Winter gas exchange between the atmosphere and snow-covered soils on Niwot Ridge,Colorado, USA

Background: There is a growing interest in understanding the gas exchange between the atmosphere and seasonally snow-covered regions, especially in light of projections that climate change will alter the timing and extent of seasonal snow cover. In snow-covered ecosystems, gas fluxes are due both to microbial activity in the snow-covered soils and to chemical and physical reactions with the various gases and/or dissolved constituents in the snowpack. Niwot Ridge, in the Colorado Rocky Mountains, has one of the most extensive sets of measurements of winter gas exchange globally.

Aims: Our goal was to examine the temporal patterns and environmental controls on Niwot Ridge of gas fluxes for gases with different sources and sinks.

Methods: Here, we review the concentrations and fluxes that have been measured for carbon dioxide, nitrous oxide, methane, nitrogen oxides, ozone, gaseous elemental mercury and volatile organic carbon compounds.

Results and Conclusions: We looked for similarities and differences among the gases, but in many cases, the origin, fate and controls of these fluxes still need to be determined. However, we believe that many of the biologically driven reactions are the result of exponential growth of a winter microbial community during the long period of stable environmental conditions under the seasonal snowpack.     

Fluxes of energy,water, and carbon dioxide from mountain ecosystems at Niwot Ridge,Colorado

Background: The eddy covariance (EC) technique provides a direct measure of water vapour and carbon dioxide fluxes between ecosystems and the atmosphere.

Aims: This review article highlights the findings of various studies that have integrated EC observations into basic meteorological, hydrological and ecological research questions in two ecosystems near Niwot Ridge, Colorado, and synthesises these studies into a catchment-scale model of water and carbon cycling, within the context of regional disturbance and environmental change.

Methods: EC was implemented continuously over subalpine forest and alpine tundra vegetation at Niwot Ridge, and resulting data were compared with discrete measurements and modelling studies.

Results: Sensible heat fluxes were generally in excess of latent heat fluxes, indicating that the forest and tundra ecosystems were moisture limited. Snow cover regulated the annual sum of primary productivity in the forest, and beneath-snow respiration represented a significant portion of ecosystem respiration at both locations.

Conclusions: Changes in the magnitude, timing, or spatial distribution of snow are likely to have the greatest impact on ecological processes in these semi-arid mountain catchments, but possibly in compensatory ways. Ultimately, the degree to which net carbon losses from alpine tundra offset forest carbon sequestration will determine the future magnitude of the Western United States carbon sink.     

Measuring Nitrogen and Phosphorus Uptake by Intact Roots of Mature Acer saccharum Marsh., Pinus resinosa Ait., and Picea abies (L.) Karst

A common method for measuring uptake by intact roots in situ is the depletion method, wherein intact fine roots are separated from soil and placed in nutrient solution. The difference between initial and final amounts of nutrient in solution is attributed to root uptake. Variations on this method include applying pretreatment solutions, training roots to grow into bags or trays, and varying concentrations of nutrient solution. We tested whether variations in methods affected measured net uptake rates of NH ₄ ⁺ , NO ₃ ⁻ , and PO ₄ ³⁻ . Intact roots of 60 year-old sugar maple (Acer saccharum Marsh.), red pine (Pinus resinosa Ait.), and Norway spruce (Picea abies (L.) Karst.) were given one of four treatments prior to measuring net uptake. “Trained” roots were grown in a sand-soil mixture. “Recovered” roots were excavated and allowed to recover in nutrient solution for two or four days (“two-day recovery” and “four-day recovery”, respectively). “No recovery” roots were excavated and used immediately in experiments. We exposed roots to three concentrations of nutrient solutions to observe the effects of initial nutrient solution concentration. Initial nutrient solution concentration was an important source of variation in measured uptake rates, and N uptake was stimulated by low antecedent concentrations. We found no significant differences in net uptake rates between pretreatments for any of the species studied, indicating that our pretreatments were not effective in improving measurement of uptake. Such pretreatments may not be necessary for measuring net uptake and may not hinder the comparison of rates measured using variations of the depletion method.     

