984 resultados para Diatoms Spatial gradients


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The spatial and temporal distribution of planktonic, sediment-associated and epiphytic diatoms among 58 sites in Biscayne Bay, Florida was examined in order to identify diatom taxa indicative of different salinity and water quality conditions, geographic locations and habitat types. Assessments were made in contrasting wet and dry seasons in order to develop robust assessment models for salinity and water quality for this region. We found that diatom assemblages differed between nearshore and offshore locations, especially during the wet season when salinity and nutrient gradients were steepest. In the dry season, habitat structure was primary determinant of diatom assemblage composition. Among a suite of physicochemical variables, water depth and sediment total phosphorus (STP) were most strongly associated with diatom assemblage composition in the dry season, while salinity and water total phosphorus (TP) were more important in the wet season. We used indicator species analysis (ISA) to identify taxa that were most abundant and frequent at nearshore and offshore locations, in planktonic, epiphytic and benthic habitats and in contrasting salinity and water quality regimes. Because surface water concentrations of salts, total phosphorus, nitrogen (TN) and organic carbon (TOC) are partly controlled by water management in this region, diatom-based models were produced to infer these variables in modern and retrospective assessments of management-driven changes. Weighted averaging (WA) and weighted averaging partial least squares (WA-PLS) regressions produced reliable estimates of salinity, TP, TN and TOC from diatoms (r2 = 0.92, 0.77, 0.77 and 0.71, respectively). Because of their sensitivity to salinity, nutrient and TOC concentrations diatom assemblages should be useful in developing protective nutrient criteria for estuaries and coastal waters of Florida.

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Intertidal flats of the estuarine macro-intertidal Baie des Veys (France) were investigated to identify spatial features of sediment and microphytobenthos (MPB) in April 2003. Gradients occurred within the domain, and patches were identified close to vegetated areas or within the oyster-farming areas where calm physical conditions and biodeposition altered the sediment and MPB landscapes. Spatial patterns of chl a content were explained primarily by the influence of sediment features, while bed elevation and compaction brought only minor insights into MPB distribution regulation. The smaller size of MPB patches compared to silt patches revealed the interplay between physical structure defining the sediment landscape, the biotic patches that they contain, and that median grain-size is the most important parameter in explaining the spatial pattern of MPB. Small-scale temporal dynamics of sediment chl a content and grain-size distribution were surveyed in parallel during 2 periods of 14 d to detect tidal and seasonal variations. Our results showed a weak relationship between mud fraction and MPB biomass in March, and this relationship fully disappeared in July. Tidal exposure was the most important parameter in explaining the summer temporal dynamics of MPB. This study reveals the general importance of bed elevation and tidal exposure in muddy habitats and that silt content was a prime governing physical factor in winter. Biostabilisation processes seemed to behave only as secondary factors that could only amplify the initial silt accumulation in summer rather than primary factors explaining spatial or long-term trends of sediment changes.

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The biomass and species composition of tropical phytoplankton in Albatross Bay, Gulf of Carpentaria, northern Australia, were examined monthly for 6 yr (1986 to 1992). Chlorophyll a (chl a) concentrations were highest (2 to 5.7 mu g l(-1)) in the wet season at inshore sites, usually coinciding with low salinities (30 to 33 ppt) and high temperatures (29 to 32 degrees C). At the offshore sites chi a concentrations were lower (0.2 to 2 mu g l(-1)) and did not vary seasonally. Nitrate and phosphate concentrations were generally low (0 to 3.68 mu M and 0.09 to 3 mu M for nitrate and phosphate respectively), whereas silicate was present in concentrations in the range 0.19 to 13 mu M. The phytoplankton community was dominated by diatoms, particularly at the inshore sites, as determined by a combination of microscopic and high-performance liquid chromatography (HPLC) pigment analyses. At the offshore sites the proportion of green flagellates increased. The cyanobacterium genus Trichodesmium and the diatom genera Chaetoceros, Rhizosolenia, Bacteriastrum and Thalassionema dominated the phytoplankton caught in 37 mu m mesh nets; however, in contrast to many other coastal areas studied worldwide there was no distinct species succession of the diatoms and only Trichodesmium showed seasonal changes in abundance. This reflects a stable phytoplankton community in waters without pulses of physical and chemical disturbances. These results are discussed in the context of the commercial prawn fishery in the Gulf of Carpentaria and the possible effect of phytoplankton on prawn larval growth and survival.

