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PubMed2004

What drives bacteria to produce a biofilm?

Nearly 40 years ago, Dr. R.J. Gibbons made the first reports of the clinical relevance of what we now know as bacterial biofilms when he published his observations of the role of polysaccharide glycocalyx formation on teeth by Streptococcus mutans [Sci. Am. 238 (1978) 86]. As the clinical relevance of bacterial biofilm formation became increasingly apparent, interest in the phenomenon exploded. Studies are rapidly shedding light on the biomolecular pathways leading to this sessile mode of growth but many fundamental questions remain. The intent of this review is to consider the reasons why bacteria switch from a free-floating to a biofilm mode of growth. The currently available wealth of data pertaining to the molecular genetics of biofilm formation in commonly studied, clinically relevant, single-species biofilms will be discussed in an effort to decipher the motivation behind the transition from planktonic to sessile growth in the human body. Four potential incentives behind the formation of biofilms by bacteria during infection are considered: (1) protection from harmful conditions in the host (defense), (2) sequestration to a nutrient-rich area (colonization), (3) utilization of cooperative benefits (community), (4) biofilms normally grow as biofilms and planktonic cultures are an in vitro artifact (biofilms as the default mode of growth).

Adaptation, Physiological↗
PubMed2004

Identification of oligopeptide permease (opp) gene cluster in Vibrio fluvialis and characterization of biofilm production by oppA knockout mutation.

Oligopeptides play important roles in bacterial nutrition and signaling. The oligopeptide permease (opp) gene cluster was cloned from Vibrio fluvialis. The V. fluvialis opp operon encodes five proteins: OppA, B, C, D and F. The deduced amino acid sequence of these proteins showed high similarity with those from other Gram-negative bacteria. To investigate whether OppA is involved in biofilm production, an oppA knockout mutant was constructed by homologous recombination. The oppA mutant produced more abundant biofilm than the wild type in BHI medium. When both strains were grown in minimal medium, we could not detect biofilm formation. However, it was found that the biofilm productivity of the oppA mutant was two folds greater than that of the wild type in minimal medium containing peptone or tryptone. This variation in biofilm production was demonstrated by scanning electron microscopy (SEM). In minimal medium containing C-sources, both strains produced some biofilm without significant difference in the biofilm productivity. Complementation of oppA gene with the plasmid pOAC2, which contains oppA ORF plus promoter regions, was sufficient to restore growth rate and biofilm to the wild type. These results suggest that the OppA protein is involved in uptake of peptides and affects biofilm productivity.

Amino Acid Sequence↗
PubMed2004

Interactions in biofilms between Listeria monocytogenes and resident microorganisms from food industry premises.

Twenty nine bacterial strains were grown as binary culture biofilms with Listeria monocytogenes to assess their influence on the settlement of the latter on stainless steel coupons. Most of the strains had been isolated from food processing plants after cleaning and disinfection and were tentatively identified by the APILAB Plus 3.3.3 database (bioMerieux). Sixteen of them decreased L. monocytogenes biofilm colony forming units (CFU) counts. Three strains, Bacillus sp. CCL 9 an unidentified Gram-positive strain CCL 59 and Pseudomonas fluorescens E9. 1, led to a 3-log difference in CFU counts between the pure L. monocytogenes biofilms and the mixed biofilms. Eleven strains had no effect and only four, Kocuria varians CCL 73, Staphylococcus capitis CCL 54, Stenotrophomonas maltophilia CCL 47 and Comamonas testosteroni CCL 24, had a positive effect, with a 0.5- to 1.0-log increase in the L. monocytogenes biofilm CFU counts. On its own, L. monocytogenes settled as single cells, but in binary biofilms, different spatial arrangements were observed: (i) with K. varians CCL 73, K. varians CCL 56 and S. capitis CCL 54, L. monocytogenes cells gathered around the microcolonies of the partner strain; (ii) with the two Gram-negative strains, C. testosteroni CCL 24 and CCL 25, L. monocytogenes cells formed its own microcolonies. No link could be found between the exopolysaccharide production capacity of the bacterial strains in pure-culture biofilms and their effect on the L. monocytogenes population in mixed biofilms. With one strain, C. testosteroni CCL 24, adding filter-sterilized supernatant from a pure-culture biofilm to a pure culture of L. monocytogenes increased the number of L. monocytogenes cells adhering to the stainless steel coupons and forming microcolonies. This study suggests that the "house flora" can have a strong effect on the likelihood of finding L. monocytogenes on inert surfaces.

