Biofilm diatom community structure: influence of temporal and substratum variability

Diatoms, which are early autotrophic colonisers, are an important constituent of the biofouling community in the marine environment. The effects of substratum and temporal variations on the fouling diatom community structure in a monsoon-influenced tropical estuary were studied. Fibreglass and glass coupons were exposed every month for a period of 4 days and the diatom population sampled at 24 h intervals, over a period of 14 months. The planktonic diatom community structure differed from the biofilm community. Pennate diatoms dominated the biofilms whilst centric diatoms were dominant in the water column. Among the biofilm diatoms, species belonging to the genera Navicula, Amphora, Nitzschia, Pleurosigma and Thalassionema were dominant. On certain occasions, the influence of planktonic blooms was also seen on the biofilm community. A comparative study of biofilms formed on the two substrata revealed significant differences in density and diversity. However species composition was almost constant. In addition to substratum variations, the biofilm diatom community structure also showed significant seasonal variations, which were attributed to physico-chemical and biological changes in both the water and substratum. Temporal variations in the tychopelagic diatoms of the water were also observed to exert an influence on the biofilm diatom community. Variations in diatom communities may determine the functional ecosystem of the benthic environment.     

Quantification of diatoms in biofilms: Standardisation of methods

Abstract

Benthic diatoms, which often dominate marine biofilms are mostly pennate along with a few centric species that have an attached mode of life. Even though the range of diatoms in biofilms is diverse, their ecology is poorly understood because of the difficulty in sampling and enumeration. Scraping or brushing are the traditional methods used for removal of diatoms from biofilms developed on solid substrata. The method of removal is the most critical step in enumerating the biofilm diatom community structure. In this study, a nylon brush and ceramic scraper were used as tools for the removal of diatoms from 1 – 4-day-old biofilms developed on fibreglass coupons and glass microscope slides. Standardisation of methods showed that the sample volume used in the analyses had the least influence on the quantification, whereas the method of removal was critical. The nylon brush was more efficient at recovering diatoms compared to a ceramic scraper. Direct microscopic enumeration of the community in the case of glass slides indicated that scraping resulted in between 30–50% underestimation. Heterogeneity in diatom community structure between replicate samples is one possible reason for such underestimation.     

Short-term response of monospecific and natural algal biofilms to copper exposure

The effect of copper additions (Cu ranging from 0 to 30?µM) on the photosynthesis of three different microalgal biofilms was studied to identify the factors that cause sensitivity differences between benthic and pelagic algae. The response of biofilms which colonized artificial substrata in the River Meuse was compared with those of two laboratory-grown monospecific biofilms, one consisting of the diatom Synedra ulna, and the other composed of a filament-forming cyanobacterium, Oscillatoria sp. The photosynthetic yield Φ_II (quantum efficiency of photosystem II) was studied with PAM (Pulse Amplitude Modulated) fluorimetry. S. ulna biofilms appeared to be the most sensitive to Cu, followed by the cyanobacteria, while natural biofilms, dominated by supposedly very sensitive diatom species such as Melosira varians and Diatoma vulgare, were the most resistant to Cu. In the highly productive biofilms, pH is suggested to play a role in lowering toxicity by helping the precipitation of cupric ions. Cu accumulation by the biofilms during the exposure period followed a linear relationship with Cu concentration, saturation not being observed; natural biofilms had an accumulation factor of 1–2.5?×?10³ relative to the concentrations in the water, while the diatoms growing unattached to the substratum had a higher concentration factor, up to 4.9?×?10³. It was concluded that the physical structure of the biofilm (package of cells and thickness), and not the species composition, was the main factor regulating the sensitivity of the biofilm to Cu toxicity during short-term exposures.     

Diatoms in Acid Mine Drainage and Their Role in the Formation of Iron-Rich Stromatolites

Adverse conditions in the acid mine drainage (AMD) system at the Green Valley mine, Indiana, limit diatom diversity to one species, Nitzschia tubicola. It is present in three distinct microbial consortia: Euglena mutabilis-dominated biofilm, diatomdominated biofilm, and diatom-exclusive biofilm. E. mutabilis dominates the most extensive biofilm, with lesser numbers of N. tubicola, other eukaryotes, and bacteria. Diatom-dominated biofilm occurs as isolated patches containing N. tubicola with minor fungal hyphae, filamentous algae, E. mutabilis, and bacteria. Diatom-exclusive biofilm is rare, composed entirely of N. tubicola.