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Metanogeenit ovat hapettomissa oloissa eläviä arkkien pääryhmään kuuluvia mikrobeja, joiden ainutlaatuisen aineenvaihdunnan seurauksena syntyy metaania. Ilmakehässä metaani on voimakas kasvihuonekaasu. Yksi suurimmista luonnon metaanilähteistä ovat kosteikot. Pohjoisten soiden metaanipäästöt vaihtelevat voimakkaasti eri soiden välillä ja yhden suon sisälläkin, riippuen muun muassa vuodenajasta, suotyypistä ja kasvillisuudesta. Väitöskirjatyössä tutkittiin metaanipäästöjen vaihtelun mikrobiologista taustaa. Tutkimuksessa selvitettiin suotyypin, vuodenajan, tuhkalannoituksen ja turvesyvyyden vaikutusta metanogeeniyhteisöihin sekä metaanintuottoon kolmella suomalaisella suolla. Lisäksi tutkittiin ei-metanogeenisia arkkeja ja bakteereita, koska ne muodostavat metaanin tuoton lähtöaineet osana hapetonta hajotusta. Mikrobiyhteisöt analysoitiin DNA- ja RNA-lähtöisillä, polymeraasiketjureaktioon (PCR) perustuvilla menetelmillä. Merkkigeeneinä käytettiin metaanin tuottoon liittyvää mcrA-geeniä sekä arkkien ja bakteerien ribosomaalista 16S RNA-geeniä. Metanogeeniyhteisöt ja metaanintuotto erosivat huomattavasti happaman ja vähäravinteisen rahkasuon sekä ravinteikkaampien sarasoiden välillä. Rahkasuolta löytyi lähes yksinomaan Methanomicrobiales-lahkon metanogeeneja, jotka tuottavat metaania vedystä ja hiilidioksidista. Sarasoiden metanogeeniyhteisöt olivat monimuotoisempia, ja niillä esiintyi myös asetaattia käyttäviä metanogeeneja. Vuodenaika vaikutti merkittävästi metaanintuottoon. Talvella havaittiin odottamattoman suuri metaanintuottopotentiaali sekä viitteitä aktiivisista metanogeeneista. Arkkiyhteisön koostumus sen sijaan vaihteli vain vähän. Tuhkalannoitus, jonka tarkoituksena on edistää puiden kasvua ojitetuilla soilla, ei merkittävästi vaikuttanut metaanintuottoon tai -tuottajiin. Ojitetun suon yhteisöt kuitenkin muuttuivat turvesyvyyden mukaan. Vertailtaessa erilaisia PCR-menetelmiä todettiin, että kolmella mcrA-geeniin kohdistuvalla alukeparilla havaittiin pääosin samat ojitetun suon metanogeenit, mutta lajien runsaussuhteet riippuvat käytetyistä alukkeista. Soilla havaitut bakteerit kuuluivat pääjaksoihin Deltaproteobacteria, Acidobacteria ja Verrucomicrobia. Lisäksi löydettiin Crenarchaeota-pääjakson ryhmiin 1.1c ja 1.3 kuuluvia ei-metanogeenisia arkkeja. Tulokset ryhmien esiintymisestä hapettomassa turpeessa antavat lähtökohdan selvittää niiden mahdollisia vuorovaikutuksia metanogeenien kanssa. Tutkimuksen tulokset osoittivat, että metanogeeniyhteisön koostumus heijastaa metaanintuottoon vaikuttavia kemiallisia tai kasvillisuuden vaihteluita kuten suotyyppiä. Soiden metanogeenien ja niiden fysiologian parempi tuntemus voi auttaa ennustamaan ympäristömuutosten vaikutusta soiden metaanipäästöihin.

