Quorum sensing

In biology, quorum sensing is the ability to

detect and to respond to cell population
density by gene regulation. As one
example, quorum sensing (QS) enables
bacteria to restrict the expression of
specific genes to the high cell densities at
which the resulting phenotypes will be
most beneficial. Many species of bacteria
use quorum sensing to coordinate gene
expression according to the density of
their local population. In similar fashion,
some social insects use quorum sensing
to determine where to nest.

In addition to its function in biological

systems, quorum sensing has several
useful applications for computing and
robotics. In general, quorum sensing can
function as a decision-making process in
any decentralized system in which the
components have: (a) a means of
assessing the number of other
components they interact with and (b) a
standard response once a threshold
number of components is detected.

Bacteria
Some of the best-known examples of
quorum sensing come from studies of
bacteria. Bacteria use quorum sensing to
regulate certain phenotype expressions,
which in turn, coordinate their behaviours.
Some common phenotypes include biofilm
formation, virulence factor expression, and
motility. Certain bacteria are able to use
quorum sensing to regulate
bioluminescence, nitrogen fixation and
sporulation.[1]

Quorum sensing functions based on the

local density of the bacterial population in
the immediate environment.[2] It can occur
within a single bacterial species, as well as
between diverse species. Both Gram-
positive and gram-negative bacteria use
quorum sensing, but there are some major
differences in their mechanisms.[3]

Mechanism

For the bacteria to use quorum sensing

constitutively, they must possess three
characteristics: to secrete a signaling
molecule, an autoinducer, to detect the
change in concentration of signaling
molecules, and to regulate gene
transcription as a response.[1] This
process is highly dependent on the
diffusion mechanism of the signaling
molecules. QS Signaling molecules are
usually secreted at a low level by individual
bacteria. At low cell density, the molecules
may just diffuse away. At high cell density,
the local concentration of signaling
molecules may exceed its threshold level,
and trigger changes in gene
expressions.[3]

Gram-positive Bacteria

Gram-positive bacteria use autoinducing

peptide (AIP) as their autoinducers.[4]

When gram-positive bacteria detect high

concentration of AIP in the environment,
AIP bind to a receptor to activate kinase.
The kinase phosphorylates a transcription
factor, which regulated gene transcription.
This is called a two-component system.

Another possible mechanism is that AIP is

transported into the cytosol, and binds
directly to a transcription factor to initiate
or inhibit transcription.[4]

Gram-negative Bacteria

Gram-negative bacteria produce N-acyl

homoserine lactones (AHL) as their
signaling molecule.[4] Usually AHLs do not
need additional processing, and bind
directly to transcription factors to regulate
gene expression.[3]

Some gram-negative bacteria may use the

two-component system as well.[4]

Quorum sensing of Gram Negative cell

Examples

Aliivibrio fischeri
Quorum sensing was first observed in
Aliivibrio fischeri, a bioluminescent
bacterium that lives as a mutualistic
symbiont in the photophore (or light-
producing organ) of the Hawaiian bobtail
squid.[5] When A. fischeri cells are free-
living (or planktonic), the autoinducer is at
low concentration, and, thus, cells do not
show luminescence. However, when the
population reaches the threshold in the
photophore (about 1011 cells/ml),
transcription of luciferase is induced,
leading to bioluminescence.

Escherichia coli
In the Gram-negative bacterium
Escherichia coli (E. coli), cell division may
be partially regulated by AI-2-mediated
quorum sensing. This species uses AI-2,
which is produced and processed by the
lsr operon. Part of it encodes an ABC
transporter, which imports AI-2 into the
cells during the early stationary (latent)
phase of growth. AI-2 is then
phosphorylated by the LsrK kinase, and
the newly produced phospho-AI-2 can be
either internalized or used to suppress
LsrR, a repressor of the lsr operon (thereby
activating the operon). Transcription of the
lsr operon is also thought to be inhibited
by dihydroxyacetone phosphate (DHAP)
through its competitive binding to LsrR.
Glyceraldehyde 3-phosphate has also been
shown to inhibit the lsr operon through
cAMP-CAPK-mediated inhibition. This
explains why, when grown with glucose, E.
coli will lose the ability to internalize AI-2
(because of catabolite repression). When
grown normally, AI-2 presence is transient.

