Bacteria: Bacteria Bacteria (/bæk
Bacteria: Bacteria Bacteria (/bæk
Etymology
The word bacteria is the plural of the New Latin bacterium, which is the latinisation of the Greek
βακτήριον (bakterion),[3] the diminutive of βακτηρία (bakteria), meaning "staff, cane",[4] because the first
ones to be discovered were rod-shaped.[5] [6]
Habitat
Bacteria are ubiquitous, living in every possible habitat on the planet including soil, underwater, deep in
Earth's crust and even such extreme environments as acidic hot springs and radioactive waste.[20][21] There
are approximately 2×1030 bacteria on Earth,[22] forming a biomass that is only exceeded by plants.[23]
They are abundant in lakes and oceans, in arctic ice, and geothermal springs[24] where they provide the
nutrients needed to sustain life by converting dissolved compounds, such as hydrogen sulphide and
methane, to energy.[25] They live on and in plants and animals. Most do not cause diseases, are beneficial
to their environments, and are essential for life.[26] The soil is a rich source of bacteria and a few grams
contain around a thousand million of them. They are all essential to soil ecology, breaking down toxic
waste and recycling nutrients. They are even found in the atmosphere and one cubic metre of air holds
around one hundred million bacterial cells. The oceans and seas harbour around 3 x 1026 bacteria which
provide up to 50% of the oxygen humans breathe.[27] Only around 2% of bacterial species have been fully
studied.[28]
Extremophile Bacteria
Habitat Species Reference
Saline, 47% salt (Dead Sea, Great Salt Lake) several species ,[28][29]
High pressure (Mariana Trench – 1200 atm) Moritella, Shewanella and others [29]
Morphology
Bacteria display a wide diversity of shapes
and sizes. Bacterial cells are about one-tenth
the size of eukaryotic cells and are typically
0.5–5.0 micrometres in length. However, a
few species are visible to the unaided eye—
for example, Thiomargarita namibiensis is up
to half a millimetre long[30] and Epulopiscium
fishelsoni reaches 0.7 mm.[31] Among the
smallest bacteria are members of the genus
Mycoplasma, which measure only
0.3 micrometres, as small as the largest
viruses.[32] Some bacteria may be even
smaller, but these ultramicrobacteria are not
well-studied.[33]
Many bacterial species exist simply as single cells; others associate in characteristic patterns: Neisseria
forms diploids (pairs), streptococci form chains, and stahphylococci group together in "bunch of grapes"
clusters. Bacteria can also group to form larger multicellular structures, such as the elongated filaments of
Actinobacteria species, the aggregates of Myxobacteria species, and the complex hyphae of Streptomyces
species.[39] These multicellular structures are often only seen in certain conditions. For example, when
starved of amino acids, myxobacteria detect surrounding cells in a process known as quorum sensing,
migrate towards each other, and aggregate to form fruiting bodies
up to 500 micrometres long and containing approximately 100,000
bacterial cells.[40] In these fruiting bodies, the bacteria perform
separate tasks; for example, about one in ten cells migrate to the
top of a fruiting body and differentiate into a specialised dormant
state called a myxospore, which is more resistant to drying and
other adverse environmental conditions.[41]
Cellular structure
Intracellular structures
Bacteria do not have a membrane-bound nucleus, and their genetic material is typically a single circular
bacterial chromosome of DNA located in the cytoplasm in an irregularly shaped body called the
nucleoid.[59] The nucleoid contains the chromosome with its associated proteins and RNA. Like all other
organisms, bacteria contain ribosomes for the production of
proteins, but the structure of the bacterial ribosome is different
from that of eukaryotes and archaea.[60]
There are broadly speaking two different types of cell wall in bacteria, that classify bacteria into Gram-
positive bacteria and Gram-negative bacteria. The names originate from the reaction of cells to the Gram
stain, a long-standing test for the classification of bacterial species.[68]
Gram-positive bacteria possess a thick cell wall containing many layers of peptidoglycan and teichoic acids.