Root growth,macro-nutrient uptake dynamics and soil fertility requirements of a high-yielding winter oilseed rape crop

Shoot growth, root growth and macro-nutrient uptake by a high-yielding (5t/ha grain) winter oilseed rape crop have been measured. Maximum rooting density in the top 20cm of soil was 9.4 cm cm⁻³ and roots reached a depth of at least 1.8 m. Maximum nutrient uptakes were 364 kg ha⁻¹ for N, 43 kg ha⁻¹ for P, 308 kg ha⁻¹ for K, 287 kg ha⁻¹ for Ca and 16 kg ha⁻¹ for Mg. A 30-day drought coincided with the flowering period and root and shoot growth, as well as nutrient uptake rates, were reduced. Nutrient concentrations in the soil solution necessary to sustain the nutrient fluxes into the root system by diffusive supply have been calculated. Peak values were in the range 10 μM for P to 87 μM for N, lower than the observed concentrations, and it was concluded that nutrient transport to roots was not a limitation to uptake by this rape crop.     

Spatial and seasonal variation in nutrient excretion by benthic invertebrates in a eutrophic reservoir

1. Nitrogen (N) and phosphorus (P) fluxes via excretion by benthic invertebrates were quantified in a eutrophic reservoir (Acton Lake, Ohio, U.S.A.). We quantified variation in nutrient fluxes seasonally (June until November 1997), spatially (three sites) and among taxa (chironomids, tubificid oligochaetes and Chaoborus). 2. The three taxa differed in spatial distribution and contribution to nutrient fluxes. Tubificids were the most abundant taxon at two oxic sites (1.5 and 4 m depth), and were exceedingly rare at an anoxic, hypolimnetic site (8 m). Chironomids were abundant only at the shallowest oxic site. Chaoborus was the only abundant taxon at the anoxic site. Total benthic invertebrate biomass was greatest at the shallowest site and lowest at the anoxic, hypolimnetic site. 3. Mass‐specific excretion rate [μmol NH₄–N or soluble reactive P (SRP) excreted mg dry mass^–1 h^–1] varied among experiments and was influenced by temperature. Differences among taxa were not significant. Thus, nutrient flux through benthic invertebrates was affected more by total invertebrate biomass and temperature than by species composition. 4. Fluxes of N and P via benthic invertebrate excretion (μmol NH₄–N or SRP m^–2 day^–1) were greatest at the oxic sites, where fluxes were dominated by the excretion of tubificids and chironomids. The N and P fluxes at the anoxic site were much lower, and were dominated by excretion by Chaoborus. The ratio of N and P excreted by the benthic invertebrate assemblage varied seasonally and was lowest at the anoxic site. 5. Comparison with other measured inputs shows that excretion by benthic invertebrates could be an important source of nutrients, especially of P. However, the relative importance of nutrient excretion by the benthos varies greatly spatially and temporally.     

Transpiration does not control growth and nutrient supply in the amphibious plant Mentha aquatica

Mentha aquatica L. was grown at different nutrient availabilities in water and in air at 60% RH. The plants were kept at 600 mmol m^?3 free CO₂ dissolved in water (40 times air equilibrium) and at 30 mmol m^?3 CO₂ in air to ensure CO₂ saturation of growth in both environments. We quantified the transpiration-independent water transport from root to shoot in submerged plants relative to the transpiration stream in emergent plants and tested the importance of transpiration in sustaining nutrient flux and shoot growth. The acropetal water flow was substantial in submerged Mentha aquatica, reaching 14% of the transpiration stream in emergent plants. The transpiration-independent mass flow of water from the roots, measured by means of tritiated water, was diverted to leaves and adventitious shoots in active growth. The plants grew well and at the same rates in water and air, but nutrient fluxes to the shoot were greater in plants grown in air than in those that were submerged when they were rooted in fertile sediments. Restricted O₂ supply to the roots of submerged plants may account for the smaller nutrient concentrations, though these exceeded the levels required to saturate growth. In hydroponics, the root medium was aerated and circulated between submerged and emergent plants to minimize differences in medium chemistry, and here the two growth forms behaved similarly and could fully exploit nutrient enrichment. It is concluded that the lack of transpiration from leaf surfaces in a vapour-saturated atmosphere, or under water, is not likely to constrain the transfer of nutrients from root to shoot in herbaceous plants. Nutrient deficiency under these environmental conditions is more likely to derive from restricted development and function of the roots in waterlogged anoxic soils or from low porewater concentrations of nutrients.     

