Bacteria ( i/bktri/; common noun bacteria, singular bacterium) constitute a

large domain of prokaryoticmicroorganisms. Typically a few micrometres in length, bacteria

have a number of shapes, ranging from spheres to rods and spirals. Bacteria were among the
first life forms to appear on Earth, and are present in most of its habitats. Bacteria inhabit soil,
water, acidic hot springs, radioactive waste,[4] and the deep portions of Earth's crust. Bacteria
also live in symbiotic and parasitic relationships with plants and animals.
There are typically 40 million bacterial cells in a gram of soil and a million bacterial cells in a
millilitre of fresh water. There are approximately 51030 bacteria on Earth,[5] forming
a biomass which exceeds that of all plants and animals.[6] Bacteria are vital in recycling
nutrients, with many of the stages in nutrient cycles dependent on these organisms, such as
the fixation of nitrogen from the atmosphere and putrefaction. In the biological communities
surrounding hydrothermal vents and cold seeps, bacteria provide the nutrients needed to
sustain life by converting dissolved compounds, such as hydrogen sulphide and methane, to
energy. On 17 March 2013, researchers reported data that suggested bacterial life forms
thrive in the Mariana Trench, which with a depth of up to 11 kilometres is the deepest part of
the Earth's oceans.[7][8] Other researchers reported related studies that microbes thrive
inside rocks up to 580 metres below the sea floor under 2.6 kilometres of ocean off the coast
of the northwestern United States.[7][9] According to one of the researchers, "You can find
microbes everywherethey're extremely adaptable to conditions, and survive wherever they
are."[7]
Most bacteria have not been characterised, and only about half of the bacterial phyla have
species that can be grownin the laboratory.[10] The study of bacteria is known as bacteriology,
a branch of microbiology.
There are approximately ten times as many bacterial cells in the human flora as there are
human cells in the body, with the largest number of the human flora being in the gut flora,
and a large number on the skin.[11] The vast majority of the bacteria in the body are rendered
harmless by the protective effects of the immune system, and some are beneficial. However,
several species of bacteria are pathogenic and cause infectious diseases,
including cholera,syphilis, anthrax, leprosy, and bubonic plague. The most common fatal
bacterial diseases are respiratory infections, with tuberculosis alone killing about 2 million
people per year, mostly in sub-Saharan Africa.[12] In developed countries, antibiotics are used
to treat bacterial infections and are also used in farming, making antibiotic resistance a
growing problem. In industry, bacteria are important in sewage treatment and the breakdown
of oil spills, the production of cheese and yogurt through fermentation, and the recovery of
gold, palladium, copper and other metals in the mining sector,[13] as well as in biotechnology,
and the manufacture of antibiotics and other chemicals.[14]
Once regarded as plants constituting the class Schizomycetes, bacteria are now classified
as prokaryotes. Unlike cells of animals and other eukaryotes, bacterial cells do not contain
a nucleus and rarely harbour membrane-boundorganelles. Although the
term bacteria traditionally included all prokaryotes, the scientific classification changed after
the discovery in the 1990s that prokaryotes consist of two very different groups of organisms
that evolved from an ancient common ancestor. These evolutionary domains are
called Bacteria and Archaea.[1]

Etymology
Life timeline
view discuss edit
-4500


-4000

-3500

-3000

-2500

-2000

-1500

-1000

-500

water
Single-celled
life
photosynthesis

Eukaryotes
Multicellular
life
Land life
Dinosaurs
Mammals
Flowers

Earliest Earth (4540)

Earliest water

Earliest life
(4100)

LHB meteorites

Earliest oxygen

Atmospheric oxygen

Oxygen Crisis

Earliest sexual reproduction

Cambrian explosion

Earliest humans

P
h
a
n
r
z
c

P
r
o
t
e
r
o
z
o
i
c

A
r
c
h
e
a
n
H
a
d
e
a
n

Axis scale: millions of years.

