不同盐度对闽江河口沉积物硝化作用的影响

选择闽江河口鳝鱼滩湿地与道庆洲湿地为对象,采集湿地沉积物在室内进行沉积物硝化培养,分析不同盐度水平对湿地沉积物硝化作用的影响.结果表明:闽江河口湿地沉积物硝化速率普遍较低,鳝鱼滩湿地沉积物最高硝化速率只有0.193 mg·kg^-1·d^-1,而道庆洲湿地沉积物硝化速率最高不超过0.050 mg·kg^-1·d^-1.盐度的升高会抑制闽江河口湿地沉积物的硝化作用.低盐度时(5),硝化速率下降的主要原因是硝化细菌活性受到抑制;随着盐度的升高(10),硝化速率略有上升但仍低于初始值,这是由于随着盐度的升高,盐度对好氧氨化细菌活性的抑制程度有所加强,导致系统产生NH_4^+-N的速率下降,从而造成好氧氨化细菌对表观硝化速率下降的贡献减少.沉积物硝化活性对于盐度的响应存在地域差异.咸水湿地(鳝鱼滩湿地)沉积物中的微生物对盐度的变化适应性较强,从而使该湿地沉积物硝化活性在高盐度条件下仍然较高.淡水湿地(道庆洲湿地)沉积物对盐度变化的适应性较弱,导致其沉积物硝化活性在高盐度条件下低于中等盐度.闽江河口鳝鱼滩与道庆洲的短叶茳芏湿地沉积物硝化作用较低的主要原因是沉积物呈酸性及淹水状态下的缺氧条件弱化了沉积物的硝化作用.两处湿地的硝化速率与硝化活性随时间的变化趋势为先升高后下降,这是由初始NH_4^+-N浓度、氧气含量和反硝化共同作用造成的.     

闽江河口短叶茳芏湿地及围垦养虾塘的沉积物CH_4和N_2O产生潜力

河口湿地围垦转化为水产养殖塘已成沿海地区一个重要的生态环境问题,深刻影响着河口地区的碳、氮生物地球化学循环。为探讨河口湿地围垦成养虾塘后对温室气体产生的影响本研究以福建省闽江河口鳝鱼滩中部的短叶茳芏(Cyperus malaccensis)沼泽湿地及围垦养虾塘为对象,于2016年5月—2017年3月通过室内厌氧培养实验测定了2个生态系统表层沉积物的甲烷(CH_4)与氧化亚氮(N_2O)产生潜力。结果表明:①短叶茳芏湿地和养虾塘的表层沉积物CH_4产生潜力分别为0.33～215.2和0.13～9.41 ng·g~(-1)·d~(-1)均值分别为40.07±23.00和2.16±0.98 ng·g~(-1)·d~(-1),呈现出秋季夏季春季冬季的时间变化特征;②短叶茳芏湿地和养虾塘表层沉积物N_2O产生潜力分别为0.43～0.80和0.45～0.84ng·g~(-1)·d~(-1)均值分别为0.58±0.01和0.61±0.01 ng·g~(-1)·d~(-1);③短叶茳芏湿地沉积物CH_4产生潜力显著高于养虾塘沉积物(P0.01),2个生态系统表层沉积物N_2O产生潜力差异不显著(P0.05)。     

闽江河口区淡水和半咸水潮汐沼泽湿地土壤产甲烷菌多样性

为认识盐度对河口潮汐沼泽湿地土壤产甲烷菌的影响,应用PCR-RFLP技术及测序分析对闽江河口区淡水-半咸水盐度梯度上分布的4个短叶茳芏潮汐沼泽湿地土壤产甲烷菌群落结构进行研究。闽江河口区短叶茳芏潮汐沼泽湿地土壤产甲烷菌群落结构受盐度影响明显,位于下洋洲和塔礁洲的短叶茳芏潮汐淡水沼泽湿地土壤产甲烷菌的香农-威纳多样性指数值分别为2.81和2.65,位于蝙蝠洲和鳝鱼滩的短叶茳芏潮汐半咸水沼泽湿地土壤产甲烷菌香农-威纳多样性指数值分别仅为2.33和2.27。系统发育分析表明:短叶茳芏沼泽湿地土壤产甲烷菌类群主要有甲烷杆菌目(Methanobacteriales),包括Methanobacterium、Methanobrevibacter和Methanobacteriaceae;甲烷微菌目(Methanomicrobiales),主要有Methanoregula,以及甲烷八叠球菌目(Methanosarcinales),主要有Methanosarcina和Methanococcoides。闽江河口区短叶茳芏潮汐淡水沼泽湿地土壤主要的优势产甲烷菌有Methanoregula、Methanosarcina和Methanobacterium,而短叶茳芏潮汐半咸水沼泽湿地土壤主要的优势产甲烷菌则转化为仅以Methanoregula为主。     

