Note 1: A biofilm is a system that can be adapted internally to
environmental conditions by its inhabitants.
Note 2: The self-produced matrix of extracellular polymeric
substances, which is also referred to as slime, is a polymeric
conglomeration generally composed of extracellular biopolymers in
various structural forms..
A biofilm comprises any group of microorganisms in which cells stick
to each other and often also to a surface. These adherent cells
become embedded within a slimy extracellular matrix that is composed
of extracellular polymeric substances (EPS). The cells within
the biofilm produce the EPS components, which are typically a
polymeric conglomeration of extracellular polysaccharides, proteins
and DNA. Because they have three-dimensional structure and
represent a community lifestyle for microorganisms, they have been
metaphorically described as "cities for microbes".
Biofilms may form on living or non-living surfaces and can be
prevalent in natural, industrial and hospital settings. The
microbial cells growing in a biofilm are physiologically distinct from
planktonic cells of the same organism, which, by contrast, are
single-cells that may float or swim in a liquid medium. Biofilms
can form on the teeth of most animals as dental plaque, where they may
cause tooth decay and gum disease.
Microbes form a biofilm in response to various different factors,
which may include cellular recognition of specific or non-specific
attachment sites on a surface, nutritional cues, or in some cases, by
exposure of planktonic cells to sub-inhibitory concentrations of
antibiotics. When a cell switches to the biofilm mode of
growth, it undergoes a phenotypic shift in behavior in which large
suites of genes are differentially regulated.
A biofilm may also be considered a hydrogel, which is a complex
polymer containing many times its dry weight in water. Biofilms are
not just bacterial slime layers but biological systems; the bacteria
organize themselves into a coordinated functional community. Biofilms
can attach to a surface such as a tooth, rock, or surface which may
include a single species or of a diverse group of microorganisms. The
biofilm bacteria are able to share nutrients and are sheltered from
harmful factors in the environment, such as desiccation, antibiotics,
and a host body's immune system. A biofilm usually begins to form when
a free-swimming bacterium attaches to a surface.[page needed]
4.1 Extracellular matrix
5.1 Dental plaque
6 Taxonomic diversity
7 Infectious diseases
7.1 Pseudomonas aeruginosa
7.2 Streptococcus pneumoniae
8 Uses and impact
8.1 In medicine
8.2 In industry
8.2.1 Food industry
8.3 In aquaculture
9 Eukaryotic biofilms
10 See also
12 External links
13 Further reading
14 External links
An iridescent biofilm on the surface of a fish tank.
The formation of a biofilm begins with the attachment of free-floating
microorganisms to a surface. It is thought that the first
colonist bacteria of a biofilm adhere to the surface initially through
weak, reversible adhesion via van der Waals forces and hydrophobic
effects. If the colonists are not immediately separated from
the surface, they can anchor themselves more permanently using cell
adhesion structures such as pili.
Hydrophobicity can also affect the ability of bacteria to form
Bacteria with increased hydrophobicity have reduced
repulsion between the extracellular matrix and the bacterium. Some
bacteria species are not able to attach to a surface on their own
successfully due to their limited motility but are instead able to
anchor themselves to the matrix or directly to other, earlier bacteria
Non-motile bacteria cannot recognize surfaces or aggregate
together as easily as motile bacteria.
During surface colonization bacteria cells are able to communicate
using quorum sensing (QS) products such as N-acyl homoserine lactone
(AHL). Once colonization has begun, the biofilm grows through a
combination of cell division and recruitment.
typically enclose bacterial biofilms. In addition to the
polysaccharides, these matrices may also contain material from the
surrounding environment, including but not limited to minerals, soil
particles, and blood components, such as erythrocytes and fibrin.
The final stage of biofilm formation is known as dispersion, and is
the stage in which the biofilm is established and may only change in
shape and size.
The development of a biofilm may allow for an aggregate cell colony
(or colonies) to be increasingly resistant to antibiotics. Cell-cell
communication or quorum sensing has been shown to be involved in the
formation of biofilm in several bacterial species.
Five stages of biofilm development: (1) Initial attachment, (2)
Irreversible attachment, (3) Maturation I, (4) Maturation II, and (5)
Dispersion. Each stage of development in the diagram is paired with a
photomicrograph of a developing P. aeruginosa biofilm. All
photomicrographs are shown to the same scale.
