Dynamics of Microbial Processes in the Stratal Waters of the Romashkinskoe Oil Field

The dynamics of the microbial processes developing in parallel with the exploitation of the Romashkinskoe oil field (Tatarstan) were studied in two areas differing in the degree of stratal water freshening. Flooding of the strata, in conjunction with purposeful measures on stratal microflora activation, was shown to increase the microbial population density and activate both methanogenesis and sulfate reduction; the latter process was limited by the low sulfate concentration. Development of anaerobic processes correlated with changes in acetate concentration in the stratal water. High mineralization (over 200 g/l) inhibited the stratal water microflora even if other conditions were favorable. Isotopic analysis of the carbonate carbon showed that the bicarbonate concentration increased in the stratal water due to microbial degradation of oil hydrocarbons and further participation of the biogenic carbon dioxide in dissolution of the carbonate cement of the oil-bearing strata. In strongly desalinated stratal water, the proportion of the newly formed bicarbonate was as high as 80%.     

Microbiological and production characteristics of the high-temperature Kongdian bed revealed during field trial of biotechnology for the enhancement of oil recovery

Microbiological technology for the enhancement of oil recovery based on the activation of the stratal microflora was tested in the high-temperature horizons of the Kongdian bed (60 degrees C) of the Dagang oil field (China). This biotechnology consists in the pumping of a water-air mixture and nitrogen and phosphorus mineral salts into the oil stratum through injection wells in order to stimulate the activity of the stratal microflora which produce oil-releasing metabolites. Monitoring of the physicochemical, microbiological, and production characteristics of the test site has revealed large changes in the ecosystem as a result of the application of biotechnology. The cell numbers of thermophilic hydrocarbon-oxidizing, fermentative, sulfate-reducing, and methanogenic microorganisms increased 10-10 000-fold. The rates of methanogenesis and sulfate reduction increased in the near-bottom zone of the injection wells and of some production wells. The microbial oil transformation was accompanied by the accumulation of bicarbonate ions, volatile fatty acids, and biosurfactants in the formation waters, as well as of CH4 and CO2 both in the gas phase and in the oil. Microbial metabolites promoted the additional recovery of oil. As a result of the application of biotechnology, the water content in the production liquid from the test site decreased, and the oil content increased. This allowed the recovery of more than 14000 tons of additional oil over 3.5 years.     

Microbiological and production characteristics of the high-temperature Kongdian petroleum reservoir revealed during field trial of biotechnology for the enhancement of oil recovery

Microbiological technology for the enhancement of oil recovery based on the activation of the stratal microflora was tested in the high-temperature horizons of the Kongdian bed (60°C) of the Dagang oil-field (China). This biotechnology consists in the pumping of a water-air mixture and nitrogen and phosphorus mineral salts into the oil stratum through injection wells in order to stimulate the activity of the stratal microflora which produce oil-releasing metabolites. Monitoring of the physicochemical, microbiological, and production characteristics of the trial site has revealed large changes in the ecosystem as a result of the application of biotechnology. The cell numbers of thermophilic hydrocarbon-oxidizing, fermentative, sulfate-reducing, and methanogenic microorganisms increased 10–10000-fold. The rates of methanogenesis and sulfate reduction increased in the near-bottom zone of the injection wells and of some production wells. The microbial oil transformation was accompanied by the accumulation of bicarbonate ions, volatile fatty acids, and biosurfactants in the formation waters, as well as of CH₄ and CO₂ both in the gas phase and in the oil. Microbial metabolites promoted the additional recovery of oil. As a result of the application of biotechnology, the water content in the production liquid from the trial site decreased, and the oil content increased. This allowed the recovery of more than 14000 tons of additional oil over 3.5 years.     

不同压力条件下油藏内源细菌群落激活过程中变性梯度凝胶电泳分析?