珠江河口湿地沉积物硝化作用强度及影响因素研究

采用现场调查和实验室模拟的方法,研究了珠江河口红树林、芦苇和光滩表层湿地沉积物氨氧化细菌(AOB)的分布、硝化强度及其主要影响因素。研究结果表明:三种不同生态环境中AOB数量差别较大。温度、pH、盐度和氨氮的浓度对沉积物硝化作用均有影响。其中盐度和pH对沉积物硝化强度影响最为显著,较高的盐度和较低的pH对硝化过程有明显的抑制作用。偏碱性环境下(pH=8.5)红树林、芦苇和光滩的硝化强度分别是偏酸性环境(pH=6.5)时2.5倍。最适宜的硝化温度在28℃左右,氨氮对氨氧化细菌的生长有促进作用,在含氨氮浓度较高的培养液中硝化强度较强。大型植物根系对硝化作用有明显的促进作用,对河口湿地氮循环具有重要的意义。     

闽江河口湿地植物与土壤灰分及其影响因子分析

以闽江河口鳝鱼滩湿地生长的芦苇(Phragmitesaustralis)和互花米草(Spartina alterniflora)为研究对象,并对相应的36个土壤剖面进行取样分析,揭示它们的(厂火)分含量分布特征,并对其影响因子进行分析。研究结果表明:闽江河口鳝鱼滩湿地芦苇和互花米草各个器官(厂火)分含量的差异性明显,芦苇各部位(厂火)分含量的平均值大于互花米草;芦苇和互花米草湿地土壤(厂火)分含量的垂直分布较为类似,芦苇湿地土壤(厂火)分合量略高于互花米草湿地土壤(厂火)分含量,但两种植物土壤(厂火)分含量的差异并不显著;芦苇和互花米草湿地土壤(厂火)分含量与土壤有机碳有极其显著的负相关关系,与土壤含水量的相关性也较明显;从单一的植物类型来看,互花米草湿地土壤(厂火)分含量与容重、碳含量、含水量和盐度呈显著的相关关系,芦苇湿地土壤(厂火)分含量与土壤pH值有明显的正相关关系。     

闽江河口互花米草不同入侵阶段湿地土壤nirK型反硝化微生物群落结构及多样性

为了明确闽江河口互花米草海向不同入侵阶段湿地土壤nirK型反硝化微生物的群落结构及多样性,在鳝鱼滩东部的互花米草分布区,由陆向海方向选择互花米草海向入侵前的光滩(MF)、入侵1～2年(SAN)和入侵6～7年的互花米草(SA)湿地为研究对象,基于高通量测序技术,测定并分析了不同互花米草入侵阶段湿地土壤nirK型反硝化微生物群落结构及多样性的差异。结果表明:互花米草海向入侵降低了土壤中nirK型反硝化微生物群落的多样性及丰富度。不同入侵阶段湿地土壤的nirK型反硝化微生物主要包括变形菌门和放线菌门,其中变形菌门占绝对优势地位。互花米草海向入侵整体改变了湿地土壤中nirK型反硝化菌属的组成特征,MF、SAN、SA湿地土壤中相对丰度最高的nirK基因菌属分别为慢生根瘤菌属、中慢生根瘤菌属和产碱杆菌属。互花米草海向入侵增加了土壤中nirK型反硝化微生物群落组成的空间异质性,尤其是在SAN样地,这主要与样地本身的环境扰动性较大及互花米草海向入侵加大了样地环境因子的空间异质性有关。互花米草海向入侵主要是通过显著改变土壤理化因子(粒度组成、pH值和含水量)和氮养分条件(全氮、NH₄⁺-N、NO₃^--N)来影响nirK型反硝化微生物的群落结构及多样性。本研究结果有助于揭示互花米草海向入侵对湿地土壤反硝化过程影响的微生物机制。     