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Habitat requirements of fish are most strict during the early life stages, and the quality and quantity of reproduction habitats lays the basis for fish production. A considerable number of fish species in the northern Baltic Sea reproduce in the shallow coastal areas, which are also the most heavily exploited parts of the brackish marine area. However, the coastal fish reproduction habitats in the northern Baltic Sea are poorly known. The studies presented in this thesis focused on the influence of environmental conditions on the distribution of coastal reproduction habitats of freshwater fish. They were conducted in vegetated littoral zone along an exposure and salinity gradient extending from the innermost bays to the outer archipelago on the south-western and southern coasts of Finland, in the northern Baltic Sea. Special emphasis was placed on reed-covered Phragmites australis shores, which form a dominant vegetation type in several coastal archipelago areas. The main aims of this research were to (1) develop and test new survey and mapping methods, (2) investigate the environmental requirements that govern the reproduction of freshwater fish in the coastal area and (3) survey, map and model the distribution of the reproduction habitats of pike (Esox lucius) and roach (Rutilus rutilus). The white plate and scoop method with a standardized sampling time and effort was demonstrated to be a functional method for sampling the early life stages of fish in dense vegetation and shallow water. Reed-covered shores were shown to form especially important reproduction habitats for several freshwater fish species, such as pike, roach, other cyprinids and burbot, in the northern Baltic Sea. The reproduction habitats of pike were limited to sheltered reed- and moss-covered shores of the inner and middle archipelago, where suitable zooplankton prey were available and the influence of the open sea was low. The reproduction habitats of roach were even more limited and roach reproduction was successful only in the very sheltered reed-covered shores of the innermost bay areas, where salinity remained low (< 4‰) during the spawning season due to freshwater inflow. After identifying the critical factors restricting the reproduction of pike and roach, the spatial distribution of their reproduction habitats was successfully mapped and modelled along the environmental gradients using only a few environmental predictor variables. Reproduction habitat maps are a valuable tool promoting the sustainable use and management of exploited coastal areas and helping to maintain the sustainability of fish populations. However, the large environmental gradients and the extensiveness of the archipelago zone in the northern Baltic Sea demand an especially high spatial resolution of the coastal predictor variables. Therefore, the current lack of accurate large-scale, high-resolution spatial data gathered at exactly the right time is a considerable limitation for predictive modelling of shallow coastal waters.

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A new formulation of the stability of boundary-layer flows in pressure gradients is presented, taking into account the spatial development of the flow and utilizing a special coordinate transformation. The formulation assumes that disturbance wavelength and eigenfunction vary downstream no more rapidly than the boundary-layer thickness, and includes all terms nominally of order R(-1) in the boundary-layer Reynolds number R. In Blasius flow, the present approach is consistent with that of Bertolotti et al. (1992) to O(R(-1)) but simpler (i.e. has fewer terms), and may best be seen as providing a parametric differential equation which can be solved without having to march in space. The computed neutral boundaries depend strongly on distance from the surface, but the one corresponding to the inner maximum of the streamwise velocity perturbation happens to be close to the parallel flow (Orr-Sommerfeld) boundary. For this quantity, solutions for the Falkner-Skan flows show the effects of spatial growth to be striking only in the presence of strong adverse pressure gradients. As a rational analysis to O(R(-1)) demands inclusion of higher-order corrections on the mean flow, an illustrative calculation of one such correction, due to the displacement effect of the boundary layer, is made, and shown to have a significant destabilizing influence on the stability boundary in strong adverse pressure gradients. The effect of non-parallelism on the growth of relatively high frequencies can be significant at low Reynolds numbers, but is marginal in other cases. As an extension of the present approach, a method of dealing with non-similar flows is also presented and illustrated. However, inherent in the transformation underlying the present approach is a lower-order non-parallel theory, which is obtained by dropping all terms of nominal order R(-1) except those required for obtaining the lowest-order solution in the critical and wall layers. It is shown that a reduced Orr-Sommerfeld equation (in transformed coordinates) already contains the major effects of non-parallelism.

