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Bacteria
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This article is about the microorganisms. For the genus, see Bacterium (genus). For other uses, see Bacteria (disambiguation).
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Bacteria
Temporal range: Archean or earlier – present
Had'n
Archean
Proterozoic
Pha.
Scanning electron micrograph of Escherichia coli rods
Scientific classification
Domain: Bacteria
Woese, Kandler & Wheelis, 1990[1]
Phyla
Acidobacteria
Actinobacteria
Aquificae
Armatimonadetes
Bacteroidetes
Caldiserica
Chlamydiae
Chlorobi
Chloroflexi
Chrysiogenetes
Cyanobacteria
Deferribacteres
Deinococcus-Thermus
Dictyoglomi
Elusimicrobia
Fibrobacteres
Firmicutes
Fusobacteria
Gemmatimonadetes
Lentisphaerae
Nitrospirae
Planctomycetes
Proteobacteria
Spirochaetes
Synergistetes
Tenericutes
Thermodesulfobacteria
Thermotogae
Verrucomicrobia
Synonyms
Eubacteria Woese & Fox, 1977[2]
Bacteria (/bækˈtɪəriə/ ( listen); common noun bacteria, singular bacterium) constitute a large domain of prokaryotic microorganisms. 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,[3] and the deep portions of Earth's crust. Bacteria also live in symbiotic and parasitic relationships with plants and animals. Most bacteria have not been characterised, and only about half of the bacterial phyla have species that can be grown in the laboratory.[4] The study of bacteria is known as bacteriology, a branch of microbiology.
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 5×1030 bacteria on Earth,[5] forming a biomass which exceeds that of all plants and animals.[6] Bacteria are vital in many stages of the nutrient cycle by recycling nutrients such as the fixation of nitrogen from the atmosphere. The nutrient cycle includes the decomposition of dead bodies and bacteria are responsible for the putrefaction stage in this process.[7] In the biological communities surrounding hydrothermal vents and cold seeps, extremophile bacteria provide the nutrients needed to sustain life by converting dissolved compounds, such as hydrogen sulphide and methane, to energy. In March 2013, data reported by researchers in October 2012, was published. It was suggested that bacteria thrive in the Mariana Trench, which with a depth of up to 11 kilometres is the deepest known part of the oceans.[8][9] 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.[8][10] According to one of the researchers, "You can find microbes everywhere—they're extremely adaptable to conditions, and survive wherever they are."[8]
The famous notion that bacterial cells in the human body outnumber human cells by a factor of 10:1 has been debunked. There are approximately 39 trillion bacterial cells in the human microbiota as personified by a "reference" 70 kg male 170 cm tall, whereas there are 30 trillion human cells in the body. This means that although they do have the upper hand in actual numbers, it is only by 30%, and not 900%.[11]
The largest number exist in the gut flora, and a large number on the skin.[12] The vast majority of the bacteria in the body are rendered harmless by the protective effects of the immune system, though many are beneficial particularly in the gut flora. 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.[13] 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,[14] as well as in biotechnology, and the manufacture of antibiotics and other chemicals.[15]
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-bound organelles. 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]
Contents
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1 Etymology
2 Origin and early evolution
3 Morphology
4 Cellular structure
4.1 Intracellular structures
4.2 Extracellular structures
4.3 Endospores
5 Metabolism
6 Growth and reproduction
7 Genomes and chromosomes
8 Genetics
8.1 DNA transfer
8.2 Bacteriophages
9 Behaviour
9.1 Secretion
9.2 Bioluminescence
9.3 Multicellularity
9.4 Movement
10 Classification and identification
11 Interactions with other organisms
11.1 Predators
11.2 Mutualists
11.3 Pathogens
12 Significance in technology and industry
13 History of bacteriology
14 See also
15 References
16 Further reading
17 External links
Etymology
Life timeline
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Orange labels: ice ages.
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 βακτήριον (bakterion),[16] the diminutive of βακτηρία (bakteria), meaning "staff, cane",[17] because the first ones to be discovered were rod-shaped.[18][19]
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.[20][21] Although bacterial fossils exist, such as stromatolites, their lack of distinctive morphology prevents 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.[22] The most recent common ancestor of bacteria and archaea was probably a hyperthermophile that lived about 2.5 billion–3.2 billion years ago.[23][24]
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.[25][26] This involved the engulfment by proto-eukaryotic cells of alphaproteobacterial 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, some eukaryotes that already contained mitochondria also engulfed cyanobacteria-like organisms, leading to the formation of chloroplasts in algae and plants.[27][28] This is known as secondary endosymbiosis.