翅碱蓬对盐沼沉积物微生物生物量及β-氨氧化细菌群落的影响——以双台河口为例

为了解翅碱蓬植被对盐沼沉积物微生物的影响,于2013年7月、8月、9月和11月对双台河口裸滩和翅碱蓬植被沉积物(10—15 cm)微生物生物量碳(MBC)、微生物生物量氮(MBN)、16S rRNA基因丰度、潜在硝化速率、β-氨氧化细菌(β-AOB)丰度及群落进行了调查。结果表明,不同采样日期裸滩沉积物MBC、翅碱蓬沉积物β-AOB amo A丰度和两种生境潜在硝化速率没有显著差异;而翅碱蓬沉积物MBC、裸滩沉积物β-AOB amo A丰度、两种生境MBN和16S rRNA基因丰度呈现时间波动。当所有采样日期的数据结合分析时,翅碱蓬植被显著影响沉积物MBC、MBN、细菌16S rRNA基因丰度、潜在硝化速率和β-AOB amo A丰度。从裸滩和翅碱蓬沉积物获得的β-AOB序列属于Nitrosospira和Nitrosomonas,翅碱蓬植被对β-AOB群落结构和多样性均有一定的影响。研究结果有助于了解翅碱蓬湿地中微生物的作用,为盐沼生境的生态修复技术提供参考。     

互花米草入侵影响下闽江河口湿地土壤有效硅的时空变化特征

2016年1—12月,选择闽江河口鳝鱼滩的短叶茳芏湿地、互花米草湿地以及二者的交错带湿地为研究对象,采用定位研究方法探讨了互花米草入侵影响下湿地土壤有效硅含量的时空变化特征。结果表明:互花米草入侵影响下3块湿地土壤有效硅含量随时间推移整体呈波动上升趋势;互花米草入侵显著提高了鳝鱼滩湿地30—60 cm土层土壤有效硅含量(P0.01),与短叶茳芏湿地相比,交错带湿地和互花米草湿地30—60 cm土层土壤有效硅含量分别增加了8.56%和19.97%,逐步线性回归分析表明土温和电导是影响其变化的重要因素(P0.01)。研究互花米草入侵影响下湿地土壤有效硅含量的变化特征,对于揭示湿地生态系统生源要素硅生物地球化学循环过程以及互花米草入侵及其扩张机制具有重要意义。     

河口潮间带沉积物重金属累积及生态风险评价

于2015年7月份在闽江口鳝鱼滩湿地布设两条样带(T1和T2),并分别选取5个采样点采集沉积物样品。因重金属Co和V对动物和人类会造成一定危害,且闽江河口潮间带Co和V的研究较少,则对其理化性质、Co和V的空间分布特征以及污染程度等进行了研究,旨在为Co和V的污染研究提供参考。结果显示,该沉积物中Co和V的平均浓度分别为15.19 mg/kg、102.94 mg/kg,均高于福建省背景浓度,与其他河口对比,Co和V浓度处于中等水平。Co和V的含量由陆向海方向略有下降。地累积指数法分析得出沉积物中Co含量呈现轻度污染状态,V呈现无污染状态。此外,Co和V的潜在生态危害系数E_rⁱ分别为7.24—14.85和1.47—4.17,表明潜在生态危害轻微,闽江河口潮间带沉积物中Co和V属于轻度污染。     

亚热带河口陆基养虾塘水体溶解性碳浓度及沉积物-水界面碳通量时空动态特征

以福建闽江和九龙江河口陆基养虾塘为研究对象,通过野外原位观测和室内模拟培养实验,开展了河口陆基养虾塘养殖期间水体溶解性有机碳(DOC)和溶解性无机碳(DIC)及养虾塘沉积物-水界面碳交换通量变化特征的研究。结果表明:时间变化上,养虾塘水体溶解性碳浓度及沉积物-水界面碳通量在闽江河口呈现8月中旬10月中旬6月中旬的特征,在九龙江河口表现为随养殖阶段推移而增加的趋势;空间变化上,闽江河口养虾塘水体溶解性碳浓度及沉积物-水界面碳通量显著高于九龙江河口;沉积物释放溶解性碳速率与水体溶解性碳浓度呈现显著正相关关系,沉积物碳释放过程是引起养虾塘水体溶解性碳浓度时空变化的重要因素。表明河口区水产养虾塘碳循环研究时需考虑不同形态碳生物地球化学循环的时空差异性。     