Bacteria↗
PubMed2006

Compositions of maple sap microflora and collection system biofilms evaluated by scanning electron microscopy and denaturing gradient gel electrophoresis.

The bacterial microflora of maple sap and biofilms in collection system tubing were studied through the use of bacterial counts, scanning electron microscopy (SEM) of surfaces and the analysis of 16S rRNA gene by denaturing gradient gel electrophoresis (DGGE). Samples were taken at five times during the 2002 and 2003 seasons in order to follow the changes in the microflora of this complex ecosystem. Bacterial counts showed the growth of bacterial populations as the season advanced. These populations were mainly composed of psychrotrophic bacteria and Pseudomonas spp. SEM results confirmed the suspected presence of biofilms on the inner surfaces of tubing samples. Bacterial colonization and biofilm formation progressively increased during the season for both lateral and main line surfaces, and biofilms were mainly composed of rod shape bacteria. The bacterial microflora profiles obtained for sap and corresponding biofilm by DGGE showed up to 12 major bands. The Shannon-Weaver index of diversity (H) calculated from DGGE bands were statistically higher for sap samples compared to biofilm. The diversity index was relatively stable or increasing for lateral line sap and biofilm samples during the season while the diversity index for sap and biofilm samples of the main line showed a decreasing profile as the season progressed. Sequence analysis of major DGGE bands revealed the predominance of bacteria from the genera Pseudomonas, Rahnella and another, unidentified genus. The results describe the composition of sap collection system microflora as well as the formation of biofilms and will be useful for further studies on factors affecting maple product quality.

Acer↗
PubMed2006

Biofilm formation and the survival of Salmonella Typhimurium on parsley.

Although several studies provide evidence that the formation of biofilms by human pathogens on plant tissue is possible, to date there is no direct evidence that biofilms enhance the resistance of plant-associated pathogens to disinfectants or biocides. We hypothesized that biofilm formation would enhance the adhesion and survival of Salmonella on leafy vegetables. To test our hypothesis, we compared the adhesion and persistence of Salmonella Typhimurium and its biofilm-deficient isogenic mutant. Following inoculation of parsley and rinsing with water or chlorine solution, both strains had similar survival properties, and up to 3-log reduction were observed, depending on chlorine concentration. This indicates that the biofilm matrix of Salmonella likely does not play a significant role in initial adhesion and survival after disinfection. After a week of storage the biofilm producing strain survived chlorination significantly better than the biofilm-deficient mutant. However, the recovery of the mutant was still elevated, indicating that although the biofilm matrix has a role in persistence of Salmonella after chlorination treatment of parsley, this is not the most important mechanism, and other mechanisms, probably the ability to penetrate the plant tissue or the pre-existing biofilms, or production of different polysaccharides other than cellulose, provide the protection.

Analysis of Variance↗
PubMed2006

Interaction between GIC and S. sanguis biofilms: antibacterial properties and changes of surface hardness.

OBJECTIVE: The aim of this study was to investigate the interaction of Streptococcus sanguis with two glass-ionomer formulations (GIC:A containing fluoride and GIC:B without fluoride) with particular reference to bacterial growth and changes in hardness of the cement with respect to time. MATERIALS AND METHODS: Discs of two water activated glass-ionomer cements were prepared according to the manufacturer's instruction. Hydroxyapatite discs (HA) were used as controls. 3D laser scanning technique was used to characterize surface roughness and area of the substrate prior to growing biofilms. Surface hardness was evaluated before and after biofilm growth. A constant depth film fermenter system (CDFF) was used to grow S. sanguis biofilms on the specimens in a similar manner to that described previously by Wilson et al. in 1995. For susceptibility measurement, specimens were removed from CDFF aseptically over periods up to 14d after the first colonization with bacteria. Counts of viable bacterial in the accumulating biofilm layer on each surface were measured and converted to colony forming units per unit surface area. To determine the effect of storage media, hardness discs were exposed to distilled water, lactic acid pH 4, lactic acid pH 5, citric acid pH 5, artificial saliva and S. sanguis biofilms. Twenty-four hours after preparing and subsequent autoclaving, specimens were transferred to a vessel containing 40 ml storage medium. The specimens were investigated for periods up to 7d. RESULTS: The viable counts of S. sanguis per mm2 on GIC:A were significantly less than those on HA and GIC:B during the first 5d (p<0.05). The viable counts of bacteria on the surface of GIC:B were lower during the initial 5d when compared to HA. Exposure of GIC:A and GIC:B to different medium produced softening to the surface of cement. It is apparent that the effects of the biofilms are significantly greater than storage in water but similar to storage in lactic acid pH 5. CONCLUSIONS: This investigation showed that the growth of S. sanguis biofilms were significantly affected by both glass-ionomer formulations, the greater reduction being noted on the surface of the fluoride containing GIC. S. sanguis biofilms produced reduction on the surface hardness of the cement equivalent to that seen after immersion in lactic acid at pH 5. This indicates that while S. sanguis biofilm is affected by the GIC, there is also a decrease in hardness of the cement indicating some cement degradation.