E. coli and Salmonella enterica do not

produce AHL signals commonly found in
other Gram-negative bacteria. However,
they have a receptor that detects AHLs
from other bacteria and change their gene
expression in accordance with the
presence of other "quorate" populations of
Gram-negative bacteria.[6]

Gram positive bacteria quorum sensing

Salmonella enterica

Salmonella encodes a LuxR homolog, SdiA,

but does not encode an AHL synthase.
SdiA detects AHLs produced by other
species of bacteria including Aeromonas
hydrophila, Hafnia alvei, and Yersinia
enterocolitica.[7] When AHL is detected,
SdiA regulates the rck operon on the
Salmonella virulence plasmid (pefI-srgD-
srgA-srgB-rck-srgC) and a single gene
horizontal acquisition in the chromosome
srgE.[8][9] Salmonella does not detect AHL
when passing through the gastrointestinal
tracts of several animal species,
suggesting that the normal microbiota
does not produce AHLs. However, SdiA
does become activated when Salmonella
transits through turtles colonized with
Aeromonas hydrophila or mice infected
with Yersinia enterocolitica.[10][11]
Therefore, Salmonella appears to use SdiA
to detect the AHL production of other
pathogens rather than the normal gut
flora.

Pseudomonas aeruginosa

The opportunistic pathogen Pseudomonas

aeruginosa uses quorum sensing to
coordinate the formation of biofilm,
swarming motility, exopolysaccharide
production, virulence, and cell
aggregation.[12] These bacteria can grow
within a host without harming it, until they
reach a threshold concentration. Then they
become aggressive, developing to the
point at which their numbers are sufficient
to overcome the host's immune system,
and form a biofilm, leading to disease
within the host as the biofilm is a
protective layer encasing the bacteria
population. Another form of gene
regulation that allows the bacteria to
rapidly adapt to surrounding changes is
through environmental signaling. Recent
studies have discovered that anaerobiosis
can significantly impact the major
regulatory circuit of quorum sensing. This
important link between quorum sensing
and anaerobiosis has a significant impact
on production of virulence factors of this
organism.[13] Garlic and ginseng
experimentally block quorum sensing in
Pseudomonas aeruginosa.[14] It is hoped
that the therapeutic enzymatic
degradation of the signaling molecules will
prevent the formation of such biofilms and
possibly weaken established biofilms.
Disrupting the signalling process in this
way is called quorum sensing inhibition.

Acinetobacter sp.

It has recently been found that

Acinetobacter sp. also show quorum
sensing activity. This bacterium, an
emerging pathogen, produces AHLs.[15]
Acinetobacter sp. shows both quorum
sensing and quorum quenching activity. It
produces AHLs and also, it can degrade
the AHL molecules as well.[15]

Aeromonas sp.

This bacterium was previously considered

a fish pathogen, but it has recently
emerged as a human pathogen.[16]
Aeromonas sp. have been isolated from
various infected sites from patients (bile,
blood, peritoneal fluid, pus, stool and
urine). All isolates produced the two
principal AHLs, N-butanoylhomoserine
lactone (C4-HSL) and N-hexanoyl
homoserine lactone (C6-HSL). It has been
documented that Aeromonas sobria has
produced C6-HSL and two additional AHLs
with N-acyl side chain longer than C6.[17]

Yersinia

The YenR and YenI proteins produced by

the gammaproteobacterium Yersinia
enterocolitica are similar to Aliivibrio
fischeri LuxR and LuxI.[18][19] YenR
activates the expression of a small non-
coding RNA, YenS. YenS inhibits YenI
expression and acylhomoserine lactone
production.[20] YenR/YenI/YenS are
involved in the control of swimming and
swarming motility.[19][20]
Molecules involved

Three-dimensional structures of proteins

involved in quorum sensing were first
published in 2001, when the crystal
structures of three LuxS orthologs were
determined by X-ray crystallography.[21] In
2002, the crystal structure of the receptor
LuxP of Vibrio harveyi with its inducer AI-2
(which is one of the few biomolecules
containing boron) bound to it was also
determined.[22] Many bacterial species,
including E. coli, an enteric bacterium and
model organism for Gram-negative
bacteria, produce AI-2. A comparative
genomic and phylogenetic analysis of 138
genomes of bacteria, archaea, and
eukaryotes found that "the LuxS enzyme
required for AI-2 synthesis is widespread
in bacteria, while the periplasmic binding
protein LuxP is present only in Vibrio
strains," leading to the conclusion that
either "other organisms may use
components different from the AI-2 signal
transduction system of Vibrio strains to
sense the signal of AI-2 or they do not
have such a quorum sensing system at
all."[23] Farnesol is used by the fungus
Candida albicans as a quorum sensing
molecule that inhibits filamentation.[24]
A database of quorum-sensing peptides is
available under the name
Quorumpeps.[25][26]