Uncertainty in Nutrient Spiraling: Sensitivity of Spiraling Indices to Small Errors in Measured Nutrient Concentration

Abstract Proper assessment of ecological data must consider uncertainty. However, reported estimation of uncertainty in calculated values of nutrient spiraling indices is rare. Interpretations based on single values of spiraling indices, may therefore be unwittingly flawed. We investigated the sources of analytical uncertainty in the nutrient concentrations used to calculate two spiraling indices, uptake length (S _w ) and uptake velocity (V _f ), and used Monte Carlo Simulation (MCS) to estimate the resultant uncertainty in index values. We also examined the effect of the level of nutrient enrichment on the magnitude of index uncertainty. Outcomes under high and low nutrient uptake capacity were compared by performing nutrient addition experiments in two streams with contrasting ambient nutrient concentrations. We found that small differences (or uncertainties) in the average plateau nutrient concentration resulted in large uncertainties in spiraling indices. The uncertainty resulted from a combination of small differences in nutrient concentrations between upstream and downstream stations (particularly for the low uptake case), the low nutrient concentration added into the stream, and the sample matrix and storage. The stream with low nutrient uptake capacity had larger relative uncertainties in S _w than when the nutrient uptake capacity was high. The presence of such errors demands that S _w and V _f values should be reported with uncertainty, rather than the normal practice of a single calculated value. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

In situ endoscopy: New insights to root growth in biopores

Abstract

A flexible videoscope with an outer diameter of 3.8 mm and equipped with a movable tip was used for in situ observation of roots growing in larger‐sized biopores. We were able to display root morphology and position inside of biopores and differentiate four categories of root segments.     

Benthic fluxes in San Francisco Bay

Measurements of benthic fluxes have been made on four occasions between February 1980 and February 1981 at a channel station and a shoal station in South San Francisco Bay, using in situ flux chambers. On each occasion replicate measurements of easily measured substances such as radon, oxygen, ammonia, and silica showed a variability (±1) of 30% or more over distances of a few meters to tens of meters, presumably due to spatial heterogeneity in the benthic community. Fluxes of radon were greater at the shoal station than at the channel station because of greater macrofaunal irrigation at the former, but showed little seasonal variability at either station. At both stations fluxes of oxygen, carbon dioxide, ammonia, and silica were largest following the spring bloom. Fluxes measured during different seasons ranged over factors of 2–3, 3, 4–5, and 3–10 (respectively), due to variations in phytoplankton productivity and temperature. Fluxes of oxygen and carbon dioxide were greater at the shoal station than at the channel station because the net phytoplankton productivity is greater there and the organic matter produced must be rapidly incorporated in the sediment column. Fluxes of silica were greater at the shoal station, probably because of the greater irrigation rates there. N + N (nitrate + nitrite) fluxes were variable in magnitude and in sign. Phosphate fluxes were too small to measure accurately. Alkalinity fluxes were similar at the two stations and are attributed primarily to carbonate dissolution at the shoal station and to sulfate reduction at the channel station. The estimated average fluxes into South Bay, based on results from these two stations over the course of a year, are (in mmol m^–2 d^–1): O₂ = –27 ± 6; TCO₂ = 23 ± 6; Alkalinity = 9 ± 2; N + N = –0.3 ± 0.5; NH₃ = 1.4 ± 0.2; PO₄ = 0.1 ± 0.4; Si = 5.6 ± 1.1. These fluxes are comparable in magnitude to those in other temperate estuaries with similar productivity, although the seasonal variability is smaller, probably because the annual temperature range in San Francisco Bay is smaller.Budgets constructed for South San Francisco Bay show that large fractions of the net annual productivity of carbon (about 90%) and silica (about 65%) are recycled by the benthos. Substantial rates of simultaneous nitrification and denitrification must occur in shoal areas, apparently resulting in conversion to N₂ of 55% of the particulate nitrogen reaching the sediments. In shoal areas, benthic fluxes can replace the water column standing stocks of ammonia in 2–6 days and silica in 17–34 days, indicating the importance of benthic fluxes in the maintenance of productivity.Pore water profiles of nutrients and Rn-222 show that macrofaunal irrigation is extremely important in transport of silica, ammonia, and alkalinity. Calculations of benthic fluxes from these profiles are less accurate, but yield results consistent with chamber measurements and indicate that most of the NH₃, SiO₂, and alkalinity fluxes are sustained by reactions occurring throughout the upper 20–40 cm of the sediment column. In contrast, O₂, CO₂, and N + N fluxes must be dominated by reactions occurring within the upper one cm of the sediment-water interface. While most data support the statements made above, a few flux measurements are contradictory and demonstrate the complexity of benthic exchange.     