In contrast, Gram-negative bacteria have a relatively thin cell wall consisting of a few layers of
peptidoglycan surrounded by a second lipid membrane containing lipopolysaccharides and lipoproteins.
Most bacteria have the Gram-negative cell wall, and only members of the Firmicutes group and
actinobacteria (previously known as the low G+C and high G+C Gram-positive bacteria, respectively)
have the alternative Gram-positive arrangement.[69] These differences in structure can produce differences
in antibiotic susceptibility; for instance, vancomycin can kill only Gram-positive bacteria and is ineffective
against Gram-negative pathogens, such as Haemophilus influenzae or Pseudomonas aeruginosa.[70] Some
bacteria have cell wall structures that are neither classically Gram-positive or Gram-negative. This includes
clinically important bacteria such as mycobacteria which have a thick peptidoglycan cell wall like a Gram-
positive bacterium, but also a second outer layer of lipids.[71]
In many bacteria, an S-layer of rigidly arrayed protein molecules covers the outside of the cell.[72] This
layer provides chemical and physical protection for the cell surface and can act as a macromolecular
diffusion barrier. S-layers have diverse functions and are known to act as virulence factors in
Campylobacter species and contain surface enzymes in Bacillus stearothermophilus.[73][74]
Flagella are rigid protein structures, about 20 nanometres in diameter and up to 20 micrometres in length,
that are used for motility. Flagella are driven by the energy released by the transfer of ions down an
electrochemical gradient across the cell membrane.[75]
Fimbriae (sometimes called "attachment pili") are fine filaments of protein, usually 2–10 nanometres in
diameter and up to several micrometres in length. They are distributed over the surface of the cell, and
resemble fine hairs when seen under the electron microscope.[76] Fimbriae are believed to be involved in
attachment to solid surfaces or to other cells, and are essential for the virulence of some bacterial
pathogens.[77] Pili (sing. pilus) are cellular appendages, slightly larger than fimbriae, that can transfer
genetic material between bacterial cells in a process called
conjugation where they are called conjugation pili or sex pili (see
bacterial genetics, below).[78] They can also generate movement
where they are called type IV pili.[79]
The assembly of these extracellular structures is dependent on bacterial secretion systems. These transfer
proteins from the cytoplasm into the periplasm or into the environment around the cell. Many types of
secretion systems are known and these structures are often essential for the virulence of pathogens, so are
intensively studied.[82]
Endospores
Metabolism
Bacteria exhibit an extremely wide variety of metabolic types.[94] The distribution of metabolic traits within
a group of bacteria has traditionally been used to define their taxonomy, but these traits often do not
correspond with modern genetic classifications.[95] Bacterial metabolism is classified into nutritional groups
on the basis of three major criteria: the source of energy, the electron donors used, and the source of carbon
used for growth.[96]
Bacteria either derive energy from light using photosynthesis (called phototrophy), or by breaking down
chemical compounds using oxidation (called chemotrophy).[97] Chemotrophs use chemical compounds as a
source of energy by transferring electrons from a given electron donor to a terminal electron acceptor in a
redox reaction. This reaction releases energy that can be used to drive metabolism. Chemotrophs are further
divided by the types of compounds they use to transfer electrons. Bacteria that use inorganic compounds
such as hydrogen, carbon monoxide, or ammonia as sources of electrons are called lithotrophs, while those
that use organic compounds are called organotrophs.[97] The compounds used to receive electrons are also
used to classify bacteria: aerobic organisms use oxygen as the terminal electron acceptor, while anaerobic
organisms use other compounds such as nitrate, sulfate, or carbon dioxide.[97]
Many bacteria get their carbon from other organic carbon, called heterotrophy. Others such as
cyanobacteria and some purple bacteria are autotrophic, meaning that they obtain cellular carbon by fixing
carbon dioxide.[98] In unusual circumstances, the gas methane can be used by methanotrophic bacteria as
both a source of electrons and a substrate for carbon anabolism.[99]
In many ways, bacterial metabolism provides traits that are useful for ecological stability and for human
society. One example is that some bacteria called diazotrophs have the ability to fix nitrogen gas using the