摘要：【目的】了解常压（1 MPa）和高压（10 MPa）条件下内源细菌激活中的细菌群落和结构的变化。【方法】利用胜利油田沾3×24井产出液水样,在常压和高压下进行富集培养后,定时取样,样品用变性梯度凝胶电泳（denature gradient gel electrophoresis,DGGE）技术分析。【结果】通过对内源微生物激活前后DGGE条带的数量和亮度变化分析表明,油藏环境中的细菌在种类和数量上并不丰富,激活剂的加入改变了菌群原有的贫营养环境,从而使一些因营养缺乏生长受抑制细菌得以大量繁殖,菌群结     

Influences of Electron Donor,Bicarbonate, and Sulfate on Bioreduction Processes and Manganese/Copper Redistributions among Minerals in a Water-Saturated Sediment

Bioreduction processes have profound influences on mobility and bioavailability of metals in soils and sediments. In this study, a series of microcosm studies were conducted to investigate bioreduction progresses (ferric iron and sulfate reduction) and their influences on manganese and copper element redistributions with the change of microbial community under various geochemical conditions. Results indicated that ethanol stimulated higher rates of bioreduction processes than acetate did. High-concentration bicarbonate and sulfate addition inhibited iron reduction but not sulfate reduction. Sequential extraction revealed that the exchangeable and carbonate bonding-iron were increased in ethanol amendment, whereas those of copper were decreased. The elevated bicarbonate concentration and sulfate addition both influenced the mobility and redistribution of metals. 16S rRNA analysis indicated that ethanol amendment stimulated the growths of microbial iron and sulfate reducer. A high concentration of bicarbonate suppressed the growth of iron reducer Geobacteraceae, but showed limited effect on sulfate reducers. This study concluded that geochemical conditions such as electron donor, bicarbonate and sulfate concentration influenced the microbial community and led to changes in bioreduction processes and metal distributions in the anerobic sediments.     

Microbiological investigations of high-temperature horizons of the Kongdian petroleum reservoir in connection with field trial of a biotechnology for enhancement of oil recovery

The physicochemical conditions and microbiological characteristics of the formation waters of the Kongdian bed of the Dagang oil field (China) were studied. It was demonstrated that this bed is a high-temperature ecosystem with formation waters characterized by low mineralization. The concentrations of nitrogen and phosphorus compounds, as well as of electron acceptors, are low. Oil and oil gas are the main organic matter sources. The bed is exploited with water-flooding. The oil stratum was inhabited mostly by anaerobic thermophilic microorganisms, including fermentative (10(2)-10(5) cells/ml), sulfate-reducing (0-10(2) cells/ml), and methanogenic (0-10(3) cells/ml) microorganisms. Aerobic bacteria were detected mainly in the near-bottom zone of injection wells. The rate of sulfate reduction varied from 0.002 to 18.940 microg S(2-) l(-1) day(-1) and the rate of methanogenesis from 0.012 to 16.235 microg CH4 l(-1) day(-1). Microorganisms with great biotechnological potential inhabited the bed. Aerobic thermophilic bacteria were capable of oxidizing oil with the formation of biomass, the products of partial oxidation of oil (volatile acids), and surfactants. During growth on the culture liquid of oiloxidizing bacteria, methanogenic communities produced methane and carbon dioxide, which also had oil-releasing capabilities. Using various labeled tracers, the primary filtration flows of injected solutions at the testing site were studied. Our comprehensive investigations allowed us to conclude that the tested method for microbial enhancement of oil recovery based on the activation of the stratal microflora can be applied in the Kongdian bed horizons.     

Potential role for microbial ureolysis in the rapid formation of carbonate tufa mounds

Modern carbonate tufa towers in the alkaline (~pH 9.5) Big Soda Lake (BSL), Nevada, exhibit rapid precipitation rates (exceeding 3 cm/year) and host diverse microbial communities. Geochemical indicators reveal that carbonate precipitation is, in part, promoted by the mixing of calcium-rich groundwater and carbonate-rich lake water, such that a microbial role for carbonate precipitation is unknown. Here, we characterize the BSL microbial communities and evaluate their potential effects on carbonate precipitation that may influence fast carbonate precipitation rates of the active tufa mounds of BSL. Small subunit rRNA gene surveys indicate a diverse microbial community living endolithically, in interior voids, and on tufa surfaces. Metagenomic DNA sequencing shows that genes associated with metabolisms that are capable of increasing carbonate saturation (e.g., photosynthesis, ureolysis, and bicarbonate transport) are abundant. Enzyme activity assays revealed that urease and carbonic anhydrase, two microbial enzymes that promote carbonate precipitation, are active in situ in BSL tufa biofilms, and urease also increased calcium carbonate precipitation rates in laboratory incubation analyses. We propose that, although BSL tufas form partially as a result of water mixing, tufa-inhabiting microbiota promote rapid carbonate authigenesis via ureolysis, and potentially via bicarbonate dehydration and CO₂ outgassing by carbonic anhydrase. Microbially induced calcium carbonate precipitation in BSL tufas may generate signatures preserved in the carbonate microfabric, such as stromatolitic layers, which could serve as models for developing potential biosignatures on Earth and elsewhere.     