Comparison of nutrient uptake between three‐dimensional simulation and an averaged root system model

Abstract

We present a new numerical approach describing nutrient uptake in three dimensions. Dynamic boundary conditions are considered at the individual root surfaces within a root system. As an example, we compare the three‐dimensional simulation results of phosphate uptake by a young maize root system to the corresponding effective solution. We show that the two solutions are similar concerning phosphate uptake and the size of the depletion zones. The presented approach makes it possible to verify simplifications that are made in the development of effective models. Furthermore, it is possible to extend existing models by including spatial heterogeneities that will increase our understanding of rhizosphere processes.     

Environmental controls of temporal and spatial variability in CO2 and CH4 fluxes in a neotropical peatland

Tropical peatlands play an important role in the global storage and cycling of carbon (C) but information on carbon dioxide (CO₂) and methane (CH₄) fluxes from these systems is sparse, particularly in the Neotropics. We quantified short and long‐term temporal and small scale spatial variation in CO₂ and CH₄ fluxes from three contrasting vegetation communities in a domed ombrotrophic peatland in Panama. There was significant variation in CO₂ fluxes among vegetation communities in the order Campnosperma panamensis > Raphia taedigera > Cyperus. There was no consistent variation among sites and no discernible seasonal pattern of CH₄ flux despite the considerable range of values recorded (e.g. ?1.0 to 12.6 mg m^?2 h^?1 in 2007). CO₂ fluxes varied seasonally in 2007, being greatest in drier periods (300–400 mg m^?2 h^?1) and lowest during the wet period (60–132 mg m^?2 h^?1) while very high emissions were found during the 2009 wet period, suggesting that peak CO₂ fluxes may occur following both low and high rainfall. In contrast, only weak relationships between CH₄ flux and rainfall (positive at the C. panamensis site) and solar radiation (negative at the C. panamensis and Cyperus sites) was found. CO₂ fluxes showed a diurnal pattern across sites and at the Cyperus sp. site CO₂ and CH₄ fluxes were positively correlated. The amount of dissolved carbon and nutrients were strong predictors of small scale within‐site variability in gas release but the effect was site‐specific. We conclude that (i) temporal variability in CO₂ was greater than variation among vegetation communities; (ii) rainfall may be a good predictor of CO₂ emissions from tropical peatlands but temporal variation in CH₄ does not follow seasonal rainfall patterns; and (iii) diurnal variation in CO₂ fluxes across different vegetation communities can be described by a Fourier model.     

Natural and man-caused factors affecting the abundance and cycling of dissolved organic substances in precambrian shield lakes

A study on nutrient regeneration processes and a measure of their fluxes at the sediment-water interface was carried out in two different stations of a shallow lagoon of the Po delta river (Italy). A few parameters on the solid fraction (grain-size, porosity, C, N) and pore water profiles of o-P, NH₃, NO _inf3 ^sup– , SiO₂, Tot-CO₂, SO _inf4 ^sup2– , Fe, Mn, Ca, Mg, pH, Eh were determined. At both stations the results were typical for fine sediments rich in organic matter. The ratio of variations of sulphate (SO _inf4 ^sup2– ) to total carbonate demonstrates the main role sulphate reduction plays on the organic matter decay. The use of the ratios of variations of sulphate (SO _inf4 ^sup2– ) to ammonia (NH₃) and of sulphate (SO _inf4 ^sup2– ) to phosphate (o-P) in pore waters enabled us to calculate the C/N/P of the decomposing organic matter. Obtained C/N/P indicated an enrichment of N and P with regard to C/N/P ratios of the solid fraction, due to the selective stripping of N and P during organic matter mineralization. This phenomenon decreases with depth, where organic matter becomes more refractory. Calculations on saturation degrees have shown the possibility of authigenic calcite, apatite and rhodochrosite precipitation in sediments. Nutrient fluxes were estimated for SiO₂, NH₃ and o-P by means of benthic chambers and modelling the pore water profiles. The model used for the calculation of fluxes allowed us to account for the bioturbation-irrigation influence near the interface, by means of a cumulative diffusion coefficient. Directly measured fluxes proved to be always significantly greater than the theoretical ones. These differences seem to be due to surface processes which do not affect pore water concentrations (degradation of fresh materials at the interface; micro-bioturbation by small gasteropoda such as Hydrobia ventrosa) and/or to the different concept of the two methods in time and space. Number, size and biomass of macrobenthic species living in the sediment underneath the benthic chambers were determined. The comparison between data on macrobenthic populations and flux values showed a good relationship between the number of organisms and benthic fluxes within each station. However, flux variations between stations are to be attributed mainly to the different arrangement of the tubes of the polychaetes Polydora ciliata in the sediment.     