Also see: Human timeline and Nature timeline

The word bacteria is the plural of the New Latin bacterium, which is the latinisation of
the Greek (bakterion),[15] the diminutive of (bakteria), meaning "staff,
cane",[16] because the first ones to be discovered were rod-shaped. [17][18]

Origin and early evolution

Further information: Timeline of evolution and Evolutionary history of life
The ancestors of modern bacteria were unicellular microorganisms that were the first forms of
life to appear on Earth, about 4 billion years ago. For about 3 billion years, most organisms
were microscopic, and bacteria and archaea were the dominant forms of life. [19][20] In 2008,
fossils of macroorganisms were discovered and named as the Francevillian biota. Although
bacterial fossils exist, such as stromatolites, their lack of distinctive morphologyprevents
them from being used to examine the history of bacterial evolution, or to date the time of
origin of a particular bacterial species. However, gene sequences can be used to reconstruct
the bacterial phylogeny, and these studies indicate that bacteria diverged first from the
archaeal/eukaryotic lineage.[21] Bacteria were also involved in the second great evolutionary
divergence, that of the archaea and eukaryotes. Here, eukaryotes resulted from the entering
of ancient bacteria into endosymbiotic associations with the ancestors of eukaryotic cells,
which were themselves possibly related to the Archaea.[22][23] This involved the engulfment

by proto-eukaryotic cells ofalphaproteobacterial symbionts to form

either mitochondria or hydrogenosomes, which are still found in all known Eukarya
(sometimes in highly reduced form, e.g. in ancient "amitochondrial" protozoa). Later on, some
eukaryotes that already contained mitochondria also engulfed cyanobacterial-like organisms.
This led to the formation ofchloroplasts in algae and plants. There are also some algae that
originated from even later endosymbiotic events. Here, eukaryotes engulfed a eukaryotic
algae that developed into a "second-generation" plastid.[24][25] This is known as secondary
endosymbiosis.

Morphology
Further information: Bacterial cell structure Cell morphology
Further information: Bacterial morphological plasticity

Bacteria display many cell morphologies and arrangements

Bacteria display a wide diversity of shapes and sizes, called morphologies. Bacterial cells are
about one-tenth the size of eukaryotic cells and are typically 0.55.0 micrometres in length.
However, a few species are visible to the unaided eyefor example,Thiomargarita
namibiensis is up to half a millimetre long[26] and Epulopiscium fishelsoni reaches 0.7 mm.
[27] Among the smallest bacteria are members of the genus Mycoplasma, which measure only
0.3 micrometres, as small as the largest viruses.[28] Some bacteria may be even smaller, but
theseultramicrobacteria are not well-studied.[29]
Most bacterial species are either spherical, called cocci (sing. coccus, from Greek kkkos,
grain, seed), or rod-shaped, called bacilli (sing. bacillus, from Latin baculus, stick). Elongation
is associated with swimming.[30] Some bacteria, called vibrio, are shaped like slightly curved
rods or comma-shaped; others can be spiral-shaped, called spirilla, or tightly coiled,
called spirochaetes. A small number of species even have tetrahedral or cuboidal shapes.
[31] More recently, some bacteria were discovered deep under Earth's crust that grow as
branching filamentous types with a star-shaped cross-section. The large surface area to
volume ratio of this morphology may give these bacteria an advantage in nutrient-poor
environments.[32]This wide variety of shapes is determined by the bacterial cell
wall and cytoskeleton, and is important because it can influence the ability of bacteria to
acquire nutrients, attach to surfaces, swim through liquids and escape predators.[33][34]

A biofilm of thermophilic bacteria in the outflow of Mickey Hot Springs,Oregon, approximately 20 mm thick.

Many bacterial species exist simply as single cells, others associate in characteristic
patterns: Neisseria form diploids (pairs),Streptococcus form chains, and Staphylococcus group
together in "bunch of grapes" clusters. Bacteria can also be elongated to form filaments, for
example the Actinobacteria. Filamentous bacteria are often surrounded by a sheath that
contains many individual cells. Certain types, such as species of the genus Nocardia, even
form complex, branched filaments, similar in appearance to fungal mycelia.[35]
Bacteria often attach to surfaces and form dense aggregations called biofilms or bacterial
mats. These films can range from a few micrometres in thickness to up to half a metre in
depth, and may contain multiple species of bacteria, protists and archaea. Bacteria living in
biofilms display a complex arrangement of cells and extracellular components, forming
secondary structures, such as microcolonies, through which there are networks of channels to
enable better diffusion of nutrients.[36][37] In natural environments, such as soil or the
surfaces of plants, the majority of bacteria are bound to surfaces in biofilms. [38] Biofilms are
also important in medicine, as these structures are often present during chronic bacterial
infections or in infections of implantedmedical devices, and bacteria protected within biofilms
are much harder to kill than individual isolated bacteria. [39]
Even more complex morphological changes are sometimes possible. 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; this type of cooperation is a simple type
of multicellular organisation. For example, about one in 10 cells migrate to the top of these
fruiting bodies and differentiate into a specialised dormant state called myxospores, which are
more resistant to drying and other adverse environmental conditions than are ordinary cells.
[41]

