast, the ratio of unidirectional fluxes is independent of blood flow, provided that permeation of the substance out of the draining capillaries is proportional to the concentration and the permeation into the opposite direction does not depend on the trans concentration. For this special case, flux ratio variations must be due to changes in permeation characteristics of the layer overlying the capillaries (i.e. epithelium, unstirred fluid layer), while unidirectional flux variations can also be attributed to changes in the absorption site blood flow rate.     

Paired-tower measurements of carbon and energy fluxes following disturbance in the boreal forest

Disturbances by fire and harvesting are thought to regulate the carbon balance of the Canadian boreal forest over scales of several decades. However, there are few direct measurements of carbon fluxes following disturbances to provide data needed to refine mathematical models. The eddy covariance technique was used with paired towers to measure fluxes simultaneously at disturbed and undisturbed sites over periods of about one week during the growing season in 1998 and 1999. Comparisons were conducted at three sites: a 1‐y‐old burned jackpine stand subjected to an intense crown fire at the International Crown Fire Modelling Experiment site near Fort Providence, North‐west Territories; a 1‐y‐old clearcut aspen area at the EMEND project near Peace River, Alberta; and a 10‐y‐old burned, mixed forest near Prince Albert National Park, Saskatchewan. Nearby mature forest stands of the same types were also measured as controls. The harvested site had lower net radiation (R_n), sensible (H) and latent (LE) heat fluxes, and greater ground heat fluxes (G) than the mature forest. Daytime CO₂ fluxes were much reduced, but night‐time CO₂ fluxes were identical to that of the mature aspen forest. It is hypothesized that the aspen roots remained alive following harvesting, and dominated soil respiration. The overall effect was that the harvested site was a carbon source of about 1.6 gC m^?2 day^?1, while the mature site was a sink of about ?3.8 gC m^?2 day^?1. The one‐year‐old burn had lower R_n, H and LE than the mature jackpine forest, and had a continuous CO₂ efflux of about 0.8 gC m^–2 day^?1 compared to the mature forest sink of ? 0.5 g C m^?2 day^?1. The carbon source was likely caused by decomposition of fire‐killed vegetation. The 10‐y‐old burned site had similar H, LE, and G to the mature mixed forest site. Although the diurnal amplitude of the CO₂ fluxes were slightly lower at the 10‐y‐old site, there was no significant difference between the daily integrals (? 1.3 gC m^?2 day^?1 at both sites). It appears that most of the change in carbon flux occurs within the first 10 years following disturbance, but more data are needed on other forest and disturbance types for the first 20 years following the disturbance event.     

Reciprocal fluxes of stream and riparian invertebrates in a coastal California basin with Mediterranean climate

Stream and riparian food webs are connected by reciprocal fluxes of invertebrates, and a growing number of studies demonstrate strong effects of these subsidies on consumers and food webs in both habitats. However, despite its importance in understanding energy flow between these habitats, seasonality of reciprocal subsidies has been examined only in a single temperate system in Japan. We measured input of terrestrial invertebrates and emergence of adult aquatic insects for 14?months in two adjacent streams in a coastal Mediterranean basin in California to assess seasonal patterns, annual fluxes, and local variation. Fluxes of terrestrial and aquatic invertebrates fluctuated seasonally and were relatively synchronous, although in the fall of 2004, terrestrial inputs peaked 1?C2?months earlier than emergence. Terrestrial inputs were similar in the two streams with annual flux of 7.9?C8.6?g dry mass?m^?2?year^?1. Emergence differed between the streams: annual emergence was 7.8?g?m^?2?year^?1 (similar to terrestrial flux) in one reach but 5.3?g?m^?2?year^?1 from the other. The presence of streambed travertine in the reach with lower emergence was the primary difference in habitat between the streams, suggesting that travertine may reduce emergence and alter net reciprocal flux. Comparison of our results with those from Japan suggests that seasonality and net annual flux of reciprocal stream-riparian subsidies vary among biomes due to differences in climate, vegetation, and geography. Our results also indicate that local factors, such as travertine, may cause reciprocal fluxes to vary at finer spatial scales.     