enzyme nitrogenase.[100] This environmentally important trait can be found in bacteria of most metabolic
types listed above.[101] This leads to the ecologically important processes of denitrification, sulfate
reduction, and acetogenesis, respectively.[102] Bacterial metabolic processes are also important in biological
responses to pollution; for example, sulfate-reducing bacteria are largely responsible for the production of
the highly toxic forms of mercury (methyl- and dimethylmercury) in the environment.[103] Non-respiratory
anaerobes use fermentation to generate energy and reducing power, secreting metabolic by-products (such
as ethanol in brewing) as waste. Facultative anaerobes can switch between fermentation and different
terminal electron acceptors depending on the environmental conditions in which they find themselves.[104]
In the laboratory, bacteria are usually grown using solid or liquid media.[109] Solid growth media, such as
agar plates, are used to isolate pure cultures of a bacterial strain. However, liquid growth media are used
when the measurement of growth or large volumes of cells are required. Growth in stirred liquid media
occurs as an even cell suspension, making the
cultures easy to divide and transfer, although
isolating single bacteria from liquid media is
difficult. The use of selective media (media
with specific nutrients added or deficient, or
with antibiotics added) can help identify
A colony of
specific organisms.[110]
Escherichia
Most laboratory techniques for growing
coli[108]
bacteria use high levels of nutrients to
produce large amounts of cells cheaply and
quickly.[109] However, in natural
environments, nutrients are limited, meaning that bacteria cannot
Many bacteria reproduce through
continue to reproduce indefinitely. This nutrient limitation has led
binary fission, which is compared to
the evolution of different growth strategies (see r/K selection mitosis and meiosis in this image.
theory). Some organisms can grow extremely rapidly when
nutrients become available, such as the formation of algal (and
cyanobacterial) blooms that often occur in lakes during the
summer.[111] Other organisms have adaptations to harsh
environments, such as the production of multiple antibiotics by
streptomyces that inhibit the growth of competing
microorganisms.[112] In nature, many organisms live in
communities (e.g., biofilms) that may allow for increased supply of
nutrients and protection from environmental stresses.[46] These
relationships can be essential for growth of a particular organism or
group of organisms (syntrophy).[113]
Genetics
Most bacteria have a single circular chromosome that can range in size from only 160,000 base pairs in the
endosymbiotic bacteria Carsonella ruddii,[119] to 12,200,000 base pairs (12.2 Mbp) in the soil-dwelling
bacteria Sorangium cellulosum.[120] There are many exceptions to this, for example some Streptomyces
and Borrelia species contain a single linear chromosome,[121][122] while some Vibrio species contain more
than one chromosome.[123] Bacteria can also contain plasmids,
small extra-chromosomal molecules of DNA that may contain
genes for various useful functions such as antibiotic resistance,
metabolic capabilities, or various virulence factors.[124]
Behaviour
Movement
Many bacteria are motile (able to move themselves) and do so using a variety of mechanisms. The best
studied of these are flagella, long filaments that are turned by a motor at the base to generate propeller-like
movement.[138] The bacterial flagellum is made of about 20 proteins, with approximately another 30
proteins required for its regulation and assembly.[138] The
flagellum is a rotating structure driven by a reversible motor at the
base that uses the electrochemical gradient across the membrane
for power.[139]
Communication
A few bacteria have chemical systems that generate light. This bioluminescence often occurs in bacteria
that live in association with fish, and the light probably serves to attract fish or other large animals.[148]
Bacteria often function as multicellular aggregates known as biofilms, exchanging a variety of molecular
signals for inter-cell communication, and engaging in coordinated multicellular behaviour.[149][150]
The communal benefits of multicellular cooperation include a cellular division of labour, accessing
resources that cannot effectively be used by single cells, collectively defending against antagonists, and
optimising population survival by differentiating into distinct cell types.[149] For example, bacteria in
biofilms can have more than 500 times increased resistance to antibacterial agents than individual
"planktonic" bacteria of the same species.[150]