Diatom distribution is influenced by seasonal and intraseasonal changes in water temperature and chemistry. Diatoms are absent in winter due to cool water temperatures. In summer, isolated patchy communities are present due to warmer water temperatures. In 2001, the diatom community expanded its distribution following a major rainfall that temporarily diluted the effluent, creating hospitable conditions for diatom growth. After several weeks when effluent returned to preexisting conditions, the diatom biofilm retreated to isolated patches, and E. mutabilis biofilm flourished.

Iron-rich stromatolites underlie the biofilms and consist of distinct laminae, recording spatial and temporal oscillations in physicochemical conditions and microbial activity. The stromatolites are composed of thin, wavy laminae with partially decayed E. mutabilis biofilm, representing microbial activity and iron precipitation under normal AMD conditions. Alternating with the wavy layers are thicker, porous, spongelike laminae composed of iron precipitated on and incorporated into radiating colonies of diatoms. These layers indicate episodic changes in water chemistry, allowing diatoms to temporarily dominate the system.     

Changes in diatom-dominated biofilms during simulated improvements in water quality: implications for diatom-based monitoring in rivers

Although benthic diatoms are used to assess river water quality, there are few data on the rate at which diatom assemblages react to changes in water quality. The aim of this study was to assess the reaction time of diatoms and to discuss the changes occurring during water quality improvement on the basis of their autecological characteristics. In order to simulate this improvement, diatom-dominated biofilms grown on artificial sandstone substrata were transferred from several polluted rivers to an unpolluted river. They were sampled three times: before transfer and 1 and 2 months after transfer. The ecology and growth-forms of the taxa explained most of the changes in species composition observed during the experiment. Adnate diatoms gradually replaced motile and stalked taxa. Gomphonema parvulum, a stalked diatom positioned vertically in the biofilm, is adapted for light and space competition in high-density algal biofilms. When transferred to an unpolluted site, this growth-form is less competitive and does not tolerate the high grazing pressure. Fistulifera saprophila is a single celled motile diatom, living in organic matrices. When the artificial substrata were transferred to the unpolluted site, this particular ecological niche disappeared quickly. On the other hand, Achnanthidium minutissimum, which is considered to be cosmopolitan and an early colonizer, increased during the first month of transfer and then decreased. It was gradually replaced by A. biasolettianum, which was the taxon best suited to this pristine stream. The changes observed differed between treatments depending on the species composition and architecture of the biofilms. In particular, biofilms dominated by stalked and motile diatoms were more quickly modified than those dominated by small motile diatoms. The diatom index reflects these changes, and its values showed that about 60 days following a water quality improvement were necessary for transferred diatom assemblages to reach diatom index values similar as those at the unpolluted river.     

Biofilm and capsule formation of the diatom Achnanthidium minutissimum are affected by a bacterium

Photoautotrophic biofilms play an important role in various aquatic habitats and are composed of prokaryotic and/or eukaryotic organisms embedded in extracellular polymeric substances (EPS). We have isolated diatoms as well as bacteria from freshwater biofilms to study organismal interactions between representative isolates. We found that bacteria have a strong impact on the biofilm formation of the pennate diatom Achnanthidium minutissimum. This alga produces extracellular capsules of insoluble EPS, mostly carbohydrates (CHO), only in the presence of bacteria (xenic culture). The EPS themselves also have a strong impact on the aggregation and attachment of the algae. In the absence of bacteria (axenic culture), A. minutissimum did not form capsules and the cells grew completely suspended. Fractionation and quantification of CHO revealed that the diatom in axenic culture produces large amounts of soluble CHO, whereas in the xenic culture mainly insoluble CHO were detected. For investigation of biofilm formation by A. minutissimum, a bioassay was established using a diatom satellite Bacteroidetes bacterium that had been shown to induce capsule formation of A. minutissimum. Interestingly, capsule and biofilm induction can be achieved by addition of bacterial spent medium, indicating that soluble hydrophobic molecules produced by the bacterium may mediate the diatom/bacteria interaction. With the designed bioassay, a reliable tool is now available to study the chemical interactions between diatoms and bacteria with consequences for biofilm formation.     