Fluxes of N2O and CH4 from soils of savannas and seasonally-dry ecosystems

Aim Savannas and seasonally‐dry ecosystems cover a significant part of the world's land surface. If undisturbed, these ecosystems might be expected to show a net uptake of methane (CH₄) and a limited emission of nitrous oxide (N₂O). Land management has the potential to change dramatically the characteristics and gas exchange of ecosystems. The present work investigates the contribution of warm climate seasonally‐dry ecosystems to the atmospheric concentration of nitrous oxide and methane, and analyses the impact of land‐use change on N₂O and CH₄ fluxes from the ecosystems in question. Location Flux data reviewed here were collected from the literature; they come from savannas and seasonally‐dry ecosystems in warm climatic regions, including South America, India, Australasia and Mediterranean areas. Methods Data on gas fluxes were collected from the literature. Two factors were considered as determinants of the variation in gas fluxes: land management and season. Land management was grouped into: (1) control, (2) ‘burned only’ and (3) managed ecosystems. The season was categorized as dry or wet. In order to avoid the possibility that the influence of soil properties on gas fluxes might confound any differences caused by land management, sites were grouped in homogeneous clusters on the basis of soil properties, using multivariate analyses. Inter‐ and intra‐cluster analysis of gas fluxes were performed, taking into account the effects of season, land management and main vegetation types. Results Soils were often acid and nutrient‐poor, with low water retention. N₂O emissions were generally very low (median flux 0.32 mg N₂O m^?2 day^?1), and no significant differences were observed between woodland savannas and managed savannas. The highest fluxes (up to 12.9 mg N₂O m^?2 day^?1) were those on relatively fertile soils with high air‐filled porosity and water retention. The effect of season on N₂O production was evident only when sites were separated in homogeneous groups on the basis of soil properties. CH₄ fluxes varied over a wide range (?22.9 to 3.15 mg CH₄ m^?2 day^?1, where the negative sign denotes removal of gas from the atmosphere), with an annual average daily flux of ?0.48 ± 0.96 (SD) mg CH₄ m^?2 day^?1 in undisturbed (control) sites. Land‐use change dramatically reduced this CH₄ sink. Managed sites were weak sinks of CH₄ in the dry season and became sources of CH₄ in the wet season. This was particularly evident for pastures. Burning alone did not reduce soil net CH₄ oxidation, but decreased N₂O production. Main conclusions Despite the low potential for N₂O production, both in natural and managed conditions, tropical seasonally‐dry ecosystems represent a significant source of N₂O (4.4 Tg N₂O year^?1) on a global scale, as a consequence of the large area they occupy. The same environments represent a potential CH₄ sink of 5.17 Tg CH₄ year^?1. However, assuming that c. 30% of the tropical land is converted to different uses, the sink would be reduced to 3.2 Tg CH₄ year^?1. The limited information on fluxes from Mediterranean ecosystems does not allow a meaningful scaling up.     

Flux ratio of valinomycin-mediated K+ fluxes across the human red cell membrane in the presence of the protonophore CCCP

Summary The ratio of valinomycin-mediated unidirectional K⁺ fluxes across the human red cell membrane, has been determined in the presence of the protonophore carbonylcyanidem-chlorophenylhydrazone, CCCP, using the K⁺ net efflux and⁴²K influx. The driving force for the net efflux (V _m –E _K ⁺) has been calculated from the membrane potential, estimated by the CCCP-mediated proton distribution and the Nernst potential for potassium ions across the membrane. An apparent driving potential for the K⁺ net efflux has been calculated from the K⁺ flux ratio, determined in experiments where the valinomycin and CCCP concentrations were varied systematically. This apparent driving force, in conjunction with the actual driving force calculated on basis of the CCCP estimated membrane potential, is used to calculate a flux ratio exponent, which represents an estimate of the deviation of valinomycin-mediated K⁺ transport from unrestricted electrodiffusion, when protonophore is present.In the present work, the flux ratio exponent is found to be 0.90 when the CCCP concentration is 5.0 m and above, while the exponent decreases to about 0.50 when no CCCP is present. The influence of CCCP upon the rate constants in the valinomycin transport cycle is discussed. The significance of this result is that red cell membrane potentials are overestimated, when calculated from valinomycin-mediated potassium isotope fluxes, using a constant field equation.     