Analysis of Variance↗
PubMed2006

Bacterial biofilms within the clinical setting: what healthcare professionals should know.

Bacterial biofilm formation is the prevailing microbial lifestyle in natural and manmade environments and occurs on all surface types. Biofilm formation develops in several phases and is influenced by various parameters, both environmental and inherent to the attaching cell. Biofilms also serve as protective niches for particular pathogens when outside a host. Although it is accepted that biofilms are ubiquitous in nature, the significance of biofilms in clinical settings, especially with regard to their role in medical-related infections, is often underestimated. It has been found that several aspects of human pathogenesis within a clinical context are directly related to biofilm development. Various types of surfaces in clinical settings are prone to biofilm development and an increased risk of disease may be a direct consequence of their formation. This review describes the process of biofilm formation, highlights the importance of bacterial associations with surfaces in clinical settings and describes various methods for biofilm visualization and control.

Bacteria↗
PubMed2006

Secretory IgA and mucin-mediated biofilm formation by environmental strains of Escherichia coli: role of type 1 pili.

Recent studies suggest the importance of secretory IgA (SIgA) and mucin in the mediation of biofilm formation by commensal bacteria within the mammalian gut. Studies using a variety of strains of Escherichia coli have indicated that the interaction between E. coli and SIgA is dependent on the type 1 pilus. In this study, the importance of the pilus in SIgA-mediated biofilm formation by a laboratory strain (MG1655) and environmental (fecal) strains of E. coli was evaluated. Transient expression of the type 1 pilus by the laboratory strain of E. coli failed to facilitate SIgA-mediated biofilm formation, whereas constitutive expression of the type 1 pilus by the laboratory strain was sufficient. In contrast, transient expression of the type 1 pilus was sufficient to facilitate SIgA-mediated biofilm formation by environmental isolates. The "threshold" for mucin-mediated biofilm formation appeared to be lower than that for SIgA-mediated biofilm formation, perhaps reflecting disparate roles of mucin and SIgA in mediating biofilm formation in the gut. These studies also provide the first procedures for the growth of bacterial biofilms on live epithelial cells in vitro, an important development that may facilitate future studies on the effects of bacterial biofilms on epithelial cells.

Adenocarcinoma↗
PubMed2006

Opening plugged tympanostomy tubes: effect of biofilm formation.

OBJECTIVE: To determine if the presence of a bacterial biofilm impacts the rate of clearing of mucoid plugs from tympanostomy tubes (TTs). STUDY DESIGN AND SETTING: Ex vivo model. Stainless steel Reuter Bobbin TTs (n = 18) were placed in growth medium with Pseudomonas aeruginosa and Streptococcus pneumoniae for 12 days to promote biofilm formation. Tympanostomy tubes (n = 18) placed in growth medium, without bacteria, for 12 days served as controls. Biofilm formation was assessed by scanning electron microscopy. All TTs were filled with middle ear mucus and allowed to dry, thereby forming a plug. TTs were placed in a model ear chamber, covered with ofloxacin otic solution, and the time to clear each plug was recorded. RESULTS: Biofilm formation was consistently encountered on TTs exposed to bacteria but in no TTs in the control group. There was a significant effect of the biofilm on plug clearance, favoring TTs without a biofilm (P = 0.0333). Although there was no significant difference in the proportion of unplugged TTs (P = 0.264), TTs with a biofilm did not clear plugs as rapidly as TTs without a biofilm (P = 0.0416). CONCLUSIONS: The presence of a biofilm may slow the time to clear mucoid TT plugs, but it does not seem to affect the overall proportion of TTs that are unplugged.