Denitrification Potential in Lake Sediment Increases Across a Gradient of Catchment Agriculture

Intensification of catchment agriculture has increased nutrient loads and accelerated eutrophication in some lakes, often resulting in episodic harmful algal blooms or prolonged periods of anoxia. The influence of catchment agriculture on lake sediment denitrification capacity as a nitrogen (N) removal mechanism in lakes is largely unknown, particularly in contrast to research on denitrification in agricultural streams and rivers. We measured denitrification enzyme activity (DEA) to assess sediment denitrification potential in seven monomictic and three polymictic lakes that range in the proportion of agriculture in the catchment from 3 to 96% to determine if there is a link between agricultural land use in the lake catchment and sediment denitrification potential. We collected sediment cores for DEA measurements over 3 weeks in austral spring 2008 (October–November). Lake Okaro, with 96% catchment agriculture, had approximately 15 times higher DEA than Lake Tikitapu, with 3% catchment agriculture (232.2 ± 55.9 vs. 15.9 ± 4.5 μg N gAFDM⁻¹ h⁻¹, respectively). Additionally, sediment denitrification potential increased with the proportion of catchment in agriculture (R ² = 0.85, P < 0.001). Our data suggest that lakes retain a high capacity to remove excess N via denitrification under increasing N loads from higher proportions of catchment agriculture. However, evidence from the literature suggests that despite a high capacity for denitrification and longer water residence times, lakes with high N loads will still remove a smaller proportion of their N load. Lakes have a denitrification potential that reflects the condition of the lake catchment, but more measurements of in situ denitrification rates across lake catchments is necessary to determine if this capacity translates to high N removal rates.     

Nitrification and denitrification response to varying periods of desiccation and inundation in a western Kansas stream

Changing environmental conditions and increased water consumption have transformed many historically perennial stream systems into intermittent systems. Multiple drying and wetting events throughout the year might impact many stream processes including nitrification and denitrification, key components of the nitrogen (N) cycle. During summer 2007, an experimental stream was used to dry and then rewet stream sediments to determine the effects of desiccation and rewetting of stream sediment on nitrification and denitrification potentials. Mean (±SE) nitrification and denitrification rates in sediment not dried (controls) were 0.431 ± 0.017 μg NO₃ ⁻–N/cm²/h and 0.016 ± 0.002 μg N₂O–N/cm²/h, respectively. As sediment samples dried, nitrification rates decreased. Rates in sediments dried less than 7 d recovered to levels equal or greater than those in the controls within 1 d of being rewetted. Denitrification rates were not affected by 1 d of drying, but samples dried greater than 1 d experienced reduced rates of denitrification. Denitrification in sediments dried 7 d or less recovered by day seven of being rewetted. Nitrification and denitrification processes failed to fully recover in sediments dried more than 7 d. These results demonstrate that alterations in stream’s hydrology can significantly affect N-cycle processes.     

辽河口滨海湿地CH4排放特征及其影响因素

采用静态箱-气相色谱法,于2016年6—11月连续观测辽河口芦苇湿地、翅碱蓬湿地和裸滩湿地的CH_4排放速率,同时测定温度、氧化还原电位(Eh)、pH值和电导率(EC)等相关环境因子的动态变化。结果表明,3种类型湿地的CH_4排放具有明显的季节变化特征,均呈先上升后下降趋势。芦苇湿地、翅碱蓬湿地(涨潮前)和裸滩湿地(涨潮前)CH_4排放通量变化范围分别为0.447—10.40、0.045—0.509 mg m~(-2) h~(-1)和0.016—0.593 mg m~(-2) h~(-1),观测期内排放通量均值相应为(3.699±3.679)、(0.165±0.156) mg m~(-2) h~(-1)和(0.198±0.191) mg m~(-2) h~(-1),不同类型湿地之间差异显著(P0.01),芦苇湿地裸滩湿地(涨潮前)翅碱蓬湿地(涨潮前)。涨潮过程中,翅碱蓬湿地和裸滩湿地的排放速率分别变化在0.009—0.353 mg m~(-2) h~(-1)和0.018—0.335 mg m~(-2) h~(-1),观测期间其排放速率均值分别为(0.119±0.132) mg m~(-2) h~(-1)和(0.131±0.103) mg m~(-2) h~(-1),明显低于涨潮前(P0.01)。不同湿地类型间CH_4排放通量与电导率(EC)呈显著负相关(P0.01)。研究结果表明,潮汐和电导率均为影响辽河口不同类型湿地中CH_4排放的关键因子。     