Sensitivity Analysis of Flux Determination in Heart by H218O -provided Labeling Using a Dynamic Isotopologue Model of Energy Transfer Pathways

To characterize intracellular energy transfer in the heart, two organ-level methods have frequently been employed: inversion and saturation transfer, and dynamic labeling. Creatine kinase (CK) fluxes obtained by following oxygen labeling have been considerably smaller than the fluxes determined by saturation transfer. It has been proposed that dynamic labeling determines net flux through CK shuttle, whereas saturation transfer measures total unidirectional flux. However, to our knowledge, no sensitivity analysis of flux determination by oxygen labeling has been performed, limiting our ability to compare flux distributions predicted by different methods. Here we analyze oxygen labeling in a physiological heart phosphotransfer network with active CK and adenylate kinase (AdK) shuttles and establish which fluxes determine the labeling state. A mathematical model consisting of a system of ordinary differential equations was composed describing enrichment in each phosphoryl group and inorganic phosphate. By varying flux distributions in the model and calculating the labeling, we analyzed labeling sensitivity to different fluxes in the heart. We observed that the labeling state is predominantly sensitive to total unidirectional CK and AdK fluxes and not to net fluxes. We conclude that measuring dynamic incorporation of into the high-energy phosphotransfer network in heart does not permit unambiguous determination of energetic fluxes with a higher magnitude than the ATP synthase rate when the bidirectionality of fluxes is taken into account. Our analysis suggests that the flux distributions obtained using dynamic labeling, after removing the net flux assumption, are comparable with those from inversion and saturation transfer.     

Predicting N2O emissions from agricultural land through related soil parameters

An empirical model of nitrous oxide emission from agricultural soils has been developed. It is based on the relationship between N₂O and three soil parameters – soil mineral N (ammonium plus nitrate) content in the topsoil, soil water‐filled pore space and soil temperature – determined in a study on a fertilized grassland in 1992 and 1993. The model gave a satisfactory prediction of seasonal fluxes in other seasons when fluxes were much higher, and also from other grassland sites and from cereal and oilseed rape crops, over a wide flux range (< 1 to > 20 kg N₂O‐N ha^?1 y^?1). However, the model underestimated emissions from potato and broccoli crops; possible reasons for this are discussed. This modelling approach, based as it is on well‐established and widely used soil measurements, has the potential to provide flux estimates from a much wider range of agricultural sites than would be possible by direct measurement of N₂O emissions.     

Net sediment N2 fluxes in a southern New England estuary: variations in space and time

Over the past three decades, Narragansett Bay has undergone various ecological changes, including significant decreases in water column chlorophyll a concentrations, benthic oxygen uptake, and benthic nutrient regeneration rates. To add to this portrait of change, we measured the net flux of N₂ across the sediment–water interface over an annual cycle using the N₂/Ar technique at seven sites in the bay for comparison with measurements made decades ago. Net denitrification rates ranged from about 10–90 μmol N₂–N m^?2 h^?1 over the year. Denitrification rates were not significantly different among sites and had no clear correlation with temperature. Net nitrogen fixation (?5 to ?650 μmol N₂–N m^?2 h^?1) was measured at three sites and only observed in summer (June–August). Neither denitrification nor nitrogen fixation exhibited a consistent relationship with sediment oxygen demand or with fluxes of nitrite, nitrate, ammonium, total dissolved inorganic nitrogen, or dissolved inorganic phosphate across all stations. In contrast to the mid-bay historical site where denitrification rates have declined, denitrification rates in the Providence River Estuary have not changed significantly over the past 30 years.     

Short circuit current in tight and leaky epithelia

The effect of short circuit current on the unidirectional fluxes of ions transported across tight and leaky epithelia was investigated. It was found that short circuiting of the frog gastric mucosa (classified as a tight epithelium) caused a decrease of the passive J_ms^C1 and a significant increase of the net Cl^? secretion. However, no significant change of H⁺ secretory rate was observed. On the other hand, short circuiting of the mouse intestine (a known leaky membrane) caused a simultaneous increase of both J_ms and J_sm fluxes of Na⁺ while the net fluxes of Na⁺ and Cl^? remained unchanged. Also, short circuiting did not change the water permeability of the mouse intestine. To explain some of these results a theoretical model is presented to demonstrate that while short circuiting can block the passive ionic movement, it will cause an increase in the energy consumption of the system and introduce certain important changes in the ionic barriers and e.m.fs. The simultaneous increase in the unidirectional fluxes of Na⁺ under short circuit conditions can best be explained by a decrease in the polarized nature of the transepithelial shunt, thereby increasing the diffusion coefficient of the ion(s). Such an increase is specially favorable to the Na⁺ rather than an anion.     