Phylogenetic diversity of aerobic saprotrophic bacteria isolated from the Daqing oil field

A diverse and active microbial community in the stratal waters of the Daqing oil field (China), which is exploited with the use of water-flooding, was found to contain aerobic chemoheterotrophic bacteria (including hydrocarbon-oxidizing ones) and anaerobic fermentative, sulfate-reducing, and methanogenic bacteria. The aerobic bacteria were most abundant in the near-bottom zones of injection wells. Twenty pure cultures of aerobic saprotrophic bacteria were isolated from the stratal waters. Under laboratory conditions, they grew at temperatures, pH, and salinity values typical of the stratal water from which they were isolated. These isolates were found to be able to utilize crude oil and a wide range of hydrocarbons, fatty acids, and alcohols. Phylogenetic analysis carried out with the use of complete 16S rRNA sequences showed that the isolates could be divided into three major groups: gram-positive bacteria with a high and a low G + C content of DNA and gram-negative bacteria of the gamma-subclass of the Proteobacteria. Gram-positive isolates belonged to the genera Bacillus, Brevibacillus, Rhodococcus, Dietzia, Kocuria, Gordonia, Cellulomonas, and Clavibacter. Gram-negative isolates belonged to the genera Pseudomonas and Acinetobacter. In their 16S rRNA sequences, many isolates were similar to the known microbial species and some probably represented new species.     

油藏本源反硝化功能菌的产气作用(N2O)及对原油物性的影响

利用叠皿夹层培养法从新疆油田采出水中筛选到8株反硝化菌株T1、D1、D44、D46、D15、S1、S2、S6,经16S rDNA序列测定鉴定分析,这8株菌分别与施氏假单胞菌(T1、D1、D44)、恶臭假单胞菌(D46、D15)和铜绿假单胞菌(S1、S2、S6)相似,相似度均达到100%。通过室内批次培养试验,评价了这8株菌利用不同碳源的反硝化产气作用(N₂O)及对原油物性的影响。结果表明: 在以蔗糖为碳源时,产气量最大,以甘油为碳源时,产N₂O气浓度最高;菌株反硝化代谢过程导致原油体积膨胀和粘度降低,膨胀率与N₂O气体浓度呈显著正相关,相关系数为0.983,但与产气体积无相关性;铜绿假单胞菌株S1、S2、S6在以甘油为唯一碳源时产生少量表面活性剂(530～730 mg·L^-1),可降低表面张力并具有乳化原油的作用,但其产气量较少,对原油的膨胀与降粘作用低于其他反硝化菌株。研究提示,在筛选采油功能菌时,菌株反硝化产N₂O气体的能力应给予足够重视。     

Phylogenetic Diversity of Aerobic Saprotrophic Bacteria Isolated from the Daqing Oil Field

A diverse and active microbial community in the stratal waters of the Daqing oil field (China), which is exploited with the use of water-flooding, was found to contain aerobic chemoheterotrophic bacteria (including hydrocarbon-oxidizing ones) and anaerobic fermentative, sulfate-reducing, and methanogenic bacteria. The aerobic bacteria were most abundant in the near-bottom zones of injection wells. Twenty pure cultures of aerobic saprotrophic bacteria were isolated from the stratal waters. Under laboratory conditions, they grew at temperatures, pH, and salinity values typical of the stratal water from which they were isolated. These isolates were found to be able to utilize crude oil and a wide range of hydrocarbons, fatty acids, and alcohols. Phylogenetic analysis carried out with the use of complete 16S rRNA sequences showed that the isolates could be divided into three major groups: gram-positive bacteria with a high and a low G+C content of DNA and gram-negative bacteria of the -subclass of the Proteobacteria. Gram-positive isolates belonged to the genera Bacillus, Brevibacillus, Rhodococcus, Dietzia, Kocuria, Gordonia, Cellulomonas, and Clavibacter. Gram-negative isolates belonged to the genera Pseudomonas and Acinetobacter. In their 16S rRNA sequences, many isolates were similar to the known microbial species and some probably represented new species.     