Morphology
Further information: Bacterial cell structure § Cell morphology
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 0.5–5.0 micrometres in length. However, a few species are visible to the unaided eye—for example, Thiomargarita namibiensis is up to half a millimetre long[29] and Epulopiscium fishelsoni reaches 0.7 mm.[30] Among the smallest bacteria are members of the genus Mycoplasma, which measure only 0.3 micrometres, as small as the largest viruses.[31] Some bacteria may be even smaller, but these ultramicrobacteria are not well-studied.[32]
Most bacterial species are either spherical, called cocci (sing. coccus, from Greek kókkos, grain, seed), or rod-shaped, called bacilli (sing. bacillus, from Latin baculus, stick).[33] 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 other unusual shapes have been described, such as star-shaped bacteria.[34] 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.[35][36]
The range of sizes shown by prokaryotes, relative to those of other organisms and biomolecules.
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 group to form larger multicellular structures, such as the elongated filaments of Actinobacteria, the aggregates of Myxobacteria, and the complex hyphae of Streptomyces.[37] 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.[38] 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.[39]
Bacteria often attach to surfaces and form dense aggregations called biofilms, and larger formations known as microbial mats. These biofilms and mats 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.[40][41] In natural environments, such as soil or the surfaces of plants, the majority of bacteria are bound to surfaces in biofilms.[42] Biofilms are also important in medicine, as these structures are often present during chronic bacterial infections or in infections of implanted medical devices, and bacteria protected within biofilms are much harder to kill than individual isolated bacteria.[43]
Cellular structure
Further information: Bacterial cell structure
Structure and contents of a typical gram-positive bacterial cell (seen by the fact that only one cell membrane is present).
Intracellular structures
The bacterial cell is surrounded by a cell membrane which is made primarily of phospholipids. This membrane encloses the contents of the cell and acts as a barrier to hold nutrients, proteins and other essential components of the cytoplasm within the cell.[44] Unlike eukaryotic cells, bacteria usually lack large membrane-bound structures in their cytoplasm such as a nucleus, mitochondria, chloroplasts and the other organelles present in eukaryotic cells.[45] However, some bacteria have protein-bound organelles in the cytoplasm which compartmentalize aspects of bacterial metabolism,[46][47] such as the carboxysome.[48] Additionally, bacteria have a multi-component cytoskeleton to control the localisation of proteins and nucleic acids within the cell, and to manage the process of cell division.[49][50][51]
Many important biochemical reactions, such as energy generation, occur due to concentration gradients across membranes, creating a potential difference analogous to a battery. The general lack of internal membranes in bacteria means these reactions, such as electron transport, occur across the cell membrane between the cytoplasm and the outside of the cell or periplasm.[52] However, in many photosynthetic bacteria the plasma membrane is highly folded and fills most of the cell with layers of light-gathering membrane.[53] These light-gathering complexes may even form lipid-enclosed structures called chlorosomes in green sulfur bacteria.[54]
An electron micrograph of Halothiobacillus neapolitanus cells with carboxysomes inside, with arrows highlighting visible carboxysomes. Scale bars indicate 100 nm.
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.[55] The nucleoid contains the chromosome with its associated proteins and RNA. The phylum Planctomycetes[56] and candidate phylum Poribacteria[57] 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.[58] 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 80S ribosomes of eukaryotic cells and without harming the host.
Some bacteria produce intracellular nutrient storage granules for later use, such as glycogen,[59] polyphosphate,[60] sulfur[61] or polyhydroxyalkanoates.[62] Certain bacterial species, such as the photosynthetic Cyanobacteria, produce internal gas vacuoles, vesicles which they use to regulate their buoyancy—allowing them to move up or down into water layers with different light intensities and nutrient levels.[63] The gas vacuoles are freely permeable to gas.[64] 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 termed mesosomes, 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.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.[65] Bacterial cell walls are different from the cell walls of plants and fungi, which are made of cellulose and chitin, respectively.[66] 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.[66] The cell wall is absent in L-form bacteria. The main pathogenic types are some species of Mycoplasma.
There are broadly speaking two different types of cell wall in bacteria, a thick one in the gram-positives 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.[67]
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.[68] 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.[69] If the bacterial cell wall is entirely removed, it is called a protoplast, whereas if it is partially removed, it is called a spheroplast. β-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. In many bacteria, an S-layer of rigidly arrayed protein molecules covers the outside of the cell.[70] 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 in Campylobacter and contain surface enzymes in Bacillus stearothermophilus.[71]
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 for motility. Flagella are driven by the energy released by the transfer of ions down an electrochemical gradient across the cell membrane.[72]
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. 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.[73] 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).[74] They can also generate movement where they are called type IV pili (see movement, below).