Biofilms are the product of a microbial developmental process. The
process is summarized by five major stages of biofilm development (see
illustration on the right):
Dispersal of cells from the biofilm colony is an essential stage of
the biofilm life cycle. Dispersal enables biofilms to spread and
colonize new surfaces. Enzymes that degrade the biofilm extracellular
matrix, such as dispersin B and deoxyribonuclease, may play a role in
Biofilm matrix degrading enzymes may be
useful as anti-biofilm agents. Recent evidence has shown that
a fatty acid messenger, cis-2-decenoic acid, is capable of inducing
dispersion and inhibiting growth of biofilm colonies. Secreted by
Pseudomonas aeruginosa, this compound induces cyclo heteromorphic
cells in several species of bacteria and the yeast Candida
Nitric oxide has also been shown to trigger the
dispersal of biofilms of several bacteria species at sub-toxic
Nitric oxide has the potential for the treatment of
patients that suffer from chronic infections caused by biofilms.
It is generally assumed that cells dispersed from biofilms immediately
go into the planktonic growth phase. However, recent studies have
shown that the physiology of dispersed cells from Pseudomonas
aeruginosa biofilms is highly different from those of planktonic and
biofilm cells. Hence, the dispersal process is a unique stage
during the transition from biofilm to planktonic lifestyle in
bacteria. Dispersed cells are found to be highly virulent against
macrophages and Caenorhabditis elegans, but highly sensitive towards
iron stress, as compared with planktonic cells.
Biofilms are usually found on solid substrates submerged in or exposed
to an aqueous solution, although they can form as floating mats on
liquid surfaces and also on the surface of leaves, particularly in
high humidity climates. Given sufficient resources for growth, a
biofilm will quickly grow to be macroscopic (visible to the naked
eye). Biofilms can contain many different types of microorganism, e.g.
bacteria, archaea, protozoa, fungi and algae; each group performs
specialized metabolic functions. However, some organisms will form
single-species films under certain conditions. The social structure
(cooperation/competition) within a biofilm depends highly on the
different species present.
The EPS matrix consists of exopolysaccharides, proteins and nucleic
acids. A large proportion of the EPS is more or less
strongly hydrated, however, hydrophobic EPS also occur; one example is
cellulose which is produced by a range of microorganisms. This
matrix encases the cells within it and facilitates communication among
them through biochemical signals as well as gene exchange. The EPS
matrix also traps extracellular enzymes and keeps them in close
proximity to the cells. Thus, the matrix represents an external
digestion system and allows for stable synergistic microconsortia of
different species (Wingender and Flemming, Nat. Rev. Microbiol. 8,
623-633). Some biofilms have been found to contain water channels that
help distribute nutrients and signalling molecules. This matrix is
strong enough that under certain conditions, biofilms can become
Bacteria living in a biofilm usually have significantly different
properties from free-floating bacteria of the same species, as the
dense and protected environment of the film allows them to cooperate
and interact in various ways. One benefit of this environment is
increased resistance to detergents and antibiotics, as the dense
extracellular matrix and the outer layer of cells protect the interior
of the community. In some cases antibiotic resistance can be increased
Lateral gene transfer
Lateral gene transfer is often facilitated within
bacterial and archaeal biofilms and leads to a more stable biofilm
DNA is a major structural component of
many different microbial biofilms. Enzymatic degradation of
DNA can weaken the biofilm structure and release
microbial cells from the surface.
However, biofilms are not always less susceptible to antibiotics. For
instance, the biofilm form of
Pseudomonas aeruginosa has no greater
resistance to antimicrobials than do stationary-phase planktonic
cells, although when the biofilm is compared to logarithmic-phase
planktonic cells, the biofilm does have greater resistance to
antimicrobials. This resistance to antibiotics in both
stationary-phase cells and biofilms may be due to the presence of
Mats of bacterial biofilm color the hot springs in Yellowstone
National Park. The longest raised mat area is about half a meter long.
Thermophilic bacteria in the outflow of Mickey Hot Springs, Oregon,
approximately 20 mm thick.