Gross precipitation and throughfall chemistry in legume species planted in Northeastern México

Plant cover modifies throughfall chemistry, and the solute concentration is dependent on the plant species at any given site. The chemistry of gross rainfall and throughfall of four endemic species planted in northeastern Mexico was evaluated from March 1996 to March 1997. Chemical solutes measured included Ca, K, Mg, Na, Fe, Mn, Cu, and Zn. Dry deposition and canopy leaching fluxes were estimated following the canopy budget model. Variance analyses tested the statistical dependence of the total and net fluxes on the species and seasons. Regression analysis tested the dependence of chemical concentrations on rainfall depth and lag time between rains. A total of 52 rainfall events were recorded during the study period summing 523 mm. Significant differences were noted on the total and net fluxes between the plant species. For total flux, average throughfall (37.8 kg ha^?1 year^?1) almost doubled the flux of solutes compared to rainfall (24.1 kg ha^?1 year^?1). Pithecellobium ebano (Berland.) C.H. Mull. (43.3 kg ha^?1 year^?1), Acacia berlandieri Benth. (38.7 kg ha^?1 year^?1), and Pithecellobium pallens (Bent.) Standl. (38.4 kg ha^?1 year^?1) recorded the highest total flux of solutes, and Acacia rigidula Benth. (30.9 kg ha^?1 year^?1) the smallest. Chemical solutes showed significant differences for total and net fluxes. Ca was the dominant cation with 48% and 52% of the total constituent flux for rainfall and throughfall, respectively. However, K, Mg and Cu approximately doubled in throughfall in contrast to gross rainfall. Species with the largest aboveground biomass had lower throughfall volumes (i.e., higher interception rates) but higher chemical solute inputs to the forest floor. Rainfall depth and lag time between rains explained part of the variation for most species, stressing the partial dependence of the washing effect and the amount of dry deposition on canopies. This research discusses the importance and the sources of incoming solutes on the studied plant species.     

Coupling heat flux dynamics with meteorological conditions in the green roof ecosystem

Green roofs can notably modify the thermal properties of the building envelope and adjacent air to bring environmental benefits. This study investigates the heat flux dynamics of the tropical green roof ecosystem to provide a scientific basis for design and management. Green roof experimental plots were established to monitor the total solar radiation, net radiation, and micrometeorological parameters. The data permit calculation of sensible and latent heat fluxes using the Bowen ratio energy balance (BREB) method. The results demonstrated the life cycle characteristics of heat flux components. The dynamic changes of sensible (H), latent (λE) and soil (G) heat fluxes were denoted by single-peak quadratic curves. Net radiation (R_n) was largely determined by quantity and variation trends of λE, reaching at 1300 h a maximum λE of 655 W m^?2 and maximum H of 369 W m^?2. Temporal heat-flux fluctuations were strongly correlated with meteorological variables. Extreme values of H and λE correlated well with precipitation and temperature (R² = 0.78). Dynamics of heat-flux magnitude and partitioning demonstrated notable differences by daily and season periods. They displayed considerable variations in flux partitioning, with Bowen ratios strongly correlated with weather conditions and vegetation types. The energy budget of the green roof ecosystem is unbalanced with a heat loss of about 15.5% caused by soil and canopy heat reserve. The passive indoor cooling effect under the green roof is attributed to the unbalanced energy closure.     

A simple apparatus for performing short-time (1–2 seconds) uptake measurements in small volumes; its application to d-glucose transport studies in brush border vesicles from rabbit jejunum and ileum

An automated procedure allows uptake measurements with incubation times as short as 0.5 s and with volumes of 10–20 μl. Using this technique the kinetic parameters K_m and V of d-glucose transport in brush border vesicles from rabbit small intestine could be determined from unidirectional fluxes. A comparison of the data obtained from jejunum and from ileum shows that the K_m for d-glucose is the same in both parts of the intestine, whereas the maximum flux is significantly larger in the jejunum.     

K+ Secretion in Rat Distal Jejunum

The Ussing chamber technique was used to measure unidirectional Rb⁺ fluxes under short-circuit conditions across tissue sheets from proximal, central, and distal jejunum of rats. Whereas the proximal and central parts of the jejunum did not show any net transport of Rb⁺, there was a net secretion of around 0.2 μmol hr⁻¹ cm⁻² in the distal segment. This secretion could not be influenced significantly by mucosal application of K⁺ channel blockers such as Ba²⁺ (5 mm), tetraethylammonium (20 mm) or quinine (1 mm). Serosal ouabain (1 mm) blocked net secretion by increasing mucoserosal flux. Blockers of H⁺/K⁺ ATPases could not alter net fluxes of Rb⁺. Stimulation of Cl⁻ secretion by forskolin (10 μm) or of Na⁺ absorption by serine (10 mm) failed to influence the observed secretion of Rb⁺. Adrenaline (10 μm) also had no effect on Rb⁺ fluxes. Blocking Na⁺/H⁺ exchange by 5-(N-Ethyl-N-isopropyl)-amilorid (100 μm) blocked net secretion by increasing mucoserosal flux, as did the addition of Na⁺ acetate (30 mm) to the mucosal solution. We conclude that the distal jejunum of the rat secretes K⁺ under short-circuit conditions. This secretion does not seem to occur via K⁺ channels, but through a pH dependent mechanism. Received: 16 February 1999/Revised: 29 June 1999     