Microbiological investigations of high-temperature horizons of the Kongdian petroleum reservoir in connection with field trial of a biotechnology for enhancement of oil recovery

The physicochemical conditions and microbiological characteristics of the formation waters of the Kongdian oilfield of the Dagang oilfield (China) were studied. It was demonstrated that this oilfield is a high-temperature ecosystem with formation waters characterized by low mineralization. The concentrations of nitrogen and phosphorus compounds, as well as of electron acceptors, are low. Oil and oil gas are the main organic matter sources. The oilfield is exploited with water-flooding. The oil stratum was inhabited mostly by anaerobic thermophilic microorganisms, including fermentative (10²–10⁵ cells/ml), sulfate-reducing (0–10² cells/ml), and methanogenic (0–10³ cells/ml) microorganisms. Aerobic bacteria were detected mainly in the near-bottom zone of injection wells. The rate of sulfate reduction varied from 0.002 to 18.940 μg S^2? l^?1 day^?1 and the rate of methanogenesis from 0.012 to 16.235 μg CH₄ l^?1 day^?1. Microorganisms with great biotechnological potential inhabited the oilfield. Aerobic thermophilic bacteria were capable of oxidizing oil with formation of biomass, the products of partial oxidation of oil (volatile acids), and surfactants. During growth on the culture liquid of oil-oxidizing bacteria, methanogenic communities produced methane and carbon dioxide, which also had oil-releasing capabilities. Using various labeled tracers, the primary filtration flows of injected solutions at the test site were studied. Our comprehensive investigations allowed us to conclude that the method for microbial enhancement of oil recovery based on the activation of the stratal microflora can be applied in the Kongdian oilfield.     

Carbon load in aquatic ecosystems affects the diversity and biomass of water biofilm consortia and the persistence of the pathogen Campylobacter jejuni within them

The influence of carbon load on autochthonous water microflora population distribution and diversity, and on the persistence of Campylobacter jejuni, was examined with a two-stage aquatic biofilm model. Serine was chosen since it is a carbon source utilised by C. jejuni and concentrations were chosen to reflect upper limits of amino acid load reported in surface water. The total viable count of the autochthonous biofilm microflora increased with increasing serine concentration (10-fold and 20-fold with 5 nM and 5 μM serine, respectively), as did the counts of the microflora in the planktonic phase. Differences in biofilm species distribution as determined by culture were small with changes in temperature or the addition of serine; but was markedly affected by serine as determined by light microscopy, becoming more luxuriant and dominated by long filamentous cells. The addition of serine to the water significantly and progressively reduced the persistence of C. jejuni, which decreased by 25% and 50% with serine concentrations of 5 nM and 5 μM respectively. We have demonstrated that carbon load affects the species diversity and density of both the planktonic and biofilm phase of aquatic autochthonous microflora. Although the survival of C. jejuni in water in a culturable form was sufficient for this to be an important vehicle for its transmission, carbon load significantly influenced survival; an increase in serine concentration significantly reduced survival.     

Microbial potential for cleaning the oiled iron scale

The possibility of using microorganisms to clean oiled iron scale of metallurgical production was investigated with the goal of recuperation. A stable microbial association growing on mineral oil as the sole carbon source was isolated from a sample from oiled iron scale taken directly from a metallurgical plant. For microbial cultures isolated from this association, the taxonomic position, as well as their morphological and cultural characteristics, were determined. The microorganisms belonged to the genera Luteimonas, Alcanivorax, Flavobacterium, and Pseudomonas. Microbial associations oxidizing mineral oil were found to contain some microorganisms incapable of its utilization, which stimulated the hydrocarbon-oxidizing microflora. Application of the isolates, as well as of the strains from microbial collections, resulted in a 58% decrease in residual oil content in treated samples of the oiled iron scale.     