The bacterial cell is surrounded by a cell membrane (also known as a lipid, cytoplasmic or
plasma membrane). This membrane encloses the contents of the cell and acts as a barrier to
hold nutrients, proteins and other essential components of thecytoplasm within the cell. As
they are prokaryotes, bacteria do not usually have membrane-bound organelles in their
cytoplasm, and thus contain few large intracellular structures. They lack a
true nucleus, mitochondria, chloroplasts and the other organelles present in eukaryotic cells.
[42] Bacteria were once seen as simple bags of cytoplasm, but structures such as
the prokaryotic cytoskeleton[43][44] and the localisation of proteins to specific locations
within the cytoplasm[43] that give bacteria some complexity have been discovered. These
subcellular levels of organisation have been called "bacterial hyperstructures". [45]
Bacterial microcompartments, such as carboxysomes,[46] provide a further level of
organisation; they are compartments within bacteria that are surrounded
by polyhedral protein shells, rather than by lipid membranes.[47] These "polyhedral
organelles" localise and compartmentalise bacterial metabolism, a function performed by the
membrane-bound organelles in eukaryotes.[48][49]

Many important biochemical reactions, such as energy generation, use concentration

gradients across membranes. The general lack of internal membranes in bacteria means
reactions such as electron transport occur across the cell membrane between the cytoplasm
and the periplasmic space.[50] However, in many photosynthetic bacteria the plasma
membrane is highly folded and fills most of the cell with layers of light-gathering membrane.
[51] These light-gathering complexes may even form lipid-enclosed structures
calledchlorosomes in green sulfur bacteria.[52] Other proteins import nutrients across the cell
membrane, or expel undesired molecules from the cytoplasm.

Carboxysomes are protein-enclosed bacterial organelles. Top left is anelectron microscope image of
carboxysomes in Halothiobacillus neapolitanus, below is an image of purified carboxysomes. On the right is a
model of their structure. Scale bars are 100 nm.[53]

Bacteria do not have a membrane-bound nucleus, and their genetic material is typically a
single circular DNA chromosome located in the cytoplasm in an irregularly shaped body called
the nucleoid.[54] The nucleoid contains the chromosome with its associated proteins and RNA.
The phylum Planctomycetes[55] and candidate phylum Poribacteria[56] may be exceptions to
the general absence of internal membranes in bacteria, because they appear to have a
double membrane around their nucleoids and contain other membrane-bound cellular
structures. Like all living organisms, bacteria contain ribosomes, often grouped in chains
called polyribosomes, for the production of proteins, but the structure of the bacterial
ribosome is different from that of eukaryotes and Archaea.[57] Bacterial ribosomes have a
sedimentation rate of 70S (measured in Svedberg units): their subunits have rates
of 30S and 50S. Some antibiotics bind specifically to 70S ribosomes and inhibit bacterial
protein synthesis. Those antibiotics kill bacteria without affecting the larger 80Sribosomes of
eukaryotic cells and without harming the host.
Some bacteria produce intracellular nutrient storage granules for later use, such asglycogen,
[58] polyphosphate,[59] sulfur[60] or polyhydroxyalkanoates.[61] Certain bacterial species, such
as the photosynthetic Cyanobacteria, produce internal gas vesicles, which they use to
regulate their buoyancyallowing them to move up or down into water layers with different
light intensities and nutrient levels.[62] Intracellular membranes called chromatophores are
also found in membranes of phototrophic bacteria. Used primarily for photosynthesis, they
contain bacteriochlorophyll pigments and carotenoids. An early idea was that bacteria might
contain membrane folds termedmesosomes, but these were later shown to be artefacts
produced by the chemicals used to prepare the cells for electron microscopy. Inclusions are
considered to be nonliving components of the cell that do not possess metabolic activity and
are not bounded by membranes. The most common inclusions are glycogen, lipid droplets,
crystals, and pigments. Volutin granules are cytoplasmic inclusions of complexed inorganic
polyphosphate. These granules are called metachromatic granules due to their displaying the
metachromatic effect; they appear red or blue when stained with the blue dyes methylene
blue or toluidine blue. Gas vacuoles, which are freely permeable to gas, are membranebound vesicles present in some species of Cyanobacteria. They allow the bacteria to control
their buoyancy. Microcompartments are widespread, membrane-bound organelles that are