Certain bacteria can produce enzymes

called lactonases that can target and
inactivate AHLs. Researchers have
developed novel molecules which block
the signalling receptors of bacteria
(Quorum quenching). mBTL is a
compound that has been shown to inhibit
quorum sensing and decrease the amount
of cell death by a significant amount.[27]
Additionally, researchers are also
examining the role of natural compounds
(such as caffeine) as potential quorum
sensing inhibitors.[28] Research in this area
has been promising and could lead to the
development of natural compounds as
effective therapeutics.

Evolution

Sequence analysis

The majority of quorum sensing systems

that fall under the "two-gene" (an
autoinducer synthase coupled with a
receptor molecule) paradigm as defined by
the Vibrio fischeri system occur in the
Gram-negative Proteobacteria. A
comparison between the Proteobacteria
phylogeny as generated by 16S ribosomal
RNA sequences and phylogenies of LuxI-,
LuxR-, or LuxS-homologs shows a notably
high level of global similarity. Overall, the
quorum sensing genes seem to have
diverged along with the Proteobacteria
phylum as a whole. This indicates that
these quorum sensing systems are quite
ancient, and arose very early in the
Proteobacteria lineage.[29][30]

Although examples of horizontal gene

transfer are apparent in LuxI, LuxR, and
LuxS phylogenies, they are relatively rare.
This result is in line with the observation
that quorum sensing genes tend to control
the expression of a wide array of genes
scattered throughout the bacterial
chromosome. A recent acquisition by
horizontal gene transfer would be unlikely
to have integrated itself to this degree.
Given that the majority of autoinducer–
synthase/receptor occurs in tandem in
bacterial genomes, it is also rare that they
switch partners and so pairs tend to co-
evolve.[30]

In quorum sensing genes of

Gammaproteobacteria, which includes
Pseudomonas aeruginosa and Escherichia
coli, the LuxI/LuxR genes form a functional
pair, with LuxI as the auto-inducer
synthase and LuxR as the receptor.
Gamma Proteobacteria are unique in
possessing quorum sensing genes, which,
although functionally similar to the
LuxI/LuxR genes, have a markedly
divergent sequence.[30] This family of
quorum-sensing homologs may have
arisen in the gamma Proteobacteria
ancestor, although the cause of their
extreme sequence divergence yet
maintenance of functional similarity has
yet to be explained. In addition, species
that employ multiple discrete quorum
sensing systems are almost all members
of the gamma Proteobacteria, and
evidence of horizontal transfer of quorum
sensing genes is most evident in this
class.[29][30]

Controversy

As quorum sensing implies a cooperative

behavior, this concept has been
challenged by the evolutionary implication
of cooperative cheaters. This is
circumvented by the concept of diffusion
sensing, which has been an alternative and
complementary model to quorum
sensing.[31] However, both explanations
face the problems of signalling in either
complex (multiple species sharing the
same space) or simple (one single cell
confined to a limited volume)
environments where the spatial
distribution of the cells can be more
important for sensing than the cell
population density. A new model,
efficiency sensing, which takes into
account both problematics, population
density and spatial confinement, has been
proposed as an alternative.[32] One of the
probable reasons for controversy is that
current terminologies (quorum sensing,
diffusion sensing, efficiency sensing) arise
from different models of how natural
selection has acted to shape and maintain
the process. Since each of the various
models does apply under some
circumstances but not others, a sensible
resolution to these controversies could be
to return the terminology of the process to
autoinduction, as originally described by
Hastings and coworkers, as this term does
not imply understanding of the function of
the process.

