Extracellular dissolved organic carbon released from phytoplankton as a source of carbon for heterotrophic bacteria in lakes of different humic content

The quantitative importance of photosynthetically produced dissolved organic carbon (PDOC) released from phytoplankton as a source of carbon for pelagic, heterotrophic bacteria was investigated in four temperate Swedish lakes, of which two had low (≈20 mg Pt 1⁻¹), and two moderately high (60–80 mg Pt 1⁻¹) humic content. The bacterial assimilation of PDOC was estimated with the ¹⁴C method, and the total production of the heterotrophic bacteria was estimated with the [3H]thymidine incorporation method. The release of PDOC from natural communities of phytoplankton was not restricted to periods of photosynthesis, but often continued during periods of darkness. Heterotrophic bacteria often assimilated the labile components of the PDOC at high rates (up to 73% of the released PDOC was assimilated during the incubation in our experiments). The contribution of PDOC to bacterial production exhibited large within-lake seasonal variations, but PDOC was at certain times, both in humic and non-humic lakes, a quantitatively very important carbon source for the heterotrophic bacteria. Under periods of comparatively low primary production, heterotrophic bacteria in humic lakes appear to utilize allochthonous, humic substances as a substrate.     

Dissolved organic matter quantity and quality from freshwater and saltwater lakes in east-central Alberta

Concentrations of dissolved organic matter (DOM) in surface waters of sub-humid to semi-arid lakes in east-central Alberta increase with increasing salinity and water residence time from about 20 to 330 mg L^–1 as dissolved organic carbon (DOC). This pattern is opposite to that observed among freshwater lakes spanning a gradient in water residence times, and is probably caused by evaporative concentration of refractory DOM. The proportion of total DOC, operationally defined as humic substances using XAD-8 resin, was high, though similar to surface waters typically referred to as "humic", and independent of salinity. Very long water residence times (hundreds of years) in saline lakes favors evapoconcentration of low-color, low molecular weight DOM, with N-content characteristic of allochthonous DOM.     

富营养化湖泊溶解性有机碳生物可利用性研究进展

富营养化湖泊溶解性有机碳(DOC)包括内源和外源性碳源,不同来源碳源在物质化学结构组成和分子量级等方面具有显著差异,进而影响到对细菌的生物可利用性和碳素在食物网中的传递效率。根据国内外文献,综述了内外源DOC在碳稳定同位素值域上的显著差异,建议通过对DOC碳稳定同位素的分析来识别富营养化湖泊中DOC的主要来源;通过对比内外源DOC在碳水化合物、结合态中性糖和腐殖质含量上的差异,并结合细菌生长参数如细菌二级生产力、细菌呼吸作用及细菌生长效率来分析内外源DOC对细菌的生物可利用性。从富营养化湖泊DOC来源的角度探讨其生物可利用性和碳素传递效率,有助于了解富营养化湖泊食物网中碳素循环特征,加强对湖泊生态学的认识,为湖泊环境治理与保护提供科学依据。     

Microbial control of dissolved organic carbon in lakes: research for the future

The dissolved organic carbon (DOC) of lakes dominates any budget of organic carbon in these systems. Limnologists are still limited by techniques and particularly by the lack of measurements of rates of microbial transformation and use of this DOC. There are now four different approaches to the study of the microbial control of DOC in lakes. The first is through measurements of the total DOC. Recent advances in measurement with high temperature combustion will likely lead to higher and more consistent measurements in freshwaters than previously. It is possible that a biologically active fraction may be identified. The second approach is through measurements of microbial incorporation and respiration of ¹⁴C-labeled organic matter. The kinetics of this process are well known but advances in measurement of the size of the substrate pool are still being made. The third approach is to use bacterial growth in batch or continuous flow experiments in order to understand how much of the total DOC can be decomposed by microbes. The assay in this approach may be microbial growth (thymidine incorporation, biomass changes) or change in the DOC (total concentrations, specific compounds, or fractions of the DOC by molecular weight). These methods are promising but are not developed enough for routine use. For example, growth measurements in the laboratory are all subject to experimental artifacts caused by changes in the DOC and in the bacterial populations. Finally, the fourth approach is through the use of isotopes of the natural DOC. In the sea this approach has given the age of the bulk DOC (¹⁴C data). In freshwaters it has great potential for differentiating between bacterial use of terrestrial DOC vs. use of algal-derived DOC (13C data). Stable isotopes are also useful for experimentally labeling DOC produced by algae and following the use of this material by bacteria.     