Glycocalyx is produced by many bacteria to surround their cells, and varies in structural complexity: ranging from a disorganised slime layer of extracellular polymeric substances to a highly structured capsule. These structures can protect cells from engulfment by eukaryotic cells such as macrophages (part of the human immune system).[75] They can also act as antigens and be involved in cell recognition, as well as aiding attachment to surfaces and the formation of biofilms.[76]
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.[77]
Endospores
Further information: Endospore
Bacillus anthracis (stained purple) growing in cerebrospinal fluid
Certain genera of gram-positive bacteria, such as Bacillus, Clostridium, Sporohalobacter, Anaerobacter, and Heliobacterium, can form highly resistant, dormant structures called endospores.[78] In almost all cases, one endospore is formed and this is not a reproductive process, although Anaerobacter can make up to seven endospores in a single cell.[79] Endospores have a central core of cytoplasm containing DNA and ribosomes surrounded by a cortex layer and protected by an impermeable and rigid coat. Dipicolinic acid is a chemical compound that composes 5% to 15% of the dry weight of bacterial spores. It is implicated as responsible for the heat resistance of the endospore.
Endospores show no detectable metabolism and can survive extreme physical and chemical stresses, such as high levels of UV light, gamma radiation, detergents, disinfectants, heat, freezing, pressure, and desiccation.[80] In this dormant state, these organisms may remain viable for millions of years,[81][82] and endospores even allow bacteria to survive exposure to the vacuum and radiation in space.[83] According to scientist Dr. Steinn Sigurdsson, "There are viable bacterial spores that have been found that are 40 million years old on Earth—and we know they're very hardened to radiation."[84] Endospore-forming bacteria can also cause disease: for example, anthrax can be contracted by the inhalation of Bacillus anthracis endospores, and contamination of deep puncture wounds with Clostridium tetani endospores causes tetanus.[85]
Metabolism
Further information: Microbial metabolism
Bacteria exhibit an extremely wide variety of metabolic types.[86] 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.[87] Bacterial metabolism is classified into nutritional groups on the basis of three major criteria: the kind of energy used for growth, the source of carbon, and the electron donors used for growth. An additional criterion of respiratory microorganisms are the electron acceptors used for aerobic or anaerobic respiration.[88]
Nutritional types in bacterial metabolism
Nutritional type
Source of energy
Source of carbon
Examples
Phototrophs Sunlight Organic compounds (photoheterotrophs) or carbon fixation (photoautotrophs) Cyanobacteria, Green sulfur bacteria, Chloroflexi, or Purple bacteria
Lithotrophs Inorganic compounds Organic compounds (lithoheterotrophs) or carbon fixation (lithoautotrophs) Thermodesulfobacteria, Hydrogenophilaceae, or Nitrospirae
Organotrophs Organic compounds Organic compounds (chemoheterotrophs) or carbon fixation (chemoautotrophs) Bacillus, Clostridium or Enterobacteriaceae
Carbon metabolism in bacteria is either heterotrophic, where organic carbon compounds are used as carbon sources, or autotrophic, meaning that cellular carbon is obtained by fixing carbon dioxide. Heterotrophic bacteria include parasitic types. Typical autotrophic bacteria are phototrophic cyanobacteria, green sulfur-bacteria and some purple bacteria, but also many chemolithotrophic species, such as nitrifying or sulfur-oxidising bacteria.[89] Energy metabolism of bacteria is either based on phototrophy, the use of light through photosynthesis, or based on chemotrophy, the use of chemical substances for energy, which are mostly oxidised at the expense of oxygen or alternative electron acceptors (aerobic/anaerobic respiration).
Filaments of photosynthetic cyanobacteria
Bacteria are further divided into lithotrophs that use inorganic electron donors and organotrophs that use organic compounds as electron donors. Chemotrophic organisms use the respective electron donors for energy conservation (by aerobic/anaerobic respiration or fermentation) and biosynthetic reactions (e.g., carbon dioxide fixation), whereas phototrophic organisms use them only for biosynthetic purposes. Respiratory organisms use chemical compounds as a source of energy by taking electrons from the reduced substrate and transferring them to a terminal electron acceptor in a redox reaction. This reaction releases energy that can be used to synthesise ATP and drive metabolism. In aerobic organisms, oxygen is used as the electron acceptor. In anaerobic organisms other inorganic compounds, such as nitrate, sulfate or carbon dioxide are used as electron acceptors. This leads to the ecologically important processes of denitrification, sulfate reduction, and acetogenesis, respectively.
Another way of life of chemotrophs in the absence of possible electron acceptors is fermentation, wherein the electrons taken from the reduced substrates are transferred to oxidised intermediates to generate reduced fermentation products (e.g., lactate, ethanol, hydrogen, butyric acid). Fermentation is possible, because the energy content of the substrates is higher than that of the products, which allows the organisms to synthesise ATP and drive their metabolism.[90][91]
These 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.[92] 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.