Modeling nitrogen cycling in a coastal fresh water sediment

Increased nitrogen (N) loading to coastal marine and freshwater systems is occurring worldwide as a result of human activities. Diagenetic processes in sediments can change the N availability in these systems, by supporting removal through denitrification and burial of organic N (N_org) or by enhancing N recycling. In this study, we use a reactive transport model (RTM) to examine N transformations in a coastal fresh water sediment and quantify N removal rates. We also assess the response of the sediment N cycle to environmental changes that may result from increased salinity which is planned to occur at the site as a result of an estuarine restoration project. Field results show that much of the N_org deposited on the sediment is currently remineralized to ammonium. A rapid removal of nitrate is observed in the sediment pore water, with the resulting nitrate reduction rate estimated to be 130 μmol N cm⁻² yr⁻¹. A model sensitivity study was conducted altering the distribution of nitrate reduction between dissimilatory nitrate reduction to ammonium (DNRA) and denitrification. These results show a 40% decline in sediment N removal as NO ₃ ⁻ reduction shifts from denitrification to DNRA. This decreased N removal leads to a shift in sediment-water exchange flux of dissolved inorganic nitrogen (DIN) from near zero with denitrification to 133 μmol N cm⁻² yr⁻¹ if DNRA is the dominant pathway. The response to salinization includes a short-term release of adsorbed ammonium. Additional changes expected to result from the estuarine restoration include: lower NO ₃ ⁻ concentrations and greater SO ₄ ²⁻ concentrations in the bottom water, decreased nitrification rates, and increased sediment mixing. The effect of these changes on net DIN flux and N removal vary based on the distribution of DNRA versus denitrification, illustrating the need for a better understanding of factors controlling this competition.     

Denitrification in the periphyton associated with plant shoots and in the sediment of a wetland system supplied with sewage treatment plant effluent

Seasonal variation in denitrification and major factors controlling this process were determined in sediment, microbial communities attached to plant shoots (periphyton) and in the water of a Phragmites and an Elodea-dominated stand of a constructed wetland system between May 1997 and February 1998. The wetland was supplied with effluent from a sewage treatment plant. The denitrification rate in periphyton on plants shoots (expressed per shoot area) was always considerably higher than in the sediment and varied with the chlorophyll-a content of the periphyton in the course of the year. The algae in the periphyton provided attachment surfaces and probably also organic compounds to the denitrifying bacteria. Decreases in periphyton biomass and denitrification rate in the Phragmites and Elodea-dominated stands during the growing season were associated with enhanced shading by Phragmites shoots or a floating layer of macro-algae and Lemna spp., respectively. Light availability and the denitrification rate of periphyton increased again after the Phragmites shoots were cut in October. Nitrate appeared to limit the denitrification rate in the sediment. Periphyton denitrification rates were mostly lower on Elodea shoots than on Phragmites shoots, in spite of the higher living algal biomass on Elodea shoots. This difference was associated with lower nitrate concentrations in the Elodea-dominated stand. In the two stands, the daily denitrification rates in periphyton on shoots of Phragmites australis (44.4–121 mg N m^–2 stand area d^–1) and Elodea nuttallii (14.8–33.1 mg N m^–2 d^–1) were clearly more important than rates in the sediment (0.5–25.5 mg N m^–2 d^–1) or the water (0.4–3.9 mg N m^–2 d^–1). The presence of few bacteria attachment sites or low organic carbon availability possibly resulted in low denitrification rates in the water. Denitrification appeared to be a major process in nitrate removal from the through-flowing water in this wetland system.     