Anti-Bacterial Agents↗
PubMed2005

Quantitative analysis of adhesion and biofilm formation on hydrophilic and hydrophobic surfaces of clinical isolates of Staphylococcus epidermidis.

Staphylococcus epidermidis is now well established as a major nosocomial pathogen associated with infections of indwelling medical devices. The major virulence factor of these organisms is their ability to adhere to devices and form biofilms. However, it has not been established that adherence and biofilm formation are closely linked phenotypes for clinical isolates. In this study, the initial adhesion to different materials (acrylic and glass) of 9 clinical isolates of S. epidermidis, along with biofilm-positive and biofilm-negative control strains, was assayed using physico-chemical interactions to analyze the basis for bacterial adherence to the substratum. X-ray photo electron spectroscopy (XPS) analysis of the cell surface elemental composition was also performed in an attempt to find a relationship between chemical composition and adhesion capabilities. Biofilm formation on the two surfaces was evaluated by dry weight measurements. Human erythrocytes were used to evaluate the ability of S. epidermidis strains to cause hemagglutination, an indicator of the production of a poly-N-acetyl glucosamine cell surface polysaccharide also involved in biofilm formation. The clinical isolates exhibited different cell wall physico-chemical properties, resulting in differing abilities to adhere to surfaces. Adhesion to hydrophobic substrata for all strains occurred to a greater extent than that to hydrophilic surfaces. Bacterial cell hydrophobicity seemed to have little or no influence on adhesion. X-ray photoelectron spectroscopy analysis showed a high ratio of oxygen/carbon for all strains, which is a common characteristic of S. epidermidis species. No relevant relationship was found between XPS data and adhesion values. All strains forming biofilms were able to agglutinate erythrocytes. However, no direct relationship was found between the amount of biofilm formed and the initial adhesion extent. These results indicate that high levels of initial adherence do not necessarily lead to thick biofilm formation. These two aspects of the pathogenesis of medical device related-infection may need to be evaluated independently to ascertain the contribution of each to the virulence of S. epidermidis causing device-related infections.

Acrylates↗
PubMed2005

Survival strategies of infectious biofilms.

Modern medicine is facing the spread of biofilm-related infections. Bacterial biofilms are difficult to detect in routine diagnostics and are inherently tolerant to host defenses and antibiotic therapies. In addition, biofilms facilitate the spread of antibiotic resistance by promoting horizontal gene transfer. We review current concepts of biofilm tolerance with special emphasis on the role of the biofilm matrix and the physiology of biofilm-embedded cells. The heterogeneity in metabolic and reproductive activity within a biofilm correlates with a non-uniform susceptibility of enclosed bacteria. Recent studies have documented similar heterogeneity in planktonic cultures. Nutritional starvation and high cell density, two key characteristics of biofilm physiology, also mediate antimicrobial tolerance in stationary-phase planktonic cultures. Advances in characterizing the role of stress response genes, quorum sensing and phase variation in stationary-phase planktonic cultures have shed new light on tolerance mechanisms within biofilm communities.

Adaptation, Physiological↗
PubMed2004

Denture stomatitis: a role for Candida biofilms.

OBJECTIVE: To assess the contribution of Candida biofilms to the etiology of denture stomatitis. STUDY DESIGN: Samples of denture acrylic were retrieved from patients with denture stomatitis and subjected to scanning electron microscopy (SEM) analysis. Oral swab and swish samples were taken from the same group of patients and representative C albicans isolates recovered were used to investigate the kinetics of biofilm development in vitro. RESULTS: Candida biofilms could be visualized by SEM directly from denture samples from patients with denture stomatitis. These biofilms showed a propensity to adhere along cracks and imperfections of the denture acrylic. C albicans clinical isolates were able to form biofilms in vitro, although differences in the extent of biofilm formation were observed for different isolates recovered from the same patient. Susceptibility testing indicated that the resulting biofilms showed increased resistance to antifungal treatment. Presence of serum and saliva conditioning films increased the initial adherence of selected isolates but had little effect in overall biofilm formation. CONCLUSIONS: Candida biofilms play a role in denture stomatitis.