Climate-induced changes in the dissolved organic carbon budgets of boreal lakes

During 20 years of climatic warming, drought and increased forest firesbetween 1970 and 1990, DOC concentrations declined by 15--25%in lakesof the Experimental Lakes Area, northwestern Ontario, allowing increasedpenetration of both UV and photosynthetically-active radiation (PAR), andcausing deeper euphotic zones and thermoclines. Decreased input to thelakes of DOC from terrestrial catchments and upstream lakes was theprimary reason for the decline, although in-lake removal also increasedslightly. Decreased streamflow caused by drought was more important thanforest fires in affecting DOC exports from catchments. Experimentalacidification of lakes caused even greater losses in DOC, by enhancing ratesof in-lake removal. DOC in Lake 302S, acidified to pH 4.5 during the1980s, declined to less than 10% of preacidificationvalues.     

海洋异养细菌利用溶解有机碳的定量评估

【目的】惰性溶解有机碳(refractory dissolved organic carbon,RDOC)是海洋总有机碳的主体组分,RDOC在深海中可保存数千年,构成了巨大的碳储库,在调节气候变化中有重要作用。但RDOC的定量评估尚未有统一的标准方法。通过测定环境中能被异养细菌利用的溶解有机碳(biodegradable DOC,BDOC)可以反过来评估RDOC的量。本文对BDOC测定中一些关键步骤进行验证,为制定海洋RDOC评估标准奠定基础。【方法】本文评估了3种过滤方式及5种滤膜对DOC测定的影响,并评估了瓶子效应和稀释效应对细菌生长和DOC利用的影响。【结果】研究发现,(1) GF/F滤膜、GF-75滤膜、聚四氟乙烯(PTFE)滤膜(孔径0.2μm)、聚碳酸酯(PC)滤膜(孔径0.2μm)和聚四氟乙烯材质针孔过滤器(HA)(孔径0.2μm) 5种滤膜不会引入DOC污染;抽滤过滤和重力过滤方式过滤效果稳定、无污染,而在线过滤效果不稳定,易污染;(2)不同大小培养体系(30–480 mL;表面积/体积比为:1.64–0.67 cm–1)之间的细菌生长速率和DOC利用量没有显著性差异;(3)培养体系稀释度越高,细菌生长速率越高,对数生长期细菌丰度及DOC利用量越低。【结论】综合考虑,建议BDOC和RDOC测定实验中采用抽滤过滤的方式及不进行稀释的培养体系;常用的滤膜和培养体积对BDOC评估无显著影响。结合研究结果,我们提出了评估海洋RDOC的方法。     

Effects of different molecular weight fractions of dissolved organic matter on the growth of bacteria,algae and protozoa from a highly humic lake

Effects of different molecular size fractions (< 1000 MW, < 10 000 MW, < 100 000 MW and <0.1 μm) of dissolved organic matter (DOM) on the growth of bacteria, algae and protozoa from a highly humic lake were investigated. DOM from catchment drainage water as well as from the lake consisted mostly (59–63%) of high molecular weight (HMW) compounds (> 10 000 MW). With excess inorganic nutrients, the growth rate and yield of bacteria were almost identical in all size fractions. However, in < 1000 MW fractions and with glucose added, a longer lag phase occurred. Without added nutrients both the growth rates and biomasses of bacteria decreased towards the smaller size fractions and the percentage of dissolved organic carbon (DOC) used during the experiment and the growth efficiency of bacteria were lower than with excess nutrients. The growth efficiency of bacteria was estimated to vary between 3–66% in different MW fractions, largely depending on the nutrient concentrations, but the highest growth efficiencies were observed in HMW fractions and with glucose. The growth of algae was clearly lowest in the < 1000 MW fraction. In dim light no net growth of algae could be found. In contrast, added nutrients substantially enhanced algal growth and in deionized water with glucose, algae achieved almost the same growth rate and biomass as in higher MW fractions of DOM. The results suggested that bacteria and some algae were favoured by DOM, but protozoans seemed to benefit only indirectly, through bacterial grazing. The utilization of DOM by bacteria and algae was strongly affected by the availability of phosphorus and nitrogen.     