Nutrient pools and fluxes of the ground vegetation in coniferous forests due to fertilizing,liming and amelioration

The effects of fertilization and amelioration treatments on some nutrient pools and fluxes of ground vegetation in mature pine and spruce stands on acid soils in South Germany are described. In N-limited pine forests with moderate canopy density and with Deschampsia flexuosa an additional N-accumulation in biomass of 20–40 kg ha^-1 occurred 3 years after pure N-fertilization. The N, P, K-cycling through ground vegetation was stimulated more than 10 years by a combined N + CaCO₃ + P treatment leading toa shift in dominance from cryptogams and Ericaceae towards Deschampsia flexuosa and ruderal species like Epilobium angustifolium. The effect of a lupine treatment (combined with initial soil preparation, liming and P supply) was far stronger than the effect of the other experimental procedures. But the fertilizer and amelioration effects on the herb layer of pine forests tended to decline after two decades for different reasons.The shade-tolerant ground vegetation in a nitrogen-saturated spruce forest was not able to prevent heavy additional nitrate losses from upper mineral soil after dolomitic liming. But the Ca, Mg and K fluxes through ground vegetation were strongly elevated in the third year after treatment.     

OLISTHODISCUS LUTEUS (CHRYSOPHYCEAE). III. UPTAKE AND UTILIZATION OF NITROGEN AND PHOSPHORUS1

Uptake and assimilation of nitrogen and phosphorus were studied in Olisthodiscus luteus Carter. A diel periodicity in nitrate reductase activity was observed in log and stationary phase cultures; there was a 10-fold difference in magnitude between maximum and minimum rates, but other cellular features such as chlorophyll a, carbon, nitrogen, C:N ratio (atoms) · cell^?1 were less variable. K_s values (~2 μM) for uptake of nitrate-N and ammonium-N were observed. Phosphorus assimilated · cell^?1· day^?1 varied with declining external phosphorus concentrations; growth rates <0.5 divisions · day^?1 were common at <0.5 μM PO_4-P. Phosphate uptake rates (K_s= 1.0–1.98 μM) varied with culture age and showed multiphasic kinetic features. Alkaline phosphatase activity was not detected. Comparisons of the nutrient dynamics of O. luteus to other phytoplankton species and the ecological implications as related to the phytoplankton community of Narragansett Bay (Rhode Island) are discussed.     

Nitrogen Cycling and Mass Balance for a Forested Catchment in the Italian Alps. Assessment of Nitrogen Status

During 1999–2001 the chemical composition and fluxes were measured in rainfall, throughfall, soil solution and stream water in a remote forested site in the Italian Alps. The analysis of temporal patterns revealed the differential behaviour of nitrogen and sulphur and suggested that different mechanisms controlled their flux. No important changes in sulphate concentration and fluxes emerged as the solution passed through the various components of the forest ecosystem, and temporal variations of SO₄ in the soil solution and stream were likely driven by the physical process of dilution. The availability of nitrate and ammonia, by contrast, was drastically reduced as throughfall water entered the soil and passed through the mineral layers, irrespective of season. The calculated hydrochemical budget based on throughfall and soil solution N fluxes revealed that ~80% N retention in the forest soil, corresponding to 12 kg ha⁻¹ yr⁻¹, despite a relatively high N deposition loading (15 kg ha⁻¹ yr⁻¹). Most of the leached nitrogen (90%) was in the organic form. Indicators of the N status of this ecosystem, such as C/N ratio in solid and solution phase of the soil and N foliage content as well as land use history were examined. Despite the strong N retention in the forested part of the catchment, the stream water N–NO₃ levels were consistently above 10 μg l⁻¹ suggesting that the Val Masino catchment as a whole was less efficient in processing atmospheric N inputs. This contrasting N behaviour illustrates the role of landscape features, such as the soil cover and vegetation type, that is characteristic of an alpine catchment.     