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Freshwater ecosystems vary in size and composition and contain a wide range of organisms which interact with each other and with the environment. These interactions are between organisms and the environment as nutrient cycling, biomass formation and transfer, maintenance of internal environment and interactions with the external environment. The range of organisms present in aquatic communities decides the generation and transfer function of biomass, which defines and characterises the system. These organisms have distinct roles as they occupy particular trophic levels, forming an interconnected system in a food chain. Availability of resources and competition would primarily determine the balance of individual species within the food web, which in turn influences the variety and proportions of the different organisms, with important implications for the overall functioning of the system. This dynamic and diverse relationship decides the physical, chemical and biological elements across spatial and temporal scales in the aquatic ecosystem, which can be recorded by regular inventorying and monitoring to maintain the integrity and conserve the ecosystem. Regular environmental monitoring, particularly water quality monitoring allows us to detect, assess and manage the overall impacts on the rivers. The appreciation of water quality is in constant flux. Water quality assessments derived through the biotic indices, i.e. assessments based on observations of the resident floral and faunal communities has gained importance in recent years. Biological evaluations provide a description of the water quality that is often not achievable from elemental analyses alone. A biological indicator (or bioindicator) is a taxon or taxa selected based on its sensitivity to a particular attribute, and then assessed to make inferences about that attribute. In other words, they are a substitute for directly measuring abiotic features or other biota. Bioindicators are evaluated through presence or absence, condition, relative abundance, reproductive success, community structure (i.e. composition and diversity), community function (i.e. trophic structure), or any combination thereof.Biological communities reflect the overall ecological integrity by integrating various stresses, thus providing a broad measure of their synergistic impacts. Aquatic communities, both plants and animals, integrate and reflect the effects of chemical and physical disturbances that occur over extended periods of time. Monitoring procedures based on the biota measure the health of a river and the ability of aquatic ecosystems to support life as opposed to simply characterising the chemical and physical components of a particular system. This is the central purpose of assessing the biological condition of aquatic communities of a river.Diatoms (Bacillariophyceae), blue green algae (Cyanophyceae), green algae (Chlorophyceae), and red algae (Rhodphyceae) are the main groups of algae in flowing water. These organisms are widely used as biological indicators of environmental health in the aquatic ecosystem because algae occupy the most basic level in the transfer of energy through natural aquatic systems. The distribution of algae in an aquatic ecosystem is directly related to the fundamental factors such as physical, chemical and biological constituents. Soft algae (all the algal groups except diatoms) have also been used as indicators of biological integrity, but they may have less efficiency than diatoms in this respect due to their highly variable morphology. The diatoms (Bacillariophyceae) comprise a ubiquitous, highly successful and distinctive group of unicellular algae with the most obvious distinguishing characteristic feature being siliceous cell walls (frustules). The photosynthetic organisms living within its photic zone are responsible for about one-half of global primary productivity. The most successful organisms are thought to be photosynthetic prokaryotes (cyanobacteria and prochlorophytes) and a class of eukaryotic unicellular algae known as diatoms. Diatoms are likely to have arisen around 240 million years ago following an endosymbiotic event between a red eukaryotic alga and a heterotrophic flagellate related to the Oomycetes.The importance of algae to riverine ecology is easily appreciated when one considers that they are primary producers that convert inorganic nutrients into biologically active organic compounds while providing physical habitat for other organisms. As primary producers, algae transform solar energy into food from which many invertebrates obtain their energy. Algae also transform inorganic nutrients, such as atmospheric nitrogen into organic forms such as ammonia and amino acids that can be used by other organisms. Algae stabilises the substrate and creates mats that form structural habitats for fish and invertebrates. Algae are a source of organic matter and provide habitat for other organisms such as non-photosynthetic bacteria, protists, invertebrates, and fish. Algae's crucial role in stream ecosystems and their excellent indicator properties make them an important component of environmental studies to assess the effects of human activities on stream health. Diatoms are used as biological indicators for a number of reasons: 1. They occur in all types of aquatic ecosystems. 2. They collectively show a broad range of tolerance along a gradient of aquatic productivity, individual species have specific water chemistry requirements. 3. They have one of the shortest generation times of all biological indicators (~2 weeks). They reproduce and respond rapidly to environmental change and provide early measures of both pollution impacts and habitat restoration. 4. It takes two to three weeks before changes are reflected to a measurable extent in the assemblage composition.