Biofilms are ubiquitous in organic life. Nearly every species of
microorganism have mechanisms by which they can adhere to surfaces and
to each other. Biofilms will form on virtually every non-shedding
surface in non-sterile aqueous or humid environments. Biofilms can
grow in the most extreme environments: from, for example, the
extremely hot, briny waters of hot springs ranging from very acidic to
very alkaline, to frozen glaciers.
Biofilms can be found on rocks and pebbles at the bottoms of most
streams or rivers and often form on the surfaces of stagnant pools of
water. Biofilms are important components of food chains in rivers and
streams and are grazed by the aquatic invertebrates upon which many
fish feed. Biofilms are found on the surface of and inside plants.
They can either contribute to crop disease or, as in the case of
nitrogen-fixing Rhizobium on roots, exist symbiotically with the
plant. Examples of crop diseases related to biofilms include
Pierce's Disease of grapes, and Bacterial Spot of
plants such as peppers and tomatoes.
Recent studies in 2003 discovered that the immune system supports
bio-film development in the large intestine. This was supported mainly
with the fact that the two most abundantly produced molecules by the
immune system also support bio-film production and are associated with
the bio-films developed in the gut. This is especially important
because the appendix holds a mass amount of these bacterial
bio-films. This discovery helps to distinguish the possible
function of the appendix and the idea that the appendix can help
reinoculate the gut with good gut flora.
In the human environment, biofilms can grow in showers very easily
since they provide a moist and warm environment for the biofilm to
thrive. Biofilms can form inside water and sewage pipes and cause
clogging and corrosion. Biofilms on floors and counters can make
sanitation difficult in food preparation areas.
Biofilm in soil can
cause bioclogging. Biofilms in cooling- or heating-water systems are
known to reduce heat transfer. Biofilms in marine engineering
systems, such as pipelines of the offshore oil and gas industry,
can lead to substantial corrosion problems.
Corrosion is mainly due to
abiotic factors; however, at least 20% of corrosion is caused by
microorganisms that are attached to the metal subsurface (i.e.,
microbially influenced corrosion).
Bacterial adhesion to boat hulls serves as the foundation for
biofouling of seagoing vessels. Once a film of bacteria forms, it is
easier for other marine organisms such as barnacles to attach. Such
fouling can reduce maximum vessel speed by up to 20%, prolonging
voyages and consuming fuel. Time in dry dock for refitting and
repainting reduces the productivity of shipping assets, and the useful
life of ships is also reduced due to corrosion and mechanical removal
(scraping) of marine organisms from ships' hulls.
Stromatolites are layered accretionary structures formed in shallow
water by the trapping, binding and cementation of sedimentary grains
by microbial biofilms, especially of cyanobacteria. Stromatolites
include some of the most ancient records of life on Earth, and are
still forming today.
Within the human body, biofilms are present on the teeth as dental
plaque, where they may cause tooth decay and gum disease. These
biofilms can either be in an uncalcified state that can be removed by
dental instruments, or a calcified state which is more difficult to
remove. Removal techniques can also include antimicrobials.
Dental plaque is an oral biofilm that adheres to the teeth and
consists of many species of both bacteria and fungi (such as
Streptococcus mutans and Candida albicans), embedded in salivary
polymers and microbial extracellular products. The accumulation of
microorganisms subjects the teeth and gingival tissues to high
concentrations of bacterial metabolites which results in dental
Biofilm on the surface of teeth is frequently subject to
oxidative stress and acid stress. Dietary carbohydrates can
cause a dramatic decrease in pH in oral biofilms to values of 4 and
below (acid stress). A pH of 4 at body temperature of 37 °C
causes depurination of DNA, leaving apurinic (AP) sites in DNA,
especially loss of guanine.
The dental plaque biofilm can result in the disease dental caries if
it is allowed to develop over time. An ecologic shift away from
balanced populations within the dental biofilm is driven by certain
(cariogenic) microbiological populations beginning to dominate when
the environment favours them. The shift to an acidogenic, aciduric,
and cariogenic microbiological population develops and is maintained
by frequent consumption of fermentable dietary carbohydrate. The
resulting activity shift in the biofilm (and resulting acid production
within the biofilm, at the tooth surface) is associated with an
imbalance between demineralization and remineralisation leading to net
mineral loss within dental hard tissues (enamel and then dentin), the
sign and symptom being a carious lesion. By preventing the dental
plaque biofilm from maturing or by returning it back to a
non-cariogenic state, dental caries can be prevented and arrested.