The effect of seasonality upon the development of lotic biofilms and microbial biostabilisation

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Hydrology and Diatom Phytoplankton of High Arctic Lakes and Ponds on Store Koldewey,Northeast Greenland

We document hydrological and phytoplankton characteristics of nine lakes and two ponds on Store Koldewey, a culturally undisturbed island off Northeast Greenland. The limnological survey included the recording of temperature, conductivity, oxygen concentration and saturation, pH, ionic composition, transparency, and the diatom phytoplankton community. In summer 2003, the lakes were cold, monomictic, thermally unstratified, alkaline and likely oligotrophic water bodies. Diatom phytoplankton was present in six lakes and consisted of four dominant species (Aulacoseira tethera, Cyclotella pseudostelligera, C. rossii, and Fragilaria tenera). The concentration of planktonic diatoms varied distinctly between the lakes. (© 2005 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

A novel biofilm channel for evaluating the adhesion of diatoms to non-biocidal coatings

Laboratory assessment of the adhesion of diatoms to non-toxic fouling-release coatings has tended to focus on single cells rather than the more complex state of a biofilm. A novel culture system based on open channel flow with adjustable bed shear stress values (0–2.4?Pa) has been used to produce biofilms of Navicula incerta. Biofilm development on glass and polydimethylsiloxane elastomer (PDMSe) showed a biphasic relationship with bed shear stress, which was characterised by regions of biofilm stability and instability reflecting cohesion between cells relative to the adhesion to the substratum. On glass, a critical shear stress of 1.3–1.4?Pa prevented biofilm development, whereas on PDMS, biofilms continued to grow at 2.4?Pa. Studies of diatom biofilms cultured on zwitterionic coatings using a bed shear stress of 0.54?Pa showed lower biomass production and adhesion strength on poly(sulfobetaine methacrylate) compared to poly(carboxybetaine methacrylate). The dynamic biofilm approach provides additional information to supplement short duration laboratory evaluations.     

The diatom assemblage of a marl core from Linton Loch

Summary

Diatom data from Linton Loch show five diatom assemblage zones, together with stratigraphic evidence indicating some aspects of the development of the lake basin during the Flandrian. Results show an early Flandrian dominance of planktonic diatom species in the lake waters with the later development of epiphytic species as the hydrosere encroached. Following deforestation in the catchment, water levels rose in the basin when benthic diatoms became dominant.     

Diatom community structure on in-service cruise ship hulls

Diatoms are an important component of marine biofilms found on ship hulls. However, there are only a few published studies that describe the presence and abundance of diatoms on ships, and none that relate to modern ship hull coatings. This study investigated the diatom community structure on two in-service cruise ships with the same cruise cycles, one coated with an antifouling (AF) system (copper self-polishing copolymer) and the other coated with a silicone fouling-release (FR) system. Biofilm samples were collected during dry docking from representative areas of the ship and these provided information on the horizontal and vertical zonation of the hull, and intact and damaged coating and niche areas. Diatoms from the genera Achnanthes, Amphora and Navicula were the most common, regardless of horizontal ship zonation and coating type. Other genera were abundant, but their presence was more dependent on the ship zonation and coating type. Samples collected from damaged areas of the hull coating had a similar community composition to undamaged areas, but with higher diatom abundance. Diatom fouling on the niche areas differed from that of the surrounding ship hull and paralleled previous studies that investigated differences in diatom community structure on static and dynamically exposed coatings; niche areas were similar to static immersion and the hull to dynamic immersion. Additionally, diatom richness was greater on the ship with the FR coating, including the identification of several new genera to the biofouling literature, viz. Lampriscus and Thalassiophysa. These results are the first to describe diatom community composition on in-service ship hulls coated with a FR system. This class of coatings appears to have a larger diatom community compared to copper-based AF systems, with new diatom genera that have the ability to stick to ship hulls and withstand hydrodynamic forces, thus creating the potential for new problematic species in the biofilm.     

西江下游浮游植物群落周年变化模式

西江肇庆段是珠江干流汇入珠三角河网水域的咽喉通道,对2009年该江段的浮游植物群落组成及变化进行系统阐析.调查期间共发现浮游植物7门,245种(包括变种、变型),其中硅藻和绿藻是浮游植物群落组成的最主要类群,分别占总种数的42.44％和34.69％,其次是裸藻和蓝藻.PCA分析结果显示,浮游植物各类群的种类丰富度和生物量的周年变化趋势主要受水温和径流量的影响.浮游植物种类丰富度的周年变化呈现明显的高温季节高,低温季节低的特征,除因大多数藻类物种直接喜好高温之外,径流量的增大有助于真浮游绿藻种类的外源注入及半浮游和偶然性浮游硅藻种类的增加.浮游植物生物量的周年变化呈现明显的双峰型,峰值分别出现在8月和11月.然而8月份第1个高峰出现之前,生物量波动不大,尽管水温的升高有助于生物量的增加,但是径流量增大所带来的稀释作用掩盖了水温上升对浮游植物生长的促进作用.真浮游植物生物量的变化趋势和数值与总浮游植物极其接近,主要得益于真浮游硅藻物种颗粒直链藻在总种群中的绝对优势地位.综上,水温升高对浮游植物种类丰富度和生物量的增长均有促进,径流量的增大虽然有助于种类丰富度的增加,但不利于生物量的增加.     