Benithic-pelagic coupling of nutrient metabolism along an estuarine eutrophication gradient

The shallow, brackish (11–18% salinity) Roskilde Fjord represents a eutrophication gradient with annual averages of chlorophyll, ranging from 3 to 25 mg chl a m^–3. Nutrient loadings in 1985 were 11.3–62.4 g N m^–2 yr^–1 and 0.4–7.3 g P m^–2 yr^–1. A simple one-layer advection-diffusion model was used to calculate mass balances for 7 boxes in the fjord. Net loss rates varied from –32.2 to 17.9 g P m^–2 yr^–1 and from –3.3 to 66.8 g N m^–2, corresponding to 74% of the external P-loading and 88% of the external N-loading to the entire estuary.Gross sedimentation rates measured by sediment traps were between 7 and 52 g p m^–2 yr^–1 and 50 and 426 g N M^–2 yr^–1, respectively. Exchangeable sediment phosphorus varied in annual average between 2.0 and 4.8 g P m^–2 and exchangeable sediment nitrogen varied from 1.9 to 33.1 g N m–1. Amplitudes in the exchangeable pools followed sedimentation peaks with delays corresponding to settling rates of 0.3 m d^–1. Short term nutrient exchange experiments performed in the laboratory with simultaneous measurements of sediment oxygen uptake showed a release pattern following the oxygen uptake, the changes in the exchangeable pools and the sedimentation peaks.The close benthic-pelagic coupling also exists for the denitrification with maxima during spring of 5 to 20 mmol N m^–2 d–1. Denitrification during the nitrogen-limited summer period suggests dependence on nitrification. Comparisons with denitrification from other shallow estuaries indicate a maximum for denitrification in estuaries of about 250 µmol N m^–2 h^–2 achieved at loading rates of about 25–125 g N m^–2 yr^–1.     

Differential metabolism of dimethylsulfoniopropionate and acrylate in saline and brackish intertidal sediments

In anoxic Spartina altemiflora—dominated sediments along a naturally occuring salinity gradient (the Cooper River estuary, South Carolina, U.S.A.), dimethylsulfoniopropionate (DMSP) was metabolized to dimethyl sulfide (DMS) and acrylate by sediment microbes. The rate of DMSP degradation and acrylate mineralization by sediment microbes was similar at all sites along this 25-km transect. However, sediments amended with acrylate (or DMSP) showed significantly higher rates of N₂ fixation (measured as acetylene reduction activity) (ARA) in the saline sediments downstream than brackish sediments. These results are consistent with the fact that acrylate stimulated the rates of both denitrification and CO₂ production in the saline sediments at the mouth of the river more than tenfold over rates in brackish sediments. Enrichment experiments indicate that microbes capable of using DMSP or acrylate were not present in upstream sediments despite the fact that microbial biomass, percent organic matter, and both glucose-stimulated ARA and denitrification were highest upstream. It appears that acrylate utilizing, N₂ fixing, and denitrifying populations are insignificant in the lower salinity sediments of the estuary. These results may reflect the availability of DMSP, which averaged 10.3 nmol g wet wt^–1 of saline sediments and levels less than our detection limit (1 m) in brackish sediments. Correspondence to: D.C. Yoch.     

Factors Controlling Sediment Denitrification in Midwestern Streams of Varying Land Use

We investigated controls on stream sediment denitrification in nine headwater streams in the Kalamazoo River Watershed, Michigan, USA. Factors influencing denitrification were determined by using experimental assays based on the chloramphenicol-amended acetylene inhibition technique. Using a coring technique, we found that sediment denitrification was highest in the top 5 cm of the benthos and was positively related to sediment organic content. To determine the effect of overlying water quality on sediment denitrification, first-order stream sediments were assayed with water from second- and third-order downstream reaches, and often showed higher denitrification rates relative to assays using site-specific water from the first-order stream reach. Denitrification was positively related to nitrate (NO₃ ⁻) concentration, suggesting that sediments may have been nutrient-limited. Using stream-incubated inorganic substrata of varying size classes, we found that finer-grained sand showed higher rates of denitrification compared to large pebbles, likely due to increased surface area per volume of substratum. Denitrification was measurable on both inorganic substrata and fine particulate organic matter loosely associated with inorganic particles, and denitrification rates were related to organic content. Using nutrient-amended denitrification assays, we found that sediment denitrification was limited by NO₃ ⁻ or dissolved organic carbon (DOC, as dextrose) variably throughout the year. The frequency and type of limitation differed with land use in the watershed: forested streams were NO₃ ⁻-limited or co-limited by both NO₃ ⁻ and DOC 92% of the time, urban streams were more often NO₃ ⁻-limited than DOC-limited, whereas agricultural stream sediments were DOC-limited or co-limited but not frequently limited by NO₃ ⁻ alone.