Nutrient (N,P, Si) fluxes between marine sediments and water column in coastal and open Adriatic

Nutrient benthic fluxes, as well as sediment phosphorus concentration at the open sea and coastal water stations of the Central and South Adriatic were studied during 1997–98. The fluxes were in the ranges: 0.16–2.67 mmol m^–2 d^–1 (silicate); –0.031–0.164 mmol m^–2 d^–1 (phosphate); –0.51–2.03 mmol m^–2 d^–1 (ammonia); and –1.32–1.62 mmol m^–2 d^–1 (nitrate + nitrite). Silicate flux showed a gradient from the coastal area to the open sea. Ammonia was the main nitrogen species in the flux at the estuary and bay stations, while the sum of nitrate and nitrite was predominant at the open sea stations. Relationships between phosphate and ammonia fluxes (r = 0.699, p<0.01) as well as phosphate and silicate (r = 0.529, p<0.01) were established.     

Temporal patterns of inorganic nitrogen uptake by mature sugar maple (Acer saccharum Marsh.) and red spruce (Picea rubens Sarg.) trees using two common approaches

Background: Plant uptake of nitrogen influences many ecosystem processes, yet uptake by trees in northern forests of the United States has not been quantified throughout the growing season.

Aims: To measure NH₄ ⁺ and NO₃ ^? uptake by mature sugar maple (Acer saccharum) and red spruce (Picea rubens) trees during the early, mid and late growing season.

Methods: At Hubbard Brook Experimental Forest, New Hampshire, we used two approaches to measure nitrogen uptake capacity by mature trees: an in situ depletion method using intact roots and an ex situ ¹⁵N tracer method using excised roots.

Results: NH₄ ⁺ uptake was greater than NO₃ ^? for both methods and tree species (P < 0.05). NH₄ ⁺ uptake was lowest during the early growing season, while NO₃ ^? uptake was lowest during the late growing season. Measured rates of NH₄ ⁺ uptake were 2–3 orders of magnitude greater using the in situ depletion method compared with the ex situ ¹⁵N tracer method.

Conclusions: These results demonstrate seasonal differences in nitrogen uptake by two dominant tree species in a northern forest and show that the method employed can significantly impact measured rates of uptake, which could have implications for understanding the magnitude of plant nitrogen uptake and for cross-study comparisons of this process.     

Xylem sap composition: A tool for investigating mineral uptake and cycling in adult spruce

Xylem sap composition of spruce is influenced by several factors, such as the sampled organ, the sampling period, the availability of soil nutrients, and the soil water potential. Based on literature data and ongoing investigations carried out with adult trees, we present an overview on the main factors influencing xylem sap concentrations of Norway spruce. Direct measurements of nutrient fluxes in the xylem sap are then used to suggest a general scheme of mineral element cycling within adult trees. In Norway spruce (Picea abies Karst.), nutrient concentration in the xylem sap was higher in twigs and fine roots compared to the bottom of the trunk, the highest concentrations beeing observed in spring during the shoot elongation. Xylem sap concentrations were higher in spruce growing at nutrient rich sites than at poor sites. The combination of twig and trunk xylem sap analysis, together with xylem flow measurements in the trunk during the course of a vegetation period allowed the quantification of mineral fluxes via xylem sap flow in the trunk and twigs. These results were compared to gross mineral uptake measurements at the same site. Ca flux in the trunk xylem sap was lower than the gross uptake of Ca. Mg flux in trunk sap was approximately equivalent to Mg gross uptake whereas P and K fluxes in trunk sap were much higher than the gross uptake. Fluxes of Ca, Mg, K and P in the twig sap were much higher than that in trunk sap. Data suggest that internal cycling is responsible for a large part of the nutrient fluxes in the xylem sap of the crown. Xylem sap composition thus appears to be a tool which can complement other sources of information on mineral uptake and cycling in adult spruce     