Characterization of a prototype industrial on-line analyzer for bicarbonate/carbonate monitoring

In many biological reactors bicarbonate is the major species determining pH buffering capacity, or alkalinity. In anaerobic digesters bicarbonate levels should be within 10 to 50 mM for stable operation. Bicarbonate alkalinity in wastewater treatment processes in routinely measured off-line titrimetrically. Recently we have described the principle of a novel on-line method of measuring bicarbonate alkalinity. In the prototype device described here, a continuous stream (15 cm(3) min(-1)) of the substrate to be monitored was saturated with gaseous CO2, acidified by the addition of excess acid, and the rate of carbon dioxide evolution, proportional to the concentration of bicarbonate/carbonate in the liquid flow, continuously measured by a sensitive gas meter. The instrument was robust and its response was satisfactory for wastewater treatment process control applications, with linearity in the range 5 to 50 mM HCO3(-), a response time in the order of 30 min, and accuracy of the order of 7% in the concentration range 5 to 50 mM sodium bicarbonate. The device was not affected by interference from volatile fatty acids, does not make use of pH probes which in many wastes are subject to fouling, and may form the basis of a digester control strategy. (c) 1994 John Wiley & Sons, Inc.     

The apparent permeability coefficient for proton flux through phosphatidylcholine vesicles is dependent on the direction of flux

A dioleoylphosphatidylcholine unilamellar vesicle model system was used to determine proton permeability. The fluorescence of the pH reporter group, pyranine, trapped within vesicles with a difference in pH across the bilayer, was digitized and analyzed with numerical integration. When H+ flux was initiated by the acidification of the external buffer (acid jump), the apparent H+ permeability was found to be a linear function of the reciprocal of the internal H+ concentration with the slope inversely proportional to the initial size of the H+ gradient. When flux was initiated by the alkalinization of the external buffer (base jump), the apparent permeability coefficient was constant for each external H+ concentration. However, the value of the apparent permeability was linearly dependent on the reciprocal of the external H+. The possibility that carbonates (carbon dioxide, carbonic acid, bicarbonate and carbonate) could be acting as proton carriers was tested by adding millimolar concentrations of bicarbonate to solutions greatly reduced in carbonates. The slopes of the graphs of apparent permeability coefficient vs. reciprocal H+ were linear functions of added bicarbonate concentration for both acid and base jump conditions. These observations were interpreted in terms of a model suggesting that carbonic acid or carbon dioxide together with bicarbonate was an efficient proton carrier across phospholipid bilayers.     

高温油藏内源微生物及其提高采收率潜力研究

大港孔店油田油藏特征、流体和微生物性质分析结果表明, 属于高温生态环境, 地层水矿化度较低, 氮、磷浓度低, 而且缺乏电子受体, 主要的有机物来源是油气。油田采用经过除油处理的油藏产出水回注方式开发, 油层中存在的微生物类型主要是厌氧嗜热菌, 包括发酵菌(102个/mL~105个/mL), 产甲烷菌(103个/mL); 好氧菌主要存在于注水井周围。硫酸盐还原菌(SRB)还原速率0.002 mg S2-/(L·d) ~18.9 mg S2-/(L·d), 产甲烷菌产甲烷速率0.012 mgCH4/(L·d)~16.2 mgCH4/(L·d)。好氧菌能够氧化油形成生物质, 部分氧化产物为挥发性脂肪酸和表面活性剂。产甲烷菌在油氧化菌液体培养基中产生CH4, CO2为好氧微生物和厌氧微生物的共同代谢产物。这些产物具有提高原油流动性的作用。用示踪剂研究了注入水渗流方向。通过综合分析, 油藏微生物具有较大的潜力, 基于激活油层菌的提高采收率方法在该油田是可行的。     

高温油藏内源微生物及其提高采收率潜力研究

大港孔店油田油藏特征、流体和微生物性质分析结果表明,属于高温生态环境,地层水矿化度较低,氮、磷浓度低,而且缺乏电子受体,主要的有机物来源是油气.油田采用经过除油处理的油藏产出水回注方式开发,油层中存在的微生物类型主要是厌氧嗜热菌,包括发酵菌(102个/mL～105个/mL),产甲烷菌(103个/mL);好氧菌主要存在于注水井周围.硫酸盐还原菌(SRB)还原速率0.002 μg S2-/(L·d)～18.9 μg S2-/(L·d),产甲烷菌产甲烷速率0.012 μgCH4/(L·d)～16.2 μgCH4/(L·d).好氧菌能够氧化油形成生物质,部分氧化产物为挥发性脂肪酸和表面活性荆.产甲烷菌在油氧化菌液体培养基中产生CH4,CO2为好氧微生物和厌氧微生物的共同代谢产物.这些产物具有提高原油流动性的作用.用示踪剂研究了注入水渗流方向.通过综合分析,油藏微生物具有较大的潜力,基于激活油层茵的提高采收率方法在该油田是可行的.     