Lithotrophic bacteria can use inorganic compounds as a source of energy. Common inorganic electron donors are hydrogen, carbon monoxide, ammonia (leading to nitrification), ferrous iron and other reduced metal ions, and several reduced sulfur compounds. In unusual circumstances, the gas methane can be used by methanotrophic bacteria as both a source of electrons and a substrate for carbon anabolism.[93] In both aerobic phototrophy and chemolithotrophy, oxygen is used as a terminal electron acceptor, whereas under anaerobic conditions inorganic compounds are used instead. Most lithotrophic organisms are autotrophic, whereas organotrophic organisms are heterotrophic.
In addition to fixing carbon dioxide in photosynthesis, some bacteria also fix nitrogen gas using the enzyme nitrogenase. This environmentally important trait can be found in most bacteria of the metabolic types listed above.[94]
Regardless of the type of metabolic process they employ, the majority of bacteria are able to take in raw materials only in the form of relatively small molecules, which enter the cell by diffusion or through molecular channels in cell membranes. The Planctomycetes are the exception (as they are in possessing membranes around their nuclear material). It has recently been shown that Gemmata obscuriglobus is able to take in large molecules via a process that in some ways resembles endocytosis, the process used by eukaryotic cells to engulf external items.[30][95]
Growth and reproduction
Many bacteria reproduce through binary fission, which is compared to mitosis and meiosis in this image.
Further information: Bacterial growth
Unlike in multicellular organisms, increases in cell size (cell growth) and reproduction by cell division are tightly linked in unicellular organisms. Bacteria grow to a fixed size and then reproduce through binary fission, a form of asexual reproduction.[96] Under optimal conditions, bacteria can grow and divide extremely rapidly, and bacterial populations can double as quickly as every 9.8 minutes.[97] In cell division, two identical clone daughter cells are produced. Some bacteria, while still reproducing asexually, form more complex reproductive structures that help disperse the newly formed daughter cells. Examples include fruiting body formation by Myxobacteria and aerial hyphae formation by Streptomyces, or budding. Budding involves a cell forming a protrusion that breaks away and produces a daughter cell.
A colony of Escherichia coli[98]
In the laboratory, bacteria are usually grown using solid or liquid media. Solid growth media, such as agar plates, are used to isolate pure cultures of a bacterial strain. However, liquid growth media are used when 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 specific organisms.[99]
Most laboratory techniques for growing bacteria use high levels of nutrients to produce large amounts of cells cheaply and quickly. However, in natural environments, nutrients are limited, meaning that bacteria cannot continue to reproduce indefinitely. This nutrient limitation has led the evolution of different growth strategies (see r/K selection 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.[100] Other organisms have adaptations to harsh environments, such as the production of multiple antibiotics by Streptomyces that inhibit the growth of competing microorganisms.[101] In nature, many organisms live in communities (e.g., biofilms) that may allow for increased supply of nutrients and protection from environmental stresses.[42] These relationships can be essential for growth of a particular organism or group of organisms (syntrophy).[102]
Bacterial growth follows four phases. When a population of bacteria first enter a high-nutrient environment that allows growth, the cells need to adapt to their new environment. The first phase of growth is the lag phase, a period of slow growth when the cells are adapting to the high-nutrient environment and preparing for fast growth. The lag phase has high biosynthesis rates, as proteins necessary for rapid growth are produced.[103] The second phase of growth is the log phase, also known as the logarithmic or exponential phase. The log phase is marked by rapid exponential growth. The rate at which cells grow during this phase is known as the growth rate (k), and the time it takes the cells to double is known as the generation time (g). During log phase, nutrients are metabolised at maximum speed until one of the nutrients is depleted and starts limiting growth. The third phase of growth is the stationary phase and is caused by depleted nutrients. The cells reduce their metabolic activity and consume non-essential cellular proteins. The stationary phase is a transition from rapid growth to a stress response state and there is increased expression of genes involved in DNA repair, antioxidant metabolism and nutrient transport.[104] The final phase is the death phase where the bacteria run out of nutrients and die.