Cryosectioning of biofilms for microscopic examination

A method for rapid and minimally disruptive embedding and sectioning of bacterial biofilms has been developed and applied to binary population biofilms of Klebsiella pneumoniae and Pseudomonas aeruginosa grown on stainless steel surfaces in continuous flow annular reactors. Biofilms were cryoembedded using a commercial tissue embedding medium. Frozen embedded biofilms could be removed easily from the substratum by gently flexing the steel coupon. Microscopic examination of the substratum surface after biofilm removal indicated that less than a monolayer of attached cells remained. Five μm thick frozen sections were cut with a cryostat and examined by light or fluorescence microscopy. The cryoembedding technique preserved biofilm structural features including an irregular surface, water channels, local protrusions up to 500 μm thick, and a well‐defined substratum interface. The method requires minimal sample processing without dehydration or prolonged fixation, and can be completed in less than 24 h.     

Biofilm community structure and the associated drag penalties of a groomed fouling release ship hull coating

Grooming is a proactive method to keep a ship’s hull free of fouling. This approach uses a frequent and gentle wiping of the hull surface to prevent the recruitment of fouling organisms. A study was designed to compare the community composition and the drag associated with biofilms formed on a groomed and ungroomed fouling release coating. The groomed biofilms were dominated by members of the Gammaproteobacteria and Alphaproteobacteria as well the diatoms Navicula, Gomphonemopsis, Cocconeis, and Amphora. Ungroomed biofilms were characterized by Phyllobacteriaceae, Xenococcaceae, Rhodobacteraceae, and the pennate diatoms Cyclophora, Cocconeis, and Amphora. The drag forces associated with a groomed biofilm (0.75 ± 0.09 N) were significantly less than the ungroomed biofilm (1.09 ± 0.06 N). Knowledge gained from this study has helped the design of additional testing which will improve grooming tool design, minimizing the growth of biofilms and thus lowering the frictional drag forces associated with groomed surfaces.     

Quantification of diatoms in biofilms: standardisation of methods

Benthic diatoms, which often dominate marine biofilms are mostly pennate along with a few centric species that have an attached mode of life. Even though the range of diatoms in biofilms is diverse, their ecology is poorly understood because of the difficulty in sampling and enumeration. Scraping or brushing are the traditional methods used for removal of diatoms from biofilms developed on solid substrata. The method of removal is the most critical step in enumerating the biofilm diatom community structure. In this study, a nylon brush and ceramic scraper were used as tools for the removal of diatoms from 1 - 4-day-old biofilms developed on fibreglass coupons and glass microscope slides. Standardisation of methods showed that the sample volume used in the analyses had the least influence on the quantification, whereas the method of removal was critical. The nylon brush was more efficient at recovering diatoms compared to a ceramic scraper. Direct microscopic enumeration of the community in the case of glass slides indicated that scraping resulted in between 30-50% underestimation. Heterogeneity in diatom community structure between replicate samples is one possible reason for such underestimation.     

The impact of culture medium on the development and physiology of biofilms of Pseudomonas fluorescens formed on polyurethane paint

Microbial biofilms cause the deterioration of polymeric coatings such as polyurethanes (PUs). In many cases, microbes have been shown to use the PU as a nutrient source. The interaction between biofilms and nutritive substrata is complex, since both the medium and the substratum can provide nutrients that affect biofilm formation and biodeterioration. Historically, studies of PU biodeterioration have monitored the planktonic cells in the medium surrounding the material, not the biofilm. This study monitored planktonic and biofilm cell counts, and biofilm morphology, in long-term growth experiments conducted with Pseudomonas fluorescens under different nutrient conditions. Nutrients affected planktonic and biofilm cell numbers differently, and neither was representative of the system as a whole. Microscopic examination of the biofilm revealed the presence of intracellular storage granules in biofilms grown in M9 but not yeast extract salts medium. These granules are indicative of nutrient limitation and/or entry into stationary phase, which may impact the biodegradative capability of the biofilm.     