Microbiological and production characteristics of the Dagang high-temperature heavy oil reservoir (block no. 1) during trials of the biotechnology for enhanced oil recovery

Microbiological and biogeochemical data on the Kongdian bed (block no. 1) of the Dagang high-temperature oilfield during trials of the biotechnology for enhanced oil recovery are reported. Oil-bearing horizons of block no. 1 are characterized by high temperature (56.9–58.4°C), complex geological conditions, and heavy oil (density 0.966–0.969 g/cm³). The biotechnology implied injecting oxygen as an air-water mixture or H₂O₂ together with aqueous solution of nitrogen and phosphorus mineral salts through injection wells in order to activate the oilfield microbial community. In the course of trials, an increase in abundance of aerobic and anaerobic microorganisms was revealed, as well as increased methanogenesis rate in formation water. Microbial oxidation of heavy oil resulted in increased concentration of mineral carbonates dissolved in formation water, changes in the stable carbon isotopic composition δ¹³C/Σ(CO₂ + HCO₃ ^- + CO₃ ^2-), formation of biosurfactants, and decreased interfacial tension of formation water. Application of the biotechnology at the Kongdian bed (block no. 1) resulted in additional recovery of 6331 t oil. Oil viscosity in the zone of production wells located at the North block of the Kongdian bed decreased by 11%. A total of 46152 t additional oil was recovered at three experimental sites of the Dagang oilfield (North block and block no. 1 of the Kongdian bed and the Gangxi bed), which is an indication of high efficiency of the technology for activation of the oilfield microflora for heavy oil replacement from high-temperature oilfields.     

Ratio [13C]/[12C] as an index for express estimation of hydrocarbon-oxidizing potential of microbiota in soil polluted with crude oil

The hydrocarbon-oxidizing potential of soil microbiota and hydrocarbon-oxidizing microorganisms introduced into soil was studied based on the quantitative and isotopic characteristics of carbon in products formed in microbial degradation of oil hydrocarbons. Comparison of CO2 production rates in native soil and that polluted with crude oil showed the intensity of microbial mineralization of soil organic matter (SOM) in the presence of oil hydrocarbons to be higher as compared with non-polluted soil, that is, revealed a priming effect ofoil. The amount of carbon of newly synthesized organic products (cell biomass and exometabolites) due to consumed petroleum was shown to significantly exceed that of SOM consumed for production of CO2. The result of microbial processes in oil-polluted soil was found to be a potent release of carbon dioxide to the atmosphere.     

Microbial activities and dissolved organic matter dynamics in oil-contaminated surface seawater from the Deepwater Horizon oil spill site

The Deepwater Horizon oil spill triggered a complex cascade of microbial responses that reshaped the dynamics of heterotrophic carbon degradation and the turnover of dissolved organic carbon (DOC) in oil contaminated waters. Our results from 21-day laboratory incubations in rotating glass bottles (roller bottles) demonstrate that microbial dynamics and carbon flux in oil-contaminated surface water sampled near the spill site two weeks after the onset of the blowout were greatly affected by activities of microbes associated with macroscopic oil aggregates. Roller bottles with oil-amended water showed rapid formation of oil aggregates that were similar in size and appearance compared to oil aggregates observed in surface waters near the spill site. Oil aggregates that formed in roller bottles were densely colonized by heterotrophic bacteria, exhibiting high rates of enzymatic activity (lipase hydrolysis) indicative of oil degradation. Ambient waters surrounding aggregates also showed enhanced microbial activities not directly associated with primary oil-degradation (β-glucosidase; peptidase), as well as a twofold increase in DOC. Concurrent changes in fluorescence properties of colored dissolved organic matter (CDOM) suggest an increase in oil-derived, aromatic hydrocarbons in the DOC pool. Thus our data indicate that oil aggregates mediate, by two distinct mechanisms, the transfer of hydrocarbons to the deep sea: a microbially-derived flux of oil-derived DOC from sinking oil aggregates into the ambient water column, and rapid sedimentation of the oil aggregates themselves, serving as vehicles for oily particulate matter as well as oil aggregate-associated microbial communities.