One type of inter-cellular communication by a molecular signal is called quorum sensing, which serves the
purpose of determining whether there is a local population density that is sufficiently high that it is
productive to invest in processes that are only successful if large numbers of similar organisms behave
similarly, as in excreting digestive enzymes or emitting light[151][152]
Quorum sensing allows bacteria to coordinate gene expression, and enables them to produce, release and
detect autoinducers or pheromones which accumulate with the growth in cell population.[153]
The identification of bacteria in the laboratory is particularly relevant in medicine, where the correct
treatment is determined by the bacterial species causing an infection. Consequently, the need to identify
human pathogens was a major impetus for the development of techniques to identify bacteria.[166]
The Gram stain, developed in 1884 by Hans Christian Gram, characterises bacteria based on the structural
characteristics of their cell walls.[167][68] The thick layers of peptidoglycan in the "Gram-positive" cell wall
stain purple, while the thin "Gram-negative" cell wall appears pink.[167] By combining morphology and
Gram-staining, most bacteria can be classified as belonging to one of four groups (Gram-positive cocci,
Gram-positive bacilli, Gram-negative cocci and Gram-negative bacilli). Some organisms are best identified
by stains other than the Gram stain, particularly mycobacteria or Nocardia, which show acid-fastness on
Ziehl–Neelsen or similar stains.[168] Other organisms may need to be identified by their growth in special
media, or by other techniques, such as serology.[169]
Culture techniques are designed to promote the growth and identify particular bacteria, while restricting the
growth of the other bacteria in the sample.[170] Often these techniques are designed for specific specimens;
for example, a sputum sample will be treated to identify organisms that cause pneumonia, while stool
specimens are cultured on selective media to identify organisms that cause diarrhea, while preventing
growth of non-pathogenic bacteria. Specimens that are normally sterile, such as blood, urine or spinal fluid,
are cultured under conditions designed to grow all possible organisms.[110][171] Once a pathogenic
organism has been isolated, it can be further characterised by its morphology, growth patterns (such as
aerobic or anaerobic growth), patterns of hemolysis, and staining.[172]
As with bacterial classification, identification of bacteria is increasingly using molecular methods,[173] and
mass spectroscopy.[174] Most bacteria have not been characterised and there are may species that cannot be
grown in the laboratory.[175] Diagnostics using DNA-based tools, such as polymerase chain reaction, are
increasingly popular due to their specificity and speed, compared to culture-based methods.[176] These
methods also allow the detection and identification of "viable but nonculturable" cells that are metabolically
active but non-dividing.[177] However, even using these improved methods, the total number of bacterial
species is not known and cannot even be estimated with any certainty. Following present classification,
there are a little less than 9,300 known species of prokaryotes, which includes bacteria and archaea;[178]
but attempts to estimate the true number of bacterial diversity have ranged from 107 to 109 total species—
and even these diverse estimates may be off by many orders of magnitude.[179][180]
Commensals
The word "commensalism" is derived from the word "commensal", meaning "eating at the same table"[183]
and all plants and animals are colonised by commensal bacteria. In humans and other animals millions of
them live on the skin, the airways, the gut and other orifices.[184][185] Referred to as "normal flora"[186] or
"commensals",[187] these bacteria usually cause no harm but can invade other sites of the body and give
rise infections. Escherichia coli is a commensal in the human gut but can cause urinary tract infections.[188]
Similarly, streptoccoci, which are part of the normal
flora of the human mouth, can cause heart
disease.[189]
Predators
Mutualists
Certain bacteria form close spatial associations that are essential for their survival. One such mutualistic
association, called interspecies hydrogen transfer, occurs between clusters of anaerobic bacteria that
consume organic acids, such as butyric acid or propionic acid, and produce hydrogen, and methanogenic
archaea that consume hydrogen.[194] The bacteria in this association are unable to consume the organic
acids as this reaction produces hydrogen that accumulates in their surroundings. Only the intimate