This can be achieved though the behavioural step of reducing the
supply of fermentable carbohydrates (i.e. sugar intake) and frequent
removal of the biofilm (i.e. toothbrushing).
A peptide pheromone quorum sensing signaling system in S. mutans
includes the Competence Stimulating Peptide (CSP) that controls
genetic competence. Genetic competence is the ability of a
cell to take up
DNA released by another cell. Competence can lead to
genetic transformation, a form of sexual interaction, favored under
conditions of high cell density and/or stress where there is maximal
opportunity for interaction between the competent cell and the DNA
released from nearby donor cells. This system is optimally expressed
when S. mutans cells reside in an actively growing biofilm. Biofilm
grown S. mutans cells are genetically transformed at a rate 10- to
600-fold higher than S. mutans growing as free-floating planktonic
cells suspended in liquid.
When the biofilm, containing S. mutans and related oral streptococci,
is subjected to acid stress, the competence regulon is induced,
leading to resistance to being killed by acid. As pointed out by
Michod et al., transformation in bacterial pathogens likely provides
for effective and efficient recombinational repair of
It appears that S. mutans can survive the frequent acid stress in oral
biofilms, in part, through the recombinational repair provided by
competence and transformation.
Many different bacteria form biofilms, including gram-positive (e.g.
Bacillus spp, Listeria monocytogenes,
Staphylococcus spp, and lactic
acid bacteria, including
Lactobacillus plantarum and Lactococcus
lactis) and gram-negative species (e.g. Escherichia coli, or
Cyanobacteria also form biofilms in
Biofilms are formed by bacteria that colonize plants, e.g. Pseudomonas
putida, Pseudomonas fluorescens, and related pseudomonads which are
common plant-associated bacteria found on leaves, roots, and in the
soil, and the majority of their natural isolates form biofilms.
Several nitrogen-fixing symbionts of legumes such as Rhizobium
Sinorhizobium meliloti form biofilms on legume roots
and other inert surfaces.
Along with bacteria, biofilms are also generated by archaea
and by a range of eukaryotic organisms, including fungi e.g.
Cryptococcus laurentii and microalgae. Among microalgae, one of
the main progenitors of biofilms are diatoms, which colonise both
fresh and marine environments worldwide.
For other species in disease-associated biofilms and biofilms arising
from eukaryotes see below.
Biofilms have been found to be involved in a wide variety of microbial
infections in the body, by one estimate 80% of all infections.
Infectious processes in which biofilms have been implicated include
common problems such as bacterial vaginosis, urinary tract infections,
catheter infections, middle-ear infections, formation of dental
plaque, gingivitis, coating contact lenses, and less common
but more lethal processes such as endocarditis, infections in cystic
fibrosis, and infections of permanent indwelling devices such as joint
prostheses, heart valves, and intervertebral disc. More
recently it has been noted that bacterial biofilms may impair
cutaneous wound healing and reduce topical antibacterial efficiency in
healing or treating infected skin wounds. Early detection of
biofilms in wounds is crucial to successful chronic wound management.
Although many techniques have developed to identify planktonic
bacteria in viable wounds, few have been able to quickly and
accurately identify bacterial biofilms. Future studies are needed to
find means of identifying and monitoring biofilm colonization at the
bedside to permit timely initiation of treatment.
It has recently been shown that biofilms are present on the removed
tissue of 80% of patients undergoing surgery for chronic sinusitis.
The patients with biofilms were shown to have been denuded of cilia
and goblet cells, unlike the controls without biofilms who had normal
cilia and goblet cell morphology. Biofilms were also found on
samples from two of 10 healthy controls mentioned. The species of
bacteria from intraoperative cultures did not correspond to the
bacteria species in the biofilm on the respective patient's tissue. In
other words, the cultures were negative though the bacteria were
present. New staining techniques are being developed to
differentiate bacterial cells growing in living animals, e.g. from
tissues with allergy-inflammations.