The relation between bacterial carbon and dissolved humic compounds in oligotrophic lakes

Abstract The mean annual biomass of planktonic bacteria showed large variations both within and between lakes. The lowest bacterial biomass was found in acidified lakes (7.8–12.1 μg C · 1⁻¹), and tended to increase with increasing water colour (up to 44.1 μg C · 1⁻¹). The highest recorded bacterial biomass was 138 μg C · 1⁻¹. The mean annual bacterial biomass equalled 23–45% of the algal biomass. Zooplankton biomass was high, compared to algal biomass (40–50%). Multiple regression analysis of 10 variables showed a strong positive correlation between bacterial biomass and humic content ( r = 0.74, P < 0.001), while other parameters, except pH, showed no correlation. The observation thus strongly supports the role of humic compounds in aquatic secondary production.     

Hydrologic control of dissolved organic matter in low-order Precambrian Shield lakes

Concentrations of dissolved organic carbon (DOC) and color were measuredas a function of time in enclosures and lakes at the Experimental Lakes Area,to calculate their net loss rates. Loss rates in enclosures werefirst order for both DOC and color, with half-times for loss of 166and 122 d, respectively. Thus, the colored, light-attenuating componentof the DOC pool is lost from water more rapidly than is bulk DOC. Loss ratesin lakes, calculated from a steady state model, were similar to values forcolor in enclosures, but for DOC in lakes were four times slower than inenclosures. In lakes, loss rate for DOC increased rapidly with decreasingwater residence time (_w) but was independent of_w when it was greater than 3 years. In lakes, theloss rate for color was independent of water residence time. Thedifference in losses of DOC and color between lakes and enclosurescould be from release of low-color DOC from sediments.     

Acidity controls on dissolved organic carbon mobility in organic soils

Dissolved organic carbon (DOC) concentrations in surface waters have increased across much of Europe and North America, with implications for the terrestrial carbon balance, aquatic ecosystem functioning, water treatment costs and human health. Over the past decade, many hypotheses have been put forward to explain this phenomenon, from changing climate and land management to eutrophication and acid deposition. Resolution of this debate has been hindered by a reliance on correlative analyses of time series data, and a lack of robust experimental testing of proposed mechanisms. In a 4 year, four‐site replicated field experiment involving both acidifying and deacidifying treatments, we tested the hypothesis that DOC leaching was previously suppressed by high levels of soil acidity in peat and organo‐mineral soils, and therefore that observed DOC increases a consequence of decreasing soil acidity. We observed a consistent, positive relationship between DOC and acidity change at all sites. Responses were described by similar hyperbolic relationships between standardized changes in DOC and hydrogen ion concentrations at all sites, suggesting potentially general applicability. These relationships explained a substantial proportion of observed changes in peak DOC concentrations in nearby monitoring streams, and application to a UK‐wide upland soil pH dataset suggests that recovery from acidification alone could have led to soil solution DOC increases in the range 46–126% by habitat type since 1978. Our findings raise the possibility that changing soil acidity may have wider impacts on ecosystem carbon balances. Decreasing sulphur deposition may be accelerating terrestrial carbon loss, and returning surface waters to a natural, high‐DOC condition.     

The effect of humic substances on the toxicity of aluminium to adult rainbow trout, Oncorhynchus mykiss (Walbaum)

Some physiological parameters were measured in adult rainbow trout during a 10-day exposure to 180 μg Al_total l⁻¹ in acid water (pH 4.7) with or without humic substances (10 mg l⁻). The fish were acclimatized to pH 5.0 for 7 days prior to the experimental treatments.
Chemical analyses revealed that, in the presence of human substances, 74–80% of the A1 was organic bound, while in the absence of humic substances most of the Al(987percnt;) occurred in the inorganic form.
Al bound to humic substances (13–150 μg l⁻¹) did not alter the plasma NaCl-concentration, nor the haematocrit value, of rainbow trout during an exposure period of 10 days. This contrasts with the high death rate obtained within 2–3 days when most of the A1 (175 μg l⁻¹) was in the inorganic form. The lethality was accompanied by a 25% decrease in the plasmaconcentration of NaCl and a doubling of the haematocrit value. Bulk analysis revealed that when the metal was present in inorganic forms the total Al content of the gills (75 μg A1 g⁻¹ wet weight) was 15 times higher than when it was present as bound to the humic substances. These experiments showed that the accumulation of A1 at the gills was accompanied by physiological disturbances, both being a function of the chemical speciation of Al.     