Optimal colonization and growth of marine benthic diatoms on artificial substrata: protocol for a routine use in bioindication

Benthic diatoms growing on hard substrata are used for their bioindication ability in freshwater quality monitoring. Artificial substrata are needed in cases where any natural substrate is present or to achieve similar sampling conditions between sites. Prior to use marine benthic diatoms for monitoring, a standardized protocol for sampling on artificial substrata must be set up. Two major types of information are required: (1) the time needed for a diatom community to be well developed and mature (climax stage); (2) the optimal growth conditions, given that the substrataum nature and texture are important parameters for the initial phase of biofilm development and can influence the future diatom assemblage. Three substrataum types were tested: frosted Plexiglass®, frosted glass, and rough enameled tiles. They were submerged for 8 weeks and sampled weekly. The experiment was conducted at five sites of distinct morphology and water chemistry, along the coastal area of Martinique Island, French West Indies. Development of diatom community was studied through biofilm dry weight, valve density, species richness, and species relative abundances. Globally, substratum type had no significant effect on any parameter. Frosted Plexiglass® was found to be the most interesting substratum because of higher valve densities and practical use. The asymptotic phase of biofilm development was encountered between 5 and 8 weeks depending on site and parameter. A compromise between community development and vandalism or loss through time was fixed to 5 weeks. This period is longer than for stream environments and is valid for tropical oligotrophic marine environments.     

Seasonal variations in the fouling diatom community structure from a monsoon influenced tropical estuary

Seasonal variations in the fouling diatom community from a monsoon influenced tropical estuary were investigated. The community composition did not differ significantly between stainless steel and polystyrene substrata due to dominance by Navicula spp. The experimental evidence suggests that Amphora, which is a dominant fouling diatom in temperate waters, ultimately dominates the community in tropical regions when conditions are favourable. These observations reveal that a faster onset of macrofouling interferes with the microfouling community wherein the faster recruiters that have a higher abundance in ambient waters, dominate the community. Seasonal variations were observed in the fouling diatom community. Navicula delicatula dominated during the post-monsoon and tychopelagic diatoms (Melosira and Odontella) were observed during the monsoon. Low diatom abundance was recorded during the pre-monsoon season. The results indicate that although the fouling diatom community composition does not vary between substrata, there is a seasonal change in the community depending on the physical, chemical and biological interactions.     

Spatial and temporal variations in moss-inhabiting summer diatom communities from Potter Peninsula (King George Island, Antarctica)

This work represents the first analysis of the spatial and temporal variations in the diversity and community structure of diatoms associated with mosses, either dry or wet, sampled during the summer season from the littoral zone of three ponds in Potter Peninsula (King George Island, South Shetland Islands). It also seeks to explore the effect moisture content in the mosses can have on the structure of the diatom assemblages. The ponds approximately had the same number of species, and differences were found between the dry and wet sampling points. The same 4 species were predominant in the three ponds (Nitzschia alpina, N. perminuta, Staurosira oldenburgiana, and Pinnularia subantarctica var. elongata), but not in the same proportions. The first three species differed from those which were found to be dominant in other basins covered by mosses in southern South America and in the sub-Antarctic region. The latter, P. subantarctica var. elongata, had only been found to be abundant in a moss sample from another South Shetland Island. The diatom assemblages were found to be more strongly related to habitat traits (e.g. distance to the sea, ponds’ areas, and the vegetation’s moisture levels) than to the physical–chemical variables measured in the water.     

Dynamics of diatom colonization process in some rivers influenced by urban pollution (Hanoi, Vietnam)

Periphytic diatom communities’ colonization patterns were studied at three sampling stations of the Red–Nhue–Tolich hydrosystem presenting different urban pollution levels by using artificial substrates for 6 weeks in dry season 2005. Structural characteristics of periphytic diatoms developed on glass substrates at each sampling site were followed and compared. This experiment showed, through various general criteria (total diatom density, dry weight biomass) and specific criteria (relative diatom abundances, indices) that the structure of benthic diatoms developed on these substrates was strongly affected by pollution as early as the second week of colonization. Communities exposed to heavily and moderately polluted sites contained species which are known to be saprophilous or tolerant to organic pollution such as Nitzschia umbonata, Nitzschia palea, Cyclotella meneghiniana, Eolimna minima. Growth inhibition of diatom communities at the heavily polluted site was mostly related to a strong increase of organic load rather than to low metallic input, though metallic (Cd and Zn) burdens have been successfully quantified in the biofilms developed at the three studied sites. Nevertheless, no significant difference in species richness and diversity index between colonization duration times was observed. Based on values of diatom indices Indice de Polluosensibilité (IPS) and diatom assemblage index to organic water pollution (DAIPo), water quality could be classified as medium quality at Red site, polluted at NT₂ and heavily polluted at TL. Thus, the use of diatoms as a tool for water assessment appears suitable for monitoring rivers in Vietnam, as it is in several other countries.