Acrylic Resins↗
PubMed2004

Influence of growth history on sloughing and erosion from biofilms.

The development of biofilms is determined by the balance of growth and detachment. But while the growth of biofilms is well studied, the influence of growth history and detachment on biofilm development is not. Here we report on laboratory scale experiments where heterotrophic biofilms were grown in a tubular reactor. Biofilm detachment was categorized based on particle size as erosion or sloughing. Erosion results in small particles and was approximated by the effluent suspended solids while sloughing was determined from the larger pieces of biomass that settled in a mixing tank. It was found that for all experiments, overall detachment was a combination of erosion and sloughing where erosion had a slightly larger contribution to the overall solids removal. However, sloughing had a significant influence on the biofilm morphology. Once the smooth biofilm surface was disturbed by a sloughing event (e.g., initiated through increasing liquid shear in the reactor), the biofilm became unstable resulting in spontaneous sloughing during subsequent operation. We propose that experimental investigations should consider sloughing events as an integral part of biofilm development rather than a disturbance.

Biofilms↗
PubMed2006

Dynamics of drinking water biofilm in flow/non-flow conditions.

Drinking water biofilm formation on polyvinyl chloride (PVC), cross-linked polyethylene (PEX), high density polyethylene (HDPE) and polypropylene (PP) was followed in three different reactors operating under stagnant or continuous flow regimes. After one week, a quasi-steady state was achieved where biofilm total cell numbers per unit surface area were not affected by fluctuations in the concentration of suspended cells. Metabolically active cells in biofilms were around 17-35% of the total cells and 6-18% were able to form colony units in R(2)A medium. Microbiological analysis showed that the adhesion material and reactor design did not affect significantly the biofilm growth. However, operating under continuous flow (0.8-1.9 Pa) or stagnant water had a significant effect on biofilm formation: in stagnant waters, biofilm grew to a less extent. By applying mass balances and an asymptotic biofilm formation model to data from biofilms grown on PVC and HDPE surfaces under turbulent flow, specific growth rates of bacteria in the biofilm were found to be similar for both materials (around 0.15 day(-1)) and much lower than the specific growth rates of suspended bacteria (around 1.8 day(-1)).

Biofilms↗
PubMed2002

Minimizing biofilm in the presence of iron oxides and humic substances.

Based upon circumstantial evidence linking elevated coliform bacteria counts in drinking water distribution systems with unlined cast iron pipe, it was hypothesized that adsorption of humic substances by iron oxide containing corrosion products (CPs) can stimulate and/or support biofilm development. Using porous media consisting of iron-oxide-coated glass beads (IOCBs) or actual iron CPs, experiments were performed to evaluate the effectiveness of different corrosion control and disinfection treatments in reducing biofilm when humic substances were the carbon source. Free chlorine was the most effective treatment in minimizing biofilm. Addition of phosphate alone did not significantly reduce biofilm using the CPs, but there was weak evidence it did using the IOCBs. The combination of free chlorine and phosphate was more effective at minimizing biofilm than free chlorine alone when CPs were the media. The presence of humic substances was a major factor when considering biofilm minimization based on results of experiments using both types of iron oxide media. The combination of humic substances and CPs led to an increase in biofilm biomass when free chlorine was not present, similar to conditions that could occur at distribution system dead-ends. Treatment to raise the pH to 9 did not reduce biofilm in experiments using both media, and actually increased biofilm in the experiment using CPs under the conditions tested.

Bacteria↗
PubMed2001

An in vitro model for studying the contributions of the Streptococcus mutans glucan-binding protein A to biofilm structure.