土壤溶解性有机碳在陆地生态系统碳循环中的作用

土壤溶解性有机碳(DOC)是有机碳库的活跃组分,在陆地生态系统碳循环中发挥重要作用.本文从碳循环重要性着手,综述了土壤DOC在土壤碳固持与温室气体排放中的作用;结合我国的现实情况(如土壤酸化、气候变暖等),探讨了土壤DOC的相关影响因素如土壤性质、环境因素、人为活动对土壤DOC的影响及作用机制,对进一步理解土壤DOC在陆地生态系统碳循环与温室气体减排中的作用具有重要意义.     

Dynamics of dissolved and particulate organic carbon in a saline and semiarid stream of southeast Spain (Chicamo stream)

Annual variations in the concentration of dissolved (DOC) and particulate organic carbon (CPOC = Coarse; FPOC = Fine; UPOC = Ultrafine) were studied in a 100 m-reach of the Chicamo stream, an intermittent saline stream in southeast Spain. DOC represented the most important fraction of organic carbon flowing in the Chicamo stream (>98%), with concentrations of about 1.7 mgC l^–1 during most of the year, reaching 2.5 mgC l^–1 in summer. One high flow episode during a rain storm in winter was characterized by a considerably increased concentration of DOC (9.4 mgC l^–1). CPOC was the dominant POC fraction. Positive and significant correlations were found for DOC and discharge, which support the idea of allochthonous inputs due to floods. There was no significant correlation between POC and discharge. No significant correlations were found for DOC or POC with the physico-chemical parameters measured, while a negative significant correlation was found between DOC and temperature. The export of total organic carbon from the drainage basin of the Chicamo stream was low (6.2 × 10^–4 gC m^–2 yr^–1) and typical of streams in arid and semi-arid regions. The results of a Principal Component Analysis defined three different phases. The first consisted of short periods, during which floods provide pulses of allochthonous organic carbon and nutrients, the second a dry phase (summer), defined by biotic interactions, during which the stream could acts as a `sink' of organic matter, and the third and final phase which is characterised by hydrological stability.     

Concentration and composition of dissolved organic carbon in streams in relation to catchment soil properties

Two adjacent catchments in the Otway Ranges of Victoria, Australia (Redwater and Clearwater) produce water with markedly different concentrations of dissolved organic carbon (DOC) during summer. Water from Redwater Creek had a DOC concentration of 32 mg L^–1, while water from Clearwater Creek had a DOC concentration of 3.8 mg L^–1. Examination of the catchments revealed that while climate, topography, vegetation and land use were similar, the soils were different. The objective of this study was to examine the relationship between the concentration and chemical composition of DOC in stream waters and the nature of soils in the two catchments. Soil mapping determined that clayey soils formed on Cretaceous sediments (Cretaceous soils) occurred throughout both catchments, but that Redwater Catchment also contained a large area (39%) of sandy soils formed on Tertiary sediments (Tertiary soils). The concentration of DOC in forest floor leachate was high in both the Tertiary and Cretaceous areas; however, the concentration of DOC in water draining areas dominated by Tertiary soils was greater than that in water draining areas dominated by Cretaceous soils. Laboratory experiments showed that the Cretaceous soils had higher adsorption capacities for forest floor leachate DOC than the Tertiary soils. The difference in DOC concentrations of the streams was therefore attributed to the difference in adsorption capacity of catchment soils for DOC. Adsorption capacities of the soils were found to be a function of their clay contents and specific surface areas.Solid-state³C nuclear magnetic resonance spectroscopy and pyrolysis-mass spectrometry were used to determine the chemical structure of DOC found in streams and forest floor leachate samples and that remaining in solution after interaction with soil. Chemistry of DOC in forest floor leachate was similar before and after interaction with soil, indicating no preferential adsorption of a particular type of carbon. Thus, differences between the chemical structure of stream DOC and forest floor leachate DOC could be attributed to microbial modifications during its movement through soils and into the streams, rather than losses by adsorption.     