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Multi-year observations from the network of ground-based observatories (ARFINET), established under the project `Aerosol Radiative Forcing over India' (ARFI) of Indian Space Research Organization and space-borne lidar `Cloud Aerosol Lidar with Orthogonal Polarization' (CALIOP) along with simulations from the chemical transport model `Goddard Chemistry Aerosol Radiation and Transport' (GOCART), are used to characterize the vertical distribution of atmospheric aerosols over the Indian landmass and its spatial structure. While the vertical distribution of aerosol extinction showed higher values close to the surface followed by a gradual decrease at increasing altitudes, a strong meridional increase is observed in the vertical spread of aerosols across the Indian region in all seasons. It emerges that the strong thermal convections cause deepening of the atmospheric boundary layer, which although reduces the aerosol concentration at lower altitudes, enhances the concentration at higher elevations by pumping up more aerosols from below and also helping the lofted particles to reach higher levels in the atmosphere. Aerosol depolarization ratios derived from CALIPSO as well as the GOCART simulations indicate the dominance of mineral dust aerosols during spring and summer and anthropogenic aerosols in winter. During summer monsoon, though heavy rainfall associated with the Indian monsoon removes large amounts of aerosols, the prevailing southwesterly winds advect more marine aerosols over to landmass (from the adjoining oceans) leading to increase in aerosol loading at lower altitudes than in spring. During spring and summer months, aerosol loading is found to be significant, even at altitudes as high as 4 km, and this is proposed to have significant impacts on the regional climate systems such as Indian monsoon. (C) 2015 Elsevier Ltd. All rights reserved.

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We analyzed data from National Marine Fisheries Service bottom trawl surveys carried out triennially from 1984 to 1996 in the Gulf of Alaska (GOA). The continental shelf and upper slope (0–500 m) of the GOA support a rich demersal fish fauna dominated by arrowtooth flounder (Atheresthes stomias), walleye pollock (Theragra chalcogramma), Pacific cod (Gadus macrocephalus), Pacific halibut (Hippoglossus stenolepis), and Pacific Ocean perch (Sebastes alutus). Average catch per unit of effort (CPUE) of all groundfish species combined increased with depth and had a significant peak near the shelf break at 150–200 m. Species richness and diversity had significant peaks at 200–300 m. The western GOA was characterized by higher CPUEs and lower species richness and diversity than the eastern GOA. Highest CPUEs were observed in Shelikof Strait, along the shelf break and upper slope south of Kodiak Island, and on the banks and in the gullies northeast of Kodiak Island. Significant differences in total CPUE among surveys suggest a 40% increase in total groundfish biomass between 1984 and 1996. A multivariate analysis of the CPUE of 72 groundfish taxa revealed strong gradients in species composition with depth and from east to west, and a weak but significant trend in species composition over time. The trend over time was associated with increases in the frequency of occurrence and CPUE of at least eight taxa, including skates (Rajidae), capelin (Mallotus villosus), three flatfish species, and Pacific Ocean perch, and decreases in frequency of occurrence and CPUE of several sculpin (Myoxocephalus spp.) species. Results are discussed in terms of spatial and temporal patterns in productivity and in the context of their ecological and management implications.

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Samples from stone surfaces were collected in pools within four unpolluted hillstreams (two shaded and two unshaded) in monsoonal Hong Kong (lat. 23 degrees N) to elucidate the extent of spatial (within and among streams) and temporal (seasonal) variations in algal biomass and assemblage composition. Sampling continued for over 12 months, incorporating the dry season when streams were at baseflow, and the wet season when spates were frequent. We anticipated that algal biomass would be lower in shaded streams and during the wet season, with associated seasonal differences in assemblage composition or relative abundance of different growth forms (e. g. erect versus prostrate). Benthic chlorophyll a (a proxy for algal biomass) varied among streams from an annual mean of 11.0-22.3 mg m(-2). Dry-season standing stocks were 18% higher than during the wet season when spate-induced disturbance reduced algal standing stocks. Algal biomass varied significantly at the stream scale, but not at the pool scale, and was lower in unshaded streams, where standing stocks may have been limited by high densities of algivorous balitorid loaches (mainly Pseudogastromyzon myersi). An overriding effect of grazers on algal biomass could also have reduced variations resulting from spate-induced disturbance. Significant differences in assemblage composition among streams, which were dominated by diatoms and cyanobacteria (totally 82 taxa) were not systematically related to shading conditions. Seasonal variations in algal assemblages were statistically significant but rather minor, and did not involve major shifts in composition or growth form caused by spate-induced disturbance. The abundance of filamentous cyanobacteria in all the streams may have been due to 'gardening' by balitorid loaches that removed erect or stalked diatoms and favoured cyanobacteria that persist through basal regeneration of filaments. This explanation requires validation through manipulative experiments.