Genomes and chromosomes
Further information: Plasmid and Genome
The genomes of thousands of bacterial species have been sequenced, with at least 9,000 sequences completed and more than 42,000 left as "permanent" drafts (as of Sep 2016).[105]
Chromosome. Most bacteria have a single circular chromosome that can range in size from only 160,000 base pairs (bp) in the endosymbiotic bacteria Candidatus Carsonella ruddii,[106] to 12,200,000 base pairs (12.2 Mbp) in the soil-dwelling bacteria Sorangium cellulosum.[107] The genes in bacterial genomes are usually a single continuous stretch of DNA and although several different types of introns do exist in bacteria, these are much rarer than in eukaryotes.[108]
Multiple chromosomes have been found in the genus Vibrio. For instance, Vibrio cholerae, the cause of cholera, and V. parahaemolyticus, have 2 chromosomes each.[109] In fact, 10% of all sequenced bacterial genomes are estimated to have 2 or more chromosomes.[109]
Some bacteria, e.g. Borrelia burgdorferi, the cause of Lyme disease, contains a single linear chromosome and several linear and circular plasmids.[110][111] The chromosome of Streptomyces lividans is linear too[112] and there is evidence that this is true in most species of Streptomyces.[110]
Plasmids are small extra-chromosomal DNAs that may contain genes for antibiotic resistance, enzymes that degrade unusual organic substrates, toxins that kill other bacteria, or virulence factors.[113] Plasmids replicate independently of chromosomes and often encode partition systems that ensure each daughter cell receives a copy of the plasmid following cell division. Partition systems are also encoded by most bacterial chromosomes. Plasmids that are maintained at a high copy number per cell typically lack partition systems because plasmids will randomly assort to both daughter cells.[114]
Genetics
Bacteria, as asexual organisms, inherit an identical copy of the parent's genes and are clonal. However, all bacteria can evolve by selection on changes to their genetic material DNA caused by genetic recombination or mutations. Mutations come from errors made during the replication of DNA or from exposure to mutagens. Mutation rates vary widely among different species of bacteria and even among different clones of a single species of bacteria.[115] Genetic changes in bacterial genomes come from either random mutation during replication or "stress-directed mutation", where genes involved in a particular growth-limiting process have an increased mutation rate.[116]
DNA transfer
Some bacteria also transfer genetic material between cells. This can occur in three main ways. First, bacteria can take up exogenous DNA from their environment, in a process called transformation. Genes can also be transferred by the process of transduction, when the integration of a bacteriophage introduces foreign DNA into the chromosome. The third method of gene transfer is conjugation, whereby DNA is transferred through direct cell contact.
Transduction of bacterial genes by bacteriophage is a consequence of infrequent errors during intracellular assembly of virus particles. Conjugation, in the much-studied E. coli system, is determined by plasmid genes that encode the machinery necessary to transfer a new copy of the plasmid DNA from one bacterial host to another. It is seldom that a conjugative plasmid integrates into the host bacterial chromosome and subsequently transfers part of the host bacterial DNA to another bacterium.
Transformation, unlike transduction or conjugation, depends on numerous bacterial gene products that specifically interact to perform this complex process.[117] In order for a bacterium to bind, take up and recombine donor DNA into its own chromosome, it must first enter a special physiological state termed competence (see Natural competence). In Bacillus subtilis, about 40 genes are required for the development of competence.[118] The length of DNA transferred during B. subtilis transformation can be between a third of a chromosome up to the whole chromosome.[119][120] Transformation appears to be common among bacterial species, and thus far at least 60 species are known to have the natural ability to become competent for transformation.[121] The development of competence in nature is usually associated with stressful environmental conditions, and seems to be an adaptation for facilitating repair of DNA damage in recipient cells.[122]
In ordinary circumstances, transduction, conjugation, and transformation involve transfer of DNA between individual bacteria of the same species, but occasionally transfer may occur between individuals of different bacterial species and this may have significant consequences, such as the transfer of antibiotic resistance.[123] In such cases, gene acquisition from other bacteria or the environment is called horizontal gene transfer and may be common under natural conditions.[124] Gene transfer is particularly important in antibiotic resistance as it allows the rapid transfer of resistance genes between different pathogens.[125]
Bacteriophages
Main article: Bacteriophage
Bacteriophages are viruses that infect bacteria. Many types of bacteriophage exist, some simply infect and lyse their host bacteria, while others insert into the bacterial chromosome. A bacteriophage can contain genes that contribute to its host's phenotype: for example, in the evolution of Escherichia coli O157:H7 and Clostridium botulinum, the toxin genes in an integrated phage converted a harmless ancestral bacterium into a lethal pathogen.[126] Bacteria resist phage infection through restriction modification systems that degrade foreign DNA,[127] and a system that uses CRISPR sequences to retain fragments of the genomes of phage that the bacteria have come into contact with in the past, which allows them to block virus replication through a form of RNA interference.[128][129] This CRISPR system provides bacteria with acquired immunity to infection.
Behaviour
Secretion
Bacteria frequently secrete chemicals into their environment in order to modify it favourably. The secretions are often proteins and may act as enzymes that digest some form of food in the environment.
Bioluminescence
Further information: Milky seas effect
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.[130]
Multicellularity
See also: Prokaryote § Sociality
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.[131][132]
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.[131] For example, bacteria in biofilms can have more than 500 times increased resistance to antibacterial agents than individual "planktonic" bacteria of the same species.[132]
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.
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.[133]
Movement
Further information: Chemotaxis, Flagellum, and Pilus
Flagellum of gram-negative bacteria. The base drives the rotation of the hook and filament.