The effect of land use on dissolved organic carbon and nitrogen uptake in streams

1. Agricultural and urban land use may increase dissolved inorganic nitrogen (DIN) concentrations in streams and saturate biotic nutrient demand, but less is known about their impacts on the cycling of organic nutrients. To assess these impacts we compared the uptake of DIN (as ammonium, NH₄⁺), dissolved organic carbon (DOC, as acetate), and dissolved organic nitrogen (DON, as glycine) in 18 low‐gradient headwater streams in southwest Michigan draining forested, agricultural, or urban land‐use types. Over 3 years, we quantified uptake in two streams in each of the three land‐use types during three seasons (spring, summer and autumn). 2. We found significantly higher NH₄⁺ demand (expressed as uptake velocity, V_f) in urban compared to forested streams and NH₄⁺V_f was greater in spring compared to summer and autumn. Acetate V_f was significantly higher than NH₄⁺ and glycine V_f, but neither acetate nor glycine V_f were influenced by land‐use type or season. 3. We examined the interaction between NH₄⁺ and acetate demand by comparing simultaneous short‐term releases of both solutes to releases of each solute individually. Acetate V_f did not change during the simultaneous release with NH₄⁺, but NH₄⁺V_f was significantly higher with increased acetate. Thus, labile DOC V_f was not limited by the availability of NH₄⁺, but NH₄⁺V_f was limited by the availability of labile DOC. In contrast, neither glycine nor NH₄⁺V_f changed when released simultaneously indicating either that overall N‐uptake was saturated or that glycine and NH₄⁺ uptake were controlled by different factors. 4. Our results suggest that labile DOC and DON uptake can be equivalent to, or even higher than NH₄⁺ uptake, a solute known to be highly bioreactive, but unlike NH₄⁺ uptake, may not differ among land‐use types and seasons. Moreover, downstream export of nitrogen may be exacerbated by limitation of NH₄⁺ uptake by the availability of labile DOC in headwater streams from the agricultural Midwestern United States. Further research is needed to identify the factors that influence cycling of DOC and DON in streams.     

森林转换对地表径流可溶性有机碳输出浓度和通量的影响

米槠次生林转换成米槠人工幼林和米槠人工促进天然更新幼林(以下简称"人促幼林")后,以这三种森林类型为研究对象,连续监测每次降雨后地表径流量及径流水中可溶性有机碳(DOC)的含量及通量,比较不同森林类型观测结果的差异,并分析降雨对实验结果的影响。结果表明:米槠人工幼林单次产流量是米槠次生林的1.5—19.0倍,观测期间总径流量为5.9倍;米槠人促幼林单次径流量和总径流量均与米槠次生林无显著差异(P0.05)。观测期间米槠次生林、人工幼林、人促幼林径流水DOC浓度值范围为5.9—18.4 mg/L,4.3—13.5 mg/L和3.2—9.9 mg/L,米槠次生林径流水浓度均值(12.6 mg/L)分别是米槠人促幼林(7.6 mg/L)和米槠人工幼林(5.3 mg/L)的1.6和2.4倍。回归分析表明,径流水中DOC浓度与降雨前土壤含水率呈显著相关;降雨前土壤含水率20.8%是一个临界值,含水率低于20.8%时,径流水DOC浓度与降雨前含水率呈显著正相关(P0.05);高于20.8%时,径流水DOC浓度与降雨前土壤含水率呈显著负相关(P0.05)。米槠人工幼林地表径流DOC输出通量是米槠次生林的0.7—5.4倍,观测期间总输出通量为2.1倍;米槠人促林DOC单次通量和观测期间总通量均与米槠次生林差异不显著(P0.05)。三种森林类型DOC输出通量均与降雨量呈显著相关(P0.05)。可见,米槠次生林转变成米槠人工幼林后DOC输出浓度降低,但径流量显著增加,导致DOC输出通量增加;而转变成米槠人促幼林后DOC输出浓度也降低,但径流量并未增加,因而并未增加DOC输出通量。     

Role of dissolved organic carbon in the attenuation of photosynthetically active and ultraviolet radiation in Adirondack lakes