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This study consisted of sampling benthic algae at 32 sites in the Gangqu River, an important upstream tributary of the Yangtze River. Our aims were to characterize the benthic algae communities and relationships with environmental variables. Among the 162 taxa observed, Achnanthes linearis and Achnanthes lanceolata var. elliptica were the dominant species (17.10% and 14.30% of the total relative abundance, respectively). Major gradients and principal patterns of variation within the environmental variables were detected by principal component analysis (PCA). Then non-metric multidimensional scaling (NMS) divided all the sites into three groups, which were validated by multi-response permutation procedures (MRPP). Canonical correspondence analysis (CCA) indicated that three environmental variables (TN, TDS, and TP) significantly affected the distribution of benthic algae. Weighted averaging regression and cross-calibration produced strong models for predicting TN and TDS concentration, which enabled selection of algae taxa as potentially sensitive indicators of certain TN and TDS levels: for TN, Achnanthes lanceolata, Achnanthes lanceolata var. elliptica, and Cymbella ventricosa var. semicircularis; for TDS, Cocconeis placentula, Cymbella alpina var. minuta, and Fragilaria virescens. The present study represents an early step in establishing baseline conditions. Further monitoring is suggested to gain a better understanding of this region.

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The present study was conducted in Lake Donghu, a suburban eutrophic lake arising from the middle reaches of the Yangtze River, China. Food composition of 32 taxa of zoobenthos was analyzed from 1251 gut samples. Macroinvertebrate primary consumers ingested mainly detritus, sand grains and diatoms. The predators primarily preyed on rotifers, crustaceans, oligochaetes and chironomid larvae. The dietary overlap was relatively high among collector taxa but low among other macroinvertebrates. Food composition and dietary overlap of macroinvertebrates changed considerably, both spatially and temporally. Food web structure differed between inshore and offshore regions of Lake Donghu. The inshore web was relatively complex and dynamic whereas the offshore web was simple and stable. Taxon-specific changes of diet seem to have little effect on the benthic food web structure in offshore waters of a eutrophic lake.

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Benthic diatom communities were sampled monthly from May 2004 to May 2005 at four different sites in the littoral zone of Lake Donghu, a shallow eutrophic lake of China. The seasonal patterns of the total abundance, which were lowest in summer and highest in spring, were found at all sites. Total densities of diatom assemblages were significantly higher at hyper-eutrophic sites than at moderately eutrophic sites. Melosira varians was the most abundant species and dominant contributor to total abundance at all sites during spring, autumn and winter, whereas Achnanthes exigua dominated benthic diatom assemblages at the site with the highest nutrient concentrations during the summer. Achnanthes lanceolata var. dubia, Gomphonema parvulum, Navicula similis, Navicula verecunda and Nitzschia amphibia were generally observed at all sites throughout the year and were dominant at higher-nutrient sites. The abundance of ambient nutrients was probably responsible for the spatial variation in biomass, composition and diversity of benthic diatom assemblages, and lake water temperature was the major factor that controlled seasonal distribution.

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A gene-clone-library-based molecular approach was used to study the nirS-encoding bacteria-environment relationship in the sediments of the eutrophic Jiaozhou Bay. Diverse nirS sequences were recovered and most of them were related to the marine cluster I group, ubiquitous in estuarine, coastal, and marine environments. Some NirS sequences were unique to the Jiaozhou Bay, such as the marine subcluster VIIg sequences. Most of the Jiaozhou Bay NirS sequences had their closest matches originally detected in estuarine and marine sediments, especially from the Chesapeake Bay, indicating similarity of the denitrifying bacterial communities in similar coastal environments in spite of geographical distance. Multivariate statistical analyses indicated that the spatial distribution of the nirS-encoding bacterial assemblages is highly correlated with environmental factors, such as sediment silt content, NH4+ concentration, and OrgC/OrgN. The nirS-encoding bacterial assemblages in the most hypernutrified stations could be easily distinguished from that of the least eutrophic station. For the first time, the sedimentological condition was found to influence the structure and distribution of the sediment denitrifying bacterial community.

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Gridded sound speed data were calculated using Del Grosso's formulation from the temperature and salinity data at the PN section in the East China Sea covering 92 cruises between February 1978 and October 2000. The vertical gradients of sound speed are mainly related to the seasonal variations, and the strong horizontal gradients are mainly related to the Kuroshio and the upwelling. The standard deviations show that great variations of sound speed exist in the upper layer and in the slope zone. Empirical orthogonal function analysis shows that contributions of surface heating and the Kuroshio to sound speed variance are almost equivalent.