The method described here for analyzing biofilms was sensitive enough to allow the detection of differences formed by pure cultures of S. mutans or a GbpA knockout strain. Other strains have also been tested, and the differences in biofilm structure were sometimes even more extensive (data not shown). The advantages of this method are that it is quick, inexpensive, and adaptable to almost any laboratory setting. The constant rotation of the cultures, which was employed to simulate salivary flow, appears to be a critical element for establishing biofilm differences. An analysis of protein profiles confirmed that the biofilm bacteria were metabolically distinct from the planktonic phase bacteria. For the strains tested, the variations in biofilm architecture could be visualized with or without magnification. Staining of the bacteria was not required, though we typically stained the biofilms with either crystal violet or Schiff's reagent. Altogether, this in vitro method for generating biofilms allowed the evaluation of visual, quantitative (confocal microscopy), and functional (antimicrobial susceptibility) differences. We have employed these methods in a reductionist approach to understanding the contribution of individual proteins to dental plaque development. These methods may also be useful in the screening of mutants that would be of greatest for testing in multispecies biofilms, animal models, or more complex biofilm models.

Bacterial Proteins↗
PubMed2003

Stimulation of Staphylococcus epidermidis growth and biofilm formation by catecholamine inotropes.

BACKGROUND: Bacterial colonisation of indwelling medical devices by coagulase-negative staphylococci is a prevalent risk in intensive-care units. Factors determining biofilm formation and progression to catheter- related infection are incompletely understood. We postulated that administration of inotropic agents via indwelling intravenous catheters may stimulate growth and formation of biofilms by Staphylococcus epidermidis. METHODS: Inocula representing physiologically relevant infecting doses of S epidermidis were incubated in a minimum medium supplemented with fresh human plasma in the presence or absence of pharmacological concentrations of norepinephrine or dobutamine. Biofilm formation on polystyrene and medical-grade silicone was examined. After incubation, cultures were assessed for growth and formation of biofilms by colony counting and scanning electronmicroscopy. The production of exopolysaccharide, a major constituent of S epidermidis biofilms, was also assessed by use of immunofluorescence microscopy. FINDINGS: Incubation of S epidermidis with catecholamine inotropes in the presence of human plasma resulted in a significant increase in growth compared with control on both polystyrene and silicone surfaces, with pronounced increases in biofilm formation as visualised by scanning electronmicroscopy. Experiments with transferrin labelled with radioactive iron showed the ability of catecholamine inotropes to facilitate acquisition of iron by S epidermidis. Immunofluorescence microscopy revealed extensive exopolysaccharide production associated with S epidermidis biofilms. INTERPRETATION: The ability of catecholamine inotropic drugs to stimulate bacterial proliferation and biofilm formation may be an aetiological factor in the development of intravascular catheter colonisation and catheter-related infection. The removal of iron from transferrin for subsequent use by S epidermidis is a possible mechanism by which catecholamine inotropes stimulate bacterial growth as biofilms.

Adult↗
PubMed2001

Staphylococcus aureus biofilm formation on different gentamicin-loaded polymethylmethacrylate bone cements.

In this in vitro study, the formation of a Staphylococcus aureus biofilm on six gentamicin-loaded bone cements (CMW1, CMW3, CMW Endurance, CMW2000, Palacos, and Palamed) was determined in a modified Robbins device over a 3 days time span and related with previously (Van de Belt et al., Biomaterials 21 (2000) 1981) measured kinetics of antibiotic release by these cement brands. The influence of gentamicin release on biofilm formation was quantified by expressing the number of colony-forming units on gentamicin-loaded cement relative to the number of viable organisms on unloaded cement of the same brand. Biofilms formed on all gentamicin-loaded cements, despite the release of antibiotics, followed a consistent pattern in time with a maximum number of colony-forming units per unit cement area found between 24 and 30 h after inoculation. None of the gentamicin-loaded cements showed a reduction in biofilm formation relative to unloaded cements within 6 h after inoculation, whereas only gentamicin-loaded CMW1 and Palacos reduced biofilm formation 24 h after inoculation. Alternatively, CMW Endurance, CMW2000, and Palamed did not exhibit any initial reductions in biofilm formation, but effects started after 72, 48, and 72 h, respectively. Biofilm reduction by gentamicin-loaded CMW3 lasted the longest from 24 to 72 h. Interestingly, each cement seemed to have a different "window-of-effectiveness" with regard to reduction in biofilm formation that did not relate with the gentamicin-release kinetics. Summarising, this study demonstrates that although gentamicin loading of bone cements yields reductions in biofilm formation, S. aureus is able to grow on gentamicin-loaded bone cements.

Biocompatible Materials↗