made of a protein shell that surrounds and encloses various enzymes. Carboxysomes are
bacterial microcompartments that contain enzymes involved in carbon
fixation. Magnetosomes are bacterial microcompartments, present in magnetotactic bacteria,
that contain magnetic crystals.

Extracellular structures
Further information: Cell envelope
In most bacteria, a cell wall is present on the outside of the cell membrane. The cell
membrane and cell wall comprise the cell envelope. A common bacterial cell wall material
is peptidoglycan (called "murein" in older sources), which is made from polysaccharide chains
cross-linked by peptides containing D-amino acids.[63] Bacterial cell walls are different from
the cell walls of plants and fungi, which are made of cellulose and chitin, respectively.[64] The
cell wall of bacteria is also distinct from that of Archaea, which do not contain peptidoglycan.
The cell wall is essential to the survival of many bacteria, and the antibiotic penicillin is able
to kill bacteria by inhibiting a step in the synthesis of peptidoglycan. [64]
There are broadly speaking two different types of cell wall in bacteria, a thick one in the grampositives and a thinner one in the gram-negatives. The names originate from the reaction of
cells to the Gram stain, a long-standing test for the classification of bacterial species. [65]
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. Lipopolysaccharides, also
called endotoxins, are composed of polysaccharides and lipid A that is responsible for much of
the toxicity of gram-negative bacteria. Most bacteria have the gram-negative cell wall, and
only the Firmicutes and Actinobacteria have the alternative gram-positive arrangement.
[66] These two groups were previously known as the low G+C and high G+C gram-positive
bacteria, respectively. 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.[67] If the bacterial cell wall is entirely removed, it is called a protoplast, whereas if
it is partially removed, it is called a spheroplast. -Lactam antibiotics, such as penicillin,
inhibit the formation of peptidoglycan cross-links in the bacterial cell wall. The
enzyme lysozyme, found in human tears, also digests the cell wall of bacteria and is the
body's main defence against eye infections.
Acid-fast bacteria, such as Mycobacteria, are resistant to decolorisation by acids
during staining procedures. The high mycolic acid content of Mycobacteria, is responsible for
the staining pattern of poor absorption followed by high retention. The most common staining
technique used to identify acid-fast bacteria is the Ziehl-Neelsen stain or acid-fast stain, in
which the acid-fast bacilli are stained bright-red and stand out clearly against a blue
background. L-form bacteria are strains of bacteria that lack cell walls. The main pathogenic
bacteria in this class is Mycoplasma (not to be confused with Mycobacteria).
In many bacteria, an S-layer of rigidly arrayed protein molecules covers the outside of the
cell.[68] This layer provides chemical and physical protection for the cell surface and can act
as a macromolecular diffusion barrier. S-layers have diverse but mostly poorly understood
functions, but are known to act as virulence factors inCampylobacter and contain
surface enzymes in Bacillus stearothermophilus.[69]

Helicobacter pylori electron micrograph, showing multiple flagella on the cell surface

Flagella are rigid protein structures, about 20 nanometres in diameter and up to

20 micrometres in length, that are used formotility. Flagella are driven by the energy released
by the transfer of ions down an electrochemical gradient across the cell membrane.[70]
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 theelectron microscope.
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.[71] 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).[72] They can also generate movement where they are
called type IV pili (see movement, below).