Interaction of quorum-sensing
molecules with mammalian cells
and its medical applications

Next to the potential antimicrobial

functionality, quorum-sensing derived
molecules, especially the peptides, are
being investigated for their use in other
therapeutic domains as well, including
immunology, central nervous system
disorders and oncology. Quorum-sensing
peptides have been demonstrated to
interact with cancer cells, as well as to
permeate the blood-brain barrier
permeation reaching the brain
parenchyma[33][34][35][36]

Archaea
Examples

Methanosaeta harundinacea 6Ac

Methanosaeta harundinacea 6Ac, a

methanogenic archaeon, produces
carboxylated acyl homoserine lactone
compounds that facilitate the transition
from growth as short cells to growth as
filaments.[37]

Plants
Quorum sensing could be described when
it was known that bacteria possess the
ability to communicate. In the last few
years, interactions between bacteria and
eukaryotic hosts, such as plants, have
been shown. These interactions are
facilitated by quorum-sensing molecules
and play a major role in maintaining the
pathogenicity of bacteria towards other
hosts, such as humans. This mechanism
can be understood by looking at the
effects of N-Acyl homoserine lactone
(AHL), one of the quorum sensing-
signalling molecules in gram-negative
bacteria, on plants. The model organism
used is Arabidopsis thaliana.[38]

The role of AHLs having long carbon-

chains (C12, C14), which have an unknown
receptor mechanism, is less well
understood than AHLs having short
carbon-chains (C4, C6, C8), which are
perceived by the G protein-coupled
receptor. A phenomenon called "AHL
priming", which is a dependent signalling
pathway, enhanced our knowledge of long-
chain AHLs. The role of quorum-sensing
molecules was better explained according
to three categories: host physiology–
based impact of quorum sensing
molecules; ecological effects; and cellular
signaling. Calcium signalling and
calmodulin have a large role in short-chain
AHLs response in Arabidopsis. Research
was also conducted on barley and crop
yam bean that reveals the AHLs
determining the detoxification enzymes
called GST were found less in yam
bean.[39]
Quorum sensing-based regulatory
systems are necessary to plant-disease-
causing bacteria. Looking towards
developing new strategies based on plant-
associated microbiomes, the aim of
further study is to improve the quantity
and quality of the food supply. Further
research into this inter-kingdom
communication also enhances the
possibility of learning about quorum
sensing in humans.[40]

Quorum quenching
Quorum quenching is the process of
preventing quorum sensing by disrupting
signalling.[41] This is achieved by
inactivating signalling enzymes, by
introducing molecules that mimic
signalling molecules and block their
receptors, or by degrading signalling
molecules themselves.[41][42][43]

Closantel and triclosan are known

inhibitors of quorum sensing enzymes.[44]
Closantel induces aggregation of the
histamine kinase sensor in two-
component signalling. The latter disrupts
the synthesis of a class of signalling
molecules known as N-acyl homoserine
lactones (AHLs) by blocking the enoyl-acyl
carrier protein (ACP) reductase.[44][45]
Two groups of well-known mimicking
molecules include halogenated furanones,
which mimic AHL molecules, and
synthetic Al peptides (AIPs), which mimic
naturally occurring AIPs. These groups
inhibit receptors from binding substrate or
decrease the concentration of receptors in
the cell.[44] Furanones have also been
found to act on AHL-dependant
transcriptional activity, whereby the half
life of the autoinducer-binding LuxR
protein is significantly shortened.[46]

Recently, a well-studied quorum quenching

bacterial strain (KM1S) was isolated and
its AHL degradation kinetic was studied
using rapid resolution liquid
chromatography (RRLC).[47] RRLC
efficiently separates components of a
mixture to a high degree of sensitivity,
based on their affinities for different liquid
phases.[48] It was found that the genome
of this strain encoded an inactivation
enzyme with distinct motifs targeting the
degradation of AHLs.[47]

Applications of quorum quenching that

have been exploited by humans include
the use of AHL-degrading bacteria in
aquacultures to limit the spread of
diseases in aquatic populations of fish,
mollusks and crustaceans.[49] This
technique has also been translated to
agriculture, to restrict the spread of
pathogenic bacteria that use quorum
sensing in plants.[49][50] Anti-biofouling is
another process that exploits quorum
quenching bacteria to mediate the
dissociation of unwanted biofilms
aggregating on wet surfaces, such as
medical devices, transportation
infrastructure and water systems.[49][51]

Social insects
Social insect colonies are an excellent
example of a decentralized system,
because no individual is in charge of
directing or making decisions for the
colony. Several groups of social insects
have been shown to use quorum sensing
in a process that resembles collective
decision-making.