association with the hydrogen-consuming archaea keeps the hydrogen concentration low enough to allow
the bacteria to grow.[195]
In soil, microorganisms that reside in the rhizosphere (a zone that includes the root surface and the soil that
adheres to the root after gentle shaking) carry out nitrogen fixation, converting nitrogen gas to nitrogenous
compounds.[196] This serves to provide an easily absorbable form of nitrogen for many plants, which
cannot fix nitrogen themselves. Many other bacteria are found as symbionts in humans and other
organisms. For example, the presence of over 1,000 bacterial species in the normal human gut flora of the
intestines can contribute to gut immunity, synthesise vitamins, such as folic acid, vitamin K and biotin,
convert sugars to lactic acid (see Lactobacillus), as well as fermenting complex undigestible
carbohydrates.[197][198][199] The presence of this gut flora also inhibits the growth of potentially
pathogenic bacteria (usually through competitive exclusion) and these beneficial bacteria are consequently
sold as probiotic dietary supplements.[200]
Nearly all animal life is dependent on bacteria for survival as only bacteria and some archaea possess the
genes and enzymes necessary to synthesize vitamin B12 , also known as cobalamin, and provide it through
the food chain. Vitamin B12 is a water-soluble vitamin that is involved in the metabolism of every cell of
the human body. It is a cofactor in DNA synthesis, and in both fatty acid and amino acid metabolism. It is
particularly important in the normal functioning of the nervous system via its role in the synthesis of
myelin.[201]
Pathogens
The body is continually exposed to many species of bacteria,
including beneficial commensals, which grow on the skin and
mucous membranes, and saprophytes, which grow mainly in the
soil and in decaying matter. The blood and tissue fluids contain
nutrients sufficient to sustain the growth of many bacteria. The
body has defence mechanisms that enable it to resist microbial
invasion of its tissues and give it a natural immunity or innate
resistance against many microorganisms.[202] Unlike some viruses,
bacteria evolve relatively slowly so many bacterial diseases also
occur in other animals.[203]
Colour-enhanced scanning electron
If bacteria form a parasitic association with other organisms, they
micrograph showing Salmonella
are classed as pathogens.[204] Pathogenic bacteria are a major typhimurium (red) invading cultured
cause of human death and disease and cause infections such as human cells
tetanus (caused by Clostridium tetani), typhoid fever, diphtheria,
syphilis, cholera, foodborne illness, leprosy (caused by
Micobacterium leprae) and tuberculosis (caused by Mycobacterium tuberculosis).[205] A pathogenic cause
for a known medical disease may only be discovered many years later, as was the case with Helicobacter
pylori and peptic ulcer disease.[206] Bacterial diseases are also important in agriculture, with bacteria
causing leaf spot, fire blight and wilts in plants, as well as Johne's disease, mastitis, salmonella and anthrax
in farm animals.[207]
Bacterial infections may be treated with antibiotics, which are classified as bacteriocidal if they kill bacteria
or bacteriostatic if they just prevent bacterial growth. There are many types of antibiotics, and each class
inhibits a process that is different in the pathogen from that found in the host. An example of how
antibiotics produce selective toxicity are chloramphenicol and puromycin, which inhibit the bacterial
ribosome, but not the structurally different eukaryotic ribosome.[215] Antibiotics are used both in treating
human disease and in intensive farming to promote animal growth, where they may be contributing to the
rapid development of antibiotic resistance in bacterial populations.[216] Infections can be prevented by
antiseptic measures such as sterilising the skin prior to piercing it with the needle of a syringe, and by
proper care of indwelling catheters. Surgical and dental instruments are also sterilised to prevent
contamination by bacteria. Disinfectants such as bleach are used to kill bacteria or other pathogens on
surfaces to prevent contamination and further reduce the risk of infection.[217]
The ability of bacteria to degrade a variety of organic compounds is remarkable and has been used in waste
processing and bioremediation. Bacteria capable of digesting the hydrocarbons in petroleum are often used
to clean up oil spills.[220] Fertiliser was added to some of the beaches in Prince William Sound in an
attempt to promote the growth of these naturally occurring bacteria after the 1989 Exxon Valdez oil spill.