大气氮沉降对森林土壤甲烷吸收和氧化亚氮排放的影响及其微生物学机制

水分非饱和的森林土壤是大气甲烷(CH4)汇和氧化亚氮(N2O)源,大气氮沉降增加是导致森林土壤碳氮气体通量不平衡的主要原因之一。土壤CH4吸收和N2O排放之间存在协同、消长和随机等复杂的耦合关系,关于氮素对两者产生过程的调节作用以及内在的微生物学机制至今尚不完全清楚。综述了森林土壤CH4吸收和N2O排放耦合过程的理论基础,土壤CH4和N2O的产生与消耗过程对增氮响应的生物化学和微生物学机制,指出各研究领域的不足和未来的研究重点。总体而言,低氮倾向于促进贫氮森林土壤CH4吸收,不改变土壤N2O的排放,而高氮显著抑制富氮森林土壤CH4吸收以及促进N2O排放。外源性氮素通过竞争抑制和毒性抑制来调控森林土壤CH4的吸收,而通过促进土壤硝化和反硝化过程来增加N2O的排放。然而,由于全球氮沉降控制试验网络分布的不均匀性、土壤碳氮通量产生过程的复杂性以及微生物分子生态学方法的局限性等原因,导致氮素对森林土壤碳氮通量的调控机制研究一直进展缓慢,未能将微生物功能群落动态与土壤碳氮通量真正地联系起来。未来研究应该从流域、生态系统和分子尺度上深入探讨土壤碳氮通量耦合作用的环境驱动机制,氮素对土壤CH4氧化和N2O产生过程的调控作用,以及增氮对土壤甲烷氧化菌和N2O产生菌活性和群落组成的影响。     

Microbial enzyme activity,nutrient uptake and nutrient limitation in forested streams

1. We measured NH₄⁺ and PO₄^?3 uptake length (S_w), uptake velocity (V_f), uptake rate (U), biofilm respiration and enzyme activity and channel geomorphology in streams draining forested catchments in the northwestern (Northern California Coast Range and Cascade Mountains) and southeastern (Appalachian and Ouachita mountains) regions of the United States. Our goal was to use measures of biofilm enzyme activity and nutrient uptake to assess nutrient limitation in forested streams across broad regional scales. 2. Geomorphological attributes, biofilm enzyme activity and NH₄⁺ uptake were significantly different among streams in the four study units. There was no study unit effect on PO₄^?3 uptake. The proportion of the stream channel in pools, % woody debris, % canopy closure, median substrate size (d₅₀), stream width (w), stream velocity (v), discharge (Q), dispersion coefficient (D) and transient storage (A_s/A) were correlated with biofilm enzyme activity and nutrient uptake in some study units. 3. Canonical correlation analyses across study units revealed significant correlations of NH₄‐V_f and PO₄‐V_f with geomorphological attributes (w, d₅₀, D, % woody debris, channel slope and % pools) and biofilm phosphatase activity. 4. The results did not support our expectation that carbon processing rates by biofilm microbial assemblages would be governed by stream nutrient availability or that resulting biofilm enzyme activity would be an indicator of nutrient uptake. However, the relative abundances of peptidases, phosphatase and glycosidases did yield insight into potential N‐, P‐ and C‐limitation of stream biofilm assemblages, and our use of biofilm enzyme activity represents a novel application for understanding nutrient limitations in forested streams. 5. Regressions of V_f and U against ambient NH₄⁺ and PO₄^?3 indicated that none of our study streams was either NH₄⁺ or PO₄^?3 saturated. The Appalachian, Ouachita and Coastal streams showed evidence of NH₄⁺ limitation; the Ouachita and Coastal streams were PO₄^?3 limited. As a correlate of nutrient limitation and saturation in streams, ratios of total aminopeptidase and phosphatase activities and the ratio of NH₄‐U to PO₄‐U indicate these forested streams are predominantly N‐limited, with only the streams draining Ouachita and Coastal catchments demonstrating appreciable levels of P‐limitation. 6. Our results comparing the stoichiometry of microbial enzyme activity with nutrient uptake ratios and with the molar ratios N and P in stream waters suggest that biological limitations are not strictly the result of stream chemistry and that the assessments of nutrient limitations in stream ecosystems should not be based on chemistry alone. 7. Our present study, along with previous work in streams, rivers and wetlands, suggests that microbial enzyme activities, especially the ratios of total peptidases to phosphatase, are useful indicators of nutrient limitations in aquatic ecosystems.