Many bacteria can move using a variety of mechanisms: flagella are used for swimming through fluids; bacterial gliding and twitching motility move bacteria across surfaces; and changes of buoyancy allow vertical motion.[134]
Swimming bacteria frequently move near 10 body lengths per second and a few as fast as 100. This makes them at least as fast as fish, on a relative scale.[135]
In bacterial gliding and twitching motility, bacteria use their type IV pili as a grappling hook, repeatedly extending it, anchoring it and then retracting it with remarkable force (>80 pN).[136]
Our observations redefine twitching motility as a rapid, highly organized mechanism of bacterial translocation by which Pseudomonas aeruginosa can disperse itself over large areas to colonize new territories. It is also now clear, both morphologically and genetically, that twitching motility and social gliding motility, such as occurs in Myxococcus xanthus, are essentially the same process.
—  Semmler, Whitchurch & Mattick (1999)[137]
Flagella are semi-rigid cylindrical structures that are rotated and function much like the propeller on a ship. Objects as small as bacteria operate a low Reynolds number and cylindrical forms are more efficient than the flat, paddle-like, forms appropriate at human-size scale.[138]
Bacterial species differ in the number and arrangement of flagella on their surface; some have a single flagellum (monotrichous), a flagellum at each end (amphitrichous), clusters of flagella at the poles of the cell (lophotrichous), while others have flagella distributed over the entire surface of the cell (peritrichous). The bacterial flagella is the best-understood motility structure in any organism and is made of about 20 proteins, with approximately another 30 proteins required for its regulation and assembly.[134] 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] This motor drives the motion of the filament, which acts as a propeller.
Many bacteria (such as E. coli) have two distinct modes of movement: forward movement (swimming) and tumbling. The tumbling allows them to reorient and makes their movement a three-dimensional random walk.[140] (See external links below for link to videos.) The flagella of a unique group of bacteria, the spirochaetes, are found between two membranes in the periplasmic space. They have a distinctive helical body that twists about as it moves.[134]
Motile bacteria are attracted or repelled by certain stimuli in behaviours called taxes: these include chemotaxis, phototaxis, energy taxis, and magnetotaxis.[141][142][143] In one peculiar group, the myxobacteria, individual bacteria move together to form waves of cells that then differentiate to form fruiting bodies containing spores.[39] The myxobacteria move only when on solid surfaces, unlike E. coli, which is motile in liquid or solid media.
Several Listeria and Shigella species move inside host cells by usurping the cytoskeleton, which is normally used to move organelles inside the cell. By promoting actin polymerisation at one pole of their cells, they can form a kind of tail that pushes them through the host cell's cytoplasm.[144]
Classification and identification
Main article: Bacterial taxonomy
Further information: Scientific classification, Systematics, Bacterial phyla, and Clinical pathology
Streptococcus mutans visualised with a Gram stain
Phylogenetic tree showing the diversity of bacteria, compared to other organisms.[145] Eukaryotes are coloured red, archaea green and bacteria blue.

Classification seeks to describe the diversity of bacterial species by naming and grouping organisms based on similarities. Bacteria can be classified on the basis of cell structure, cellular metabolism or on differences in cell components, such as DNA, fatty acids, pigments, antigens and quinones.[99] While these schemes allowed the identification and classification of bacterial strains, it was unclear whether these differences represented variation between distinct species or between strains of the same species. This uncertainty was due to the lack of distinctive structures in most bacteria, as well as lateral gene transfer between unrelated species.[146] Due to lateral gene transfer, some closely related bacteria can have very different morphologies and metabolisms. To overcome this uncertainty, modern bacterial classification emphasises molecular systematics, using genetic techniques such as guanine cytosine ratio determination, genome-genome hybridisation, as well as sequencing genes that have not undergone extensive lateral gene transfer, such as the rRNA gene.[147] Classification of bacteria is determined by publication in the International Journal of Systematic Bacteriology,[148] and Bergey's Manual of Systematic Bacteriology.[149] The International Committee on Systematic Bacteriology (ICSB) maintains international rules for the naming of bacteria and taxonomic categories and for the ranking of them in the International Code of Nomenclature of Bacteria.
The term "bacteria" was traditionally applied to all microscopic, single-cell prokaryotes. However, molecular systematics showed prokaryotic life to consist of two separate domains, originally called Eubacteria and Archaebacteria, but now called Bacteria and Archaea that evolved independently from an ancient common ancestor.[1] The archaea and eukaryotes are more closely related to each other than either is to the bacteria. These two domains, along with Eukarya, are the basis of the three-domain system, which is currently the most widely used classification system in microbiology.[150] However, due to the relatively recent introduction of molecular systematics and a rapid increase in the number of genome sequences that are available, bacterial classification remains a changing and expanding field.[4][151] For example, a few biologists argue that the Archaea and Eukaryotes evolved from gram-positive bacteria.[152]
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.