These efforts were effective on beaches that were not too thickly covered in oil. Bacteria are also used for
the bioremediation of industrial toxic wastes.[221] In the chemical industry, bacteria are most important in
the production of enantiomerically pure chemicals for use as pharmaceuticals or agrichemicals.[222]
Bacteria can also be used in the place of pesticides in the biological pest control. This commonly involves
Bacillus thuringiensis (also called BT), a Gram-positive, soil dwelling bacterium. Subspecies of this
bacteria are used as a Lepidopteran-specific insecticides under trade names such as Dipel and
Thuricide.[223] Because of their specificity, these pesticides are regarded as environmentally friendly, with
little or no effect on humans, wildlife, pollinators and most other beneficial insects.[224][225]
Because of their ability to quickly grow and the relative ease with which they can be manipulated, bacteria
are the workhorses for the fields of molecular biology, genetics and biochemistry. By making mutations in
bacterial DNA and examining the resulting phenotypes, scientists can determine the function of genes,
enzymes and metabolic pathways in bacteria, then apply this knowledge to more complex organisms.[226]
This aim of understanding the biochemistry of a cell reaches its most complex expression in the synthesis of
huge amounts of enzyme kinetic and gene expression data into mathematical models of entire organisms.
This is achievable in some well-studied bacteria, with models of Escherichia coli metabolism now being
produced and tested.[227][228] This understanding of bacterial metabolism and genetics allows the use of
biotechnology to bioengineer bacteria for the production of therapeutic proteins, such as insulin, growth
factors, or antibodies.[229][230]
Because of their importance for research in general, samples of bacterial strains are isolated and preserved
in Biological Resource Centers. This ensures the availability of the strain to scientists worldwide.[231]
History of bacteriology
Bacteria were first observed by the Dutch microscopist Antonie van Leeuwenhoek in 1676, using a single-
lens microscope of his own design. He then published his observations in a series of letters to the Royal
Society of London.[232] Bacteria were Leeuwenhoek's most remarkable microscopic discovery. They were
just at the limit of what his simple lenses could make out and, in one of the most striking hiatuses in the
history of science, no one else would see them again for over a century.[233] His observations had also
included protozoans which he called animalcules, and his findings were looked at again in the light of the
more recent findings of cell theory.[234]
Christian Gottfried Ehrenberg introduced the word "bacterium" in
1828.[235] In fact, his Bacterium was a genus that contained non-
spore-forming rod-shaped bacteria,[236] as opposed to Bacillus, a
genus of spore-forming rod-shaped bacteria defined by Ehrenberg
in 1835.[237]
Robert Koch, a pioneer in medical microbiology, worked on cholera, anthrax and tuberculosis. In his
research into tuberculosis Koch finally proved the germ theory, for which he received a Nobel Prize in
1905.[240] In Koch's postulates, he set out criteria to test if an organism is the cause of a disease, and these
postulates are still used today.[241]
Ferdinand Cohn is said to be a founder of bacteriology, studying bacteria from 1870. Cohn was the first to
classify bacteria based on their morphology.[242][243]
Though it was known in the nineteenth century that bacteria are the cause of many diseases, no effective
antibacterial treatments were available.[244] In 1910, Paul Ehrlich developed the first antibiotic, by
changing dyes that selectively stained Treponema pallidum—the spirochaete that causes syphilis—into
compounds that selectively killed the pathogen.[245] Ehrlich had been awarded a 1908 Nobel Prize for his
work on immunology, and pioneered the use of stains to detect and identify bacteria, with his work being
the basis of the Gram stain and the Ziehl–Neelsen stain.[246]
A major step forward in the study of bacteria came in 1977 when Carl Woese recognised that archaea have
a separate line of evolutionary descent from bacteria.[247] This new phylogenetic taxonomy depended on
the sequencing of 16S ribosomal RNA, and divided prokaryotes into two evolutionary domains, as part of
the three-domain system.[2]
See also
Genetically modified bacteria
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External links
On-line text book on bacteriology (2015) (http://www.textbookofbacteriology.net/)
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