Interaction of benthic microalgae and macrofauna in the control of benthic metabolism, nutrient fluxes and denitrification in a shallow sub-tropical coastal embayment (western Moreton Bay, Australia)

Benthic biogeochemistry and macrofauna were investigated six times over 1 year in a shallow sub-tropical embayment. Benthic fluxes of oxygen (annual mean ?918 μmol O₂ m^?2 h^?1), ammonium (NH₄ ⁺), nitrate (NO₃ ^?), dissolved organic nitrogen, dinitrogen gas (N₂), and dissolved inorganic phosphorus were positively related to OM supply (N mineralisation) and inversely related to benthic light (N assimilation). Ammonium (NH₄ ⁺), NO₃ ^? and N₂ fluxes (annual means +14.6, +15.9 and 44.6 μmol N m^?2 h^?1) accounted for 14, 16 and 53 % of the annual benthic N remineralisation respectively. Denitrification was dominated by coupled nitrification–denitrification throughout the study. Potential assimilation of nitrogen by benthic microalgae (BMA) accounted for between 1 and 30 % of remineralised N, and was greatest during winter when bottom light was higher. Macrofauna biomass tended to be highest at intermediate benthic respiration rates (?1,000 μmol O₂ m^?2 h^?1), and appeared to become limited as respiration increased above this point. While bioturbation did not significantly affect net fluxes, macrofauna biomass was correlated with increased light rates of NH₄ ⁺ flux which may have masked reductions in NH₄ ⁺ flux associated with BMA assimilation during the light. Peaks in net N₂ fluxes at intermediate respiration rates are suggested to be associated with the stimulation of potential denitrification sites due to bioturbation by burrowing macrofauna. NO₃ ^? fluxes suggest that nitrification was not significantly limited within respiration range measured during this study, however comparisons with other parts of Moreton Bay suggest that limitation of coupled nitrification–denitrification may occur in sub-tropical systems at respiration rates exceeding ?1,500 μmol O₂ m^?2 h^?1.     

Seasonal variations of CO2 and water vapour exchange rates over a temperate deciduous forest

Long-term and direct measurements of CO₂ and water vapour exchange are needed over forested ecosystems to determine their net annual fluxes of carbon dioxide and water. Such measurements are also needed to parameterize and test biogeochemical, ecological and hydrological assessment models. Responding to this need, eddy covariance measurements of CO₂ and water vapour were made ever a deciduous forest growing near Oak Ridge, TN, between April 1993 and April 1994. Periodic measurements were made of leaf area index, stomatal resistance, soil moisture and pre-dawn leaf water potential to characterize the gas exchange capacity of the canopy. Four factors had a disproportionate influence on the seasonal variation of CO₂ flux densities. These factors were photon flux densities (during the growing season), temperature (during the dormant season), leaf area index and the occurrence of drought The drought period occurred during the peak of the growing season and caused a significant decline in daily and hourly CO₂ flux densities, relative to observations over the stand when soil moisture was plentiful. The annual net uptake of carbon was calculated by integrating flux measurements and filling missing and spurious data with the relations obtained between measured CO₂ fluxes and environmental forcing variables. The net flux of carbon for the period between April 1993 and April 1994 was -525 g C m^?2 y^?1. This value represents a net flux of carbon from the atmosphere and into the forest. The net annual carbon exchange of this southern temperate broadleaved forest exceeded values measured over a northern temperate forest (which experiences a shorter growing season and has less leaf area) by 200 g C m^?2 y^?1 (cf. Wofsy et al 1993). The seasonal variation of canopy evaporation (latent heat flux) was controlled mostly by changes in leaf area and net radiation. A strong depression in evaporation rates was not observed during the drought Over a broadleaved forest large vapour pressure deficits promote evaporation and trees in a mixed stand are able to tap a variety of deep and shallow water sources.