The Gram stain, developed in 1884 by Hans Christian Gram, characterises bacteria based on the structural characteristics of their cell walls.[67] The thick layers of peptidoglycan in the "gram-positive" cell wall stain purple, while the thin "gram-negative" cell wall appears pink. 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.[153] Other organisms may need to be identified by their growth in special media, or by other techniques, such as serology.
Culture techniques are designed to promote the growth and identify particular bacteria, while restricting the growth of the other bacteria in the sample. 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 diarrhoea, 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.[99][154] 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.
As with bacterial classification, identification of bacteria is increasingly using molecular methods. Diagnostics using DNA-based tools, such as polymerase chain reaction, are increasingly popular due to their specificity and speed, compared to culture-based methods.[155] These methods also allow the detection and identification of "viable but nonculturable" cells that are metabolically active but non-dividing.[156] 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;[157] 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.[158][159]
Interactions with other organisms
Overview of bacterial infections and main species involved.[160][161]
Further information: Microbes in human culture
Despite their apparent simplicity, bacteria can form complex associations with other organisms. These symbiotic associations can be divided into parasitism, mutualism and commensalism. Due to their small size, commensal bacteria are ubiquitous and grow on animals and plants exactly as they will grow on any other surface. However, their growth can be increased by warmth and sweat, and large populations of these organisms in humans are the cause of body odour.
Predators
Some species of bacteria kill and then consume other microorganisms, these species are called predatory bacteria.[162] These include organisms such as Myxococcus xanthus, which forms swarms of cells that kill and digest any bacteria they encounter.[163] Other bacterial predators either attach to their prey in order to digest them and absorb nutrients, such as Vampirovibrio chlorellavorus,[164] or invade another cell and multiply inside the cytosol, such as Daptobacter.[165] These predatory bacteria are thought to have evolved from saprophages that consumed dead microorganisms, through adaptations that allowed them to entrap and kill other organisms.[166]
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.[167] 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.
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.[168] 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.[169][170][171] 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.[172]
Pathogens
Main article: Pathogenic bacteria
Colour-enhanced scanning electron micrograph showing Salmonella typhimurium (red) invading cultured human cells
If bacteria form a parasitic association with other organisms, they are classed as pathogens. Pathogenic bacteria are a major cause of human death and disease and cause infections such as tetanus, typhoid fever, diphtheria, syphilis, cholera, foodborne illness, leprosy and tuberculosis. A pathogenic cause for a known medical disease may only be discovered many years after, as was the case with Helicobacter pylori and peptic ulcer disease. 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.
Each species of pathogen has a characteristic spectrum of interactions with its human hosts. Some organisms, such as Staphylococcus or Streptococcus, can cause skin infections, pneumonia, meningitis and even overwhelming sepsis, a systemic inflammatory response producing shock, massive vasodilation and death.[173] Yet these organisms are also part of the normal human flora and usually exist on the skin or in the nose without causing any disease at all. Other organisms invariably cause disease in humans, such as the Rickettsia, which are obligate intracellular parasites able to grow and reproduce only within the cells of other organisms. One species of Rickettsia causes typhus, while another causes Rocky Mountain spotted fever. Chlamydia, another phylum of obligate intracellular parasites, contains species that can cause pneumonia, or urinary tract infection and may be involved in coronary heart disease.[174] Finally, some species, such as Pseudomonas aeruginosa, Burkholderia cenocepacia, and Mycobacterium avium, are opportunistic pathogens and cause disease mainly in people suffering from immunosuppression or cystic fibrosis.[175][176]
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.[177] 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.[178] 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.
Significance in technology and industry
Further information: Economic importance of bacteria
Bacteria, often lactic acid bacteria, such as Lactobacillus and Lactococcus, in combination with yeasts and moulds, have been used for thousands of years in the preparation of fermented foods, such as cheese, pickles, soy sauce, sauerkraut, vinegar, wine and yogurt.[179][180]
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.[181] 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.[182] In the chemical industry, bacteria are most important in the production of enantiomerically pure chemicals for use as pharmaceuticals or agrichemicals.[183]
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.[184] 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.[185][186]
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.[187] 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.[188][189] 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.[190][191]
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.
History of bacteriology
For the history of microbiology, see Microbiology. For the history of bacterial classification, see Bacterial taxonomy. For the natural history of Bacteria, see Last universal common ancestor.
Antonie van Leeuwenhoek, the first microbiologist and the first person to observe bacteria using a microscope.
Bacteria were first observed by the Dutch microscopist Antonie van Leeuwenhoek in 1676, using a single-lens microscope of his own design.[192] He then published his observations in a series of letters to the Royal Society of London.[193][194][195] 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.[196] 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.