1. We surveyed eighty-five lakes located in the Adirondack Mountain Region of New York State, U.S.A., to characterize the attenuation of photosynthetically active (PAR) and ultraviolet radiation (UVR) in relation to dissolved organic carbon (DOC) concentrations and pH. Attenuation of PAR was quantified in situ . Attenuation was also inferred by measuring the light absorption of filtered lake water samples at wavelengths (300, 340 and 440 nm) representing UV-B, UV-A and PAR.
2. Substantial variation in transparency was observed among lakes in this region. Attenuation depths ( z _1%) for PAR ranged from 0.5 to greater than 20 m, while inferred values for UV-B and UV-A ranged from a few centimetres to > 5 m. Median values of UV-A penetration (0.75 m) and UV-B penetration (0.45 m) corresponded to 11% (UV-A) and 6% (UV-B) of lake maximum depth.
3. Much of the variation in PAR and UVR attenuation was explained by differences in lake DOC. Univariate power models based solely on DOC accounted for 85% (PAR), 90% (UV-A) and 91% (UV-B) of the variation in absorption.
4. Attenuation and absorption coefficients were generally lower for recently acidified lakes compared to acidic and circumneutral lakes which have not undergone recent acidification. However, differences among these three groups of lakes were not statistically significant. Our results suggest that the effects of acidification on the optical properties of a regional population of lakes, even in an area experiencing widespread acidification, are relatively subtle in comparison with other factors contributing to inter-lake variability.
5. The presence of near-shore wetlands is probably a key factor influencing regional variability in DOC and light climate among Adirondack lakes. Temporal variability in climatic factors influencing wetland DOC production and export may mask more subtle influences on lake DOC associated with anthropogenic acidification.     

Growth response of four freshwater algal species to dissolved organic nitrogen of different concentration and complexity

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Productivity of clear and humic lakes: nutrients,phytoplankton, bacteria

The relationships between long-term surface average concentrations of humic acids measured as water colour, dissolved organic carbon (DOC) or Secchi disk transparency and trophic state variables were studied with literature data from more than 600 freshwater lakes. The geometric means of summer surface average nutrient (phosphorus and nitrogen) concentration, phytoplankton biomass (chlorophyll concentration), and hypolimnetic anoxia (anoxic factor) were significantly higher in coloured than in clear lakes. The regressions of colour or DOC on these trophic state variables were positive and significant throughout a range of three orders of magnitude. Phytoplankton or primary productivity was higher in coloured lakes, when expressed per volume of epilimnion. Annual integral primary productivity expressed on an areal basis was smaller in coloured lakes, probably a reflection of shallower phototrophic depths in these lakes. There is evidence that annual integral bacteria productivity is much higher in coloured lakes for two reasons: first, epilimnetic bacteria production was ca. four times higher in coloured lakes, second, other studies have shown that hypolimnetic bacteria production is commonly higher than epilimnetic production, especially in anoxic hypolimnia that are frequent in coloured lakes. All volumetrically expressed variables indicated higher productivity in coloured lakes. In addition, high bacteria productivity reflects a different food chain involving mixotrophs, possibly compensating for low light conditions. Our analyses indicate that primary and secondary productivity is as high as or higher than in clear lakes. This revised version was published online in July 2006 with corrections to the Cover Date.     

Distribution of dissolved organic carbon in lakes of different trophic types

The distributions of dissolved organic carbon (DOC) in the warm season were elucidated in ten lakes of different trophic types in Japan, Russia, and China. DOC showed similar vertical distributions in all the lakes in summer when thermal stratification occurred. DOC in the epilimnion was higher than the value of 0.8mgCl^–1 found in the hypolimnion. In three Japanese lakes, hypolimnion DOC was negatively correlated with apparent oxygen utilization (AOU), reflecting the net oxidation of DOC using the dissolved oxygen in lake water. The DOC:O₂ ratios (0.115–0.179), calculated by the slopes of the regression lines of DOC versus AOU in hypolimnion water, were as low as those of deep-sea water, which indicates low bioavailability of lake water DOC for heterotrophic bacteria. DOC and conductivity did not correlate well except in two Japanese lakes: one showed a positive correlation and the other a negative correlation, indicating DOC loading from the inflowing rivers. Eutrophic lakes tended to have higher DOC values than meso- and oligotrophic lakes, and DOC values in the surface water negatively correlated with Secchi depths.