Research has shown that sub-therapeutic levels of β-lactam
antibiotics induce biofilm formation in
Staphylococcus aureus. This
sub-therapeutic level of antibiotic may result from the use of
antibiotics as growth promoters in agriculture, or during the normal
course of antibiotic therapy. The biofilm formation induced by
low-level methicillin was inhibited by DNase, suggesting that the
sub-therapeutic levels of antibiotic also induce extracellular DNA
release. Moreover, from an evolutionary point of view, the
creation of the tragedy of the commons in pathogenic microbes may
provide advanced therapeutic ways for chronic infections caused by
biofilms via genetically engineered invasive cheaters who can invade
wild-types ‘cooperators’ of pathogenic bacteria until cooperator
populations go to extinction or overall population ‘cooperators and
cheaters ’ go to extinction.
P. aeruginosa represents a commonly used biofilm model organism since
it is involved in different types of biofilm-associated
infections. Examples of such infections include chronic wounds,
chronic otitis media, chronic prostatitis and chronic lung infections
in cystic fibrosis (CF) patients. About 80% of CF patients have
chronic lung infection, caused mainly by P. aeruginosa growing in a
non-surface attached biofilms surround by PMN. The infection
remains present despite aggressive antibiotic therapy and is a common
cause of death in CF patients due to constant inflammatory damage to
S. pneumoniae is the main cause of community-acquired pneumonia and
meningitis in children and the elderly, and of septicemia in
HIV-infected persons. When S. pneumonia grows in biofilms, genes are
specifically expressed that respond to oxidative stress and induce
competence. Formation of a biofilm depends on competence
stimulating peptide (CSP). CSP also functions as a quorum-sensing
peptide. It not only induces biofilm formation, but also increases
virulence in pneumonia and meningitis.
It has been proposed that competence development and biofilm formation
is an adaptation of S. pneumoniae to survive the defenses of the
host. In particular, the host’s polymorphonuclear leukocytes
produce an oxidative burst to defend against the invading bacteria,
and this response can kill bacteria by damaging their DNA. Competent
S. pneumoniae in a biofilm have the survival advantage that they can
more easily take up transforming
DNA from nearby cells in the biofilm
to use for recombinational repair of oxidative damages in their DNA.
Competent S. pneumoniae can also secrete an enzyme (murein hydrolase)
that destroys non-competent cells (fratricide) causing
DNA to be
released into the surrounding medium for potential use by the
Uses and impact
Infections associated with the biofilm growth usually are challenging
to eradicate. It is mostly due to the fact that mature biofilms
display tolerance towards antibiotics and the immune response.
Biofilms often form on the inert surfaces of implanted devices such as
catheters, prosthetic cardiac valves and intrauterine devices.
The rapidly expanding worldwide industry for biomedical devices and
tissue engineering related products is already at $180 billion per
year, yet this industry continues to suffer from microbial
colonization. No matter the sophistication, microbial infections can
develop on all medical devices and tissue engineering constructs.
60-70% of nosocomial or hospital acquired infections are associated
with the implantation of a biomedical device. This leads to 2
million cases annually in the U.S., costing the healthcare system over
$5 billion in additional healthcare expenses.
Biofilms can also be harnessed for constructive purposes. For example,
many sewage treatment plants include a secondary treatment stage in
which waste water passes over biofilms grown on filters, which extract
and digest organic compounds. In such biofilms, bacteria are mainly
responsible for removal of organic matter (BOD), while protozoa and
rotifers are mainly responsible for removal of suspended solids (SS),
including pathogens and other microorganisms. Slow sand filters rely
on biofilm development in the same way to filter surface water from
lake, spring or river sources for drinking purposes. What we regard as
clean water is effectively a waste material to these microcellular
organisms. Biofilms can help eliminate petroleum oil from contaminated
oceans or marine systems. The oil is eliminated by the
hydrocarbon-degrading activities of microbial communities, in
particular by a remarkable recently discovered group of specialists,
the so-called hydrocarbonoclastic bacteria (HCB). Biofilms are
used in microbial fuel cells (MFCs) to generate electricity from a
variety of starting materials, including complex organic waste and
renewable biomass. Biofilms are also relevant for the
improvement of metal dissolution in bioleaching industry
Biofilms have become problematic in several food industries due to the
ability to form on plants and during industrial processes.