Examples

Ants

Colonies of the ant Temnothorax

albipennis nest in small crevices between
rocks. When the rocks shift and the nest is
broken up, these ants must quickly choose
a new nest to move into. During the first
phase of the decision-making process, a
small portion of the workers leave the
destroyed nest and search for new
crevices. When one of these scout ants
finds a potential nest, she assesses the
quality of the crevice based on a variety of
factors including the size of the interior,
the number of openings (based on light
level), and the presence or absence of
dead ants.[52][53] The worker then returns
to the destroyed nest, where she waits for
a short period before recruiting other
workers to follow her to the nest that she
has found, using a process called tandem
running. The waiting period is inversely
related to the quality of the site; for
instance, a worker that has found a poor
site will wait longer than a worker that
encountered a good site.[54] As the new
recruits visit the potential nest site and
make their own assessment of its quality,
the number of ants visiting the crevice
increases. During this stage, ants may be
visiting many different potential nests.
However, because of the differences in the
waiting period, the number of ants in the
best nest will tend to increase at the
greatest rate. Eventually, the ants in this
nest will sense that the rate at which they
encounter other ants has exceeded a
particular threshold, indicating that the
quorum number has been reached.[55]
Once the ants sense a quorum, they return
to the destroyed nest and begin rapidly
carrying the brood, queen, and fellow
workers to the new nest. Scouts that are
still tandem-running to other potential
sites are also recruited to the new nest,
and the entire colony moves. Thus,
although no single worker may have
visited and compared all of the available
options, quorum sensing enables the
colony as a whole to quickly make good
decisions about where to move.

Honey bees

Honey bees (Apis mellifera) also use

quorum sensing to make decisions about
new nest sites. Large colonies reproduce
through a process called swarming, in
which the queen leaves the hive with a
portion of the workers to form a new nest
elsewhere. After leaving the nest, the
workers form a swarm that hangs from a
branch or overhanging structure. This
swarm persists during the decision-
making phase until a new nest site is
chosen.

The quorum sensing process in honey

bees is similar to the method used by
Temnothorax ants in several ways. A small
portion of the workers leave the swarm to
search out new nest sites, and each
worker assesses the quality of the cavity it
finds. The worker then returns to the
swarm and recruits other workers to her
cavity using the honey bee waggle dance.
However, instead of using a time delay, the
number of dance repetitions the worker
performs is dependent on the quality of
the site. Workers that found poor nests
stop dancing sooner, and can therefore be
recruited to the better sites. Once the
visitors to a new site sense that a quorum
number (usually 10–20 bees) has been
reached, they return to the swarm and
begin using a new recruitment method
called piping. This vibration signal causes
the swarm to take off and fly to the new
nest location. In an experimental test, this
decision-making process enabled honey
bee swarms to choose the best nest site in
four out of five trials.[56][57]

Computing and robotics

Quorum sensing can be a useful tool for
improving the function of self-organizing
networks such as the SECOAS (Self-
Organizing Collegiate Sensor)
environmental monitoring system. In this
system, individual nodes sense that there
is a population of other nodes with similar
data to report. The population then
nominates just one node to report the
data, resulting in power savings.[58] Ad-hoc
wireless networks can also benefit from
quorum sensing, by allowing the system to
detect and respond to network
conditions.[59]

Quorum sensing can also be used to

coordinate the behavior of autonomous
robot swarms. Using a process similar to
that used by Temnothorax ants, robots can
make rapid group decisions without the
direction of a controller.[60]

See also
Cell signaling
Collective behavior
 Interspecies quorum sensing
Microbial intelligence
Pheromone
Signal Transduction
Swarm intelligence

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Further reading
Dedicated issue of Philosophical
Transactions B on quorum sensing
(2007). Some articles are freely
available.

External links
The Quorum Sensing Website
Cell-to-Cell Communication in Bacteria
 The SECOAS project—Development of a
Self-Organising, Wireless Sensor
Network for Environmental Monitoring
Measurement of Space: From Ants to
Robots
Instant insight into quorum sensing
from the Royal Society of Chemistry
Bonnie Bassler: Discovering bacteria's
amazing communication system
Bonnie Bassler's seminar: "Cell-Cell
Communication in Bacteria"

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