Bacteria can survive long periods of time in water, animal manure, and
soil, causing biofilm formation on plants or in the processing
equipment. The buildup of biofilms can affect the heat flow across
a surface and increase surface corrosion and frictional resistance of
fluids. These can lead to a loss of energy in a system and overall
loss of products. Along with economic problems, biofilm formation
on food poses a health risk to consumers due to the ability to make
the food more resistant to disinfectants As a result, from 1996 to
Center for Disease Control and Prevention
Center for Disease Control and Prevention estimated 48
million foodborne illnesses per year. Biofilms have been connected
to about 80% of bacterial infections in the United States.
In produce, microorganisms attach to the surfaces and biofilms develop
internally. During the washing process, biofilms resist
sanitization and allow bacteria to spread across the produce. This
problem is also found in ready-to-eat foods, because the foods go
through limited cleaning procedures before consumption Due to the
perishability of dairy products and limitations in cleaning
procedures, resulting in the buildup of bacteria, dairy is susceptible
to biofilm formation and contamination. The bacteria can spoil
the products more readily and contaminated products pose a health risk
to consumers. One bacteria that can be found in various industries and
is a major cause of foodborne disease is Salmonella. Large amounts
of salmonella contamination can be found in the poultry processing
industry as about 50% of salmonella strains can produce biofilms on
Salmonella increases the risk of foodborne
illnesses when the poultry products are not cleaned and cooked
Salmonella is also found in the seafood industry where
biofilms form from seafood borne pathogens on the seafood itself as
well as in water. Shrimp products are commonly affected by
salmonella because of unhygienic processing and handling
techniques The preparation practices of shrimp and other seafood
products can allow for bacteria buildup on the products.
New forms of cleaning procedures are being tested in order to reduce
biofilm formation in these processes which will lead to safer and more
productive food processing industries. These new forms of cleaning
procedures also have a profound effect on the environment, often
releasing toxic gases into the groundwater reservoirs.
A bioflim from the Dead sea
In shellfish and algae farms, biofouling species tend to block nets
and cages and ultimately outcompete the farmed species for space and
food. Bacterial biofilms start the colonization process by
creating microenvironments that more favorable for biofouling species.
In the marine environment, biofilms could reduce the hydrodynamic
efficiency of ships and propellers, lead to pipeline blockage and
sensor malfunction, and increase the weight of appliances deployed in
seawater. Numerous studies have shown that biofilm can be a
reservoir for potentially pathogenic bacteria in freshwater
aquaculture. As mentioned previously, biofilms can be
difficult to eliminate even when antibiotics or chemicals are used in
high doses. The role that biofilm plays as reservoirs of
bacterial fish pathogens regarding has not been explored in detail but
it certainly deserves to be studied.
See also: Phototrophic biofilms
Along with bacteria, biofilms are often initiated and produced by
Eukaryotes. The biofilms produced by eukaryotes is usually occupied by
bacteria and other Eukaryotes alike, however the surface is cultivated
and EPS is secreted initially by the Eukaryote. Both fungi
and microalgae are known to form biofilms in such a way. Biofilms of
fungal origin are important aspects of human infection and fungal
pathagenicity, as the fungal infection is more resistant to
In the environment, fungal biofilms are an area of ongoing research.
One key area of research are fungal biofilms on plants. For example,
in the soil, plant associated fungi including mycorrhiza have been
shown to decompose organic matter, protect plants from bacterial
Biofilms in aquatic environments are often founded by diatoms. The
exact purpose of these biofilms is unknown, however there is evidence
that the EPS produced by diatoms facilitates both cold and salinity
stress. These Eukaryotes interact with a diverse range of
other organisms within a region known as the phycosphere, but
importantly are the bacteria associated with diatoms, as it has been
shown that although diatoms excrete EPS, they only do so when
interacting with certain bacteria species.
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Thickness analysis, organic and mineral proportion of biofilms in
order to decide a treatment strategy
Biofilm Archive of
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