Christian Gottfried Ehrenberg introduced the word "bacterium" in 1828.[197] In fact, his Bacterium was a genus that contained non-spore-forming rod-shaped bacteria,[198] as opposed to Bacillus, a genus of spore-forming rod-shaped bacteria defined by Ehrenberg in 1835.[199]
Louis Pasteur demonstrated in 1859 that the growth of microorganisms causes the fermentation process, and that this growth is not due to spontaneous generation. (Yeasts and moulds, commonly associated with fermentation, are not bacteria, but rather fungi.) Along with his contemporary Robert Koch, Pasteur was an early advocate of the germ theory of disease.[200]
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.[201] 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.[202]
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.[203][204]
Though it was known in the nineteenth century that bacteria are the cause of many diseases, no effective antibacterial treatments were available.[205] 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.[206] 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.[207]
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.[2] 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.[1]
See also
Bacteriotherapy
Genetically modified bacteria
List of bacterial orders
Panspermia
Polysaccharide encapsulated bacteria
Psychrotrophic bacteria
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Jump up ^ Cavalier-Smith T (2002). "The neomuran origin of archaebacteria, the negibacterial root of the universal tree and bacterial megaclassification". Int J Syst Evol Microbiol. 52 (Pt 1): 7–76. doi:10.1099/00207713-52-1-7. PMID 11837318.
Jump up ^ Woods GL, Walker DH (1996). "Detection of infection or infectious agents by use of cytologic and histologic stains". Clinical Microbiology Reviews. 9 (3): 382–404. PMC 172900  . PMID 8809467.
Jump up ^ Weinstein M (1994). "Clinical importance of blood cultures". Clin Lab Med. 14 (1): 9–16. PMID 8181237.
Jump up ^ Louie M, Louie L, Simor AE (8 August 2000). "The role of DNA amplification technology in the diagnosis of infectious diseases". CMAJ. 163 (3): 301–9. PMC 80298  . PMID 10951731.
Jump up ^ Oliver J (2005). "The viable but nonculturable state in bacteria". J Microbiol. 43 Spec No: 93–100. PMID 15765062. Archived from the original on 28 September 2007.
Jump up ^ Euzéby JP (8 December 2011). "Number of published names". List of Prokaryotic names with Standing in Nomenclature. Archived from the original on 19 January 2012. Retrieved 10 December 2011.
Jump up ^ Curtis TP, Sloan WT, Scannell JW (2002). "Estimating prokaryotic diversity and its limits". Proceedings of the National Academy of Sciences of the United States of America. 99 (16): 10494–9. Bibcode:2002PNAS...9910494C. doi:10.1073/pnas.142680199. PMC 124953  . PMID 12097644.
Jump up ^ Schloss PD, Handelsman J (2004). "Status of the microbial census". Microbiology and Molecular Biology Reviews. 68 (4): 686–91. doi:10.1128/MMBR.68.4.686-691.2004. PMC 539005  . PMID 15590780.
Jump up ^ Fisher B, Harvey RP, Champe PC (2007). Lippincott's Illustrated Reviews: Microbiology (Lippincott's Illustrated Reviews Series). Hagerstwon, MD: Lippincott Williams & Wilkins. pp. Chapter 33, pages 367–392. ISBN 0-7817-8215-5.
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Jump up ^ Martin MO (2002). "Predatory prokaryotes: an emerging research opportunity". Journal of Microbiology and Biotechnology. 4 (5): 467–77. PMID 12432957.
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Jump up ^ Gromov BV (1972). "Electron Microscope Study of Parasitism by Bdellovibrio Chorellavorus Bacteria on Cells of the Green Alga Chorella Vulgaris". Tsitologiya. 14 (2): 256–60.
Jump up ^ Guerrero R, Pedros-Alio C, Esteve I, Mas J, Chase D, Margulis L (April 1986). "Predatory prokaryotes: predation and primary consumption evolved in bacteria". Proceedings of the National Academy of Sciences of the United States of America. 83 (7): 2138–42. Bibcode:1986PNAS...83.2138G. doi:10.1073/pnas.83.7.2138. PMC 323246  . PMID 11542073.
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Jump up ^ Fish DN (2002). "Optimal antimicrobial therapy for sepsis". Am J Health Syst Pharm. 59 Suppl 1: S13–9. PMID 11885408.
Jump up ^ Belland RJ, Ouellette SP, Gieffers J, Byrne GI (2004). "Chlamydia pneumoniae and atherosclerosis". Cell Microbiol. 6 (2): 117–27. doi:10.1046/j.1462-5822.2003.00352.x. PMID 14706098.
Jump up ^ Heise ER (1982). "Diseases associated with immunosuppression". Environmental Health Perspectives. 43: 9–19. doi:10.2307/3429162. JSTOR 3429162. PMC 1568899  . PMID 7037390.
Jump up ^ Saiman L (2004). "Microbiology of early CF lung disease". Paediatric Respiratory Reviews. 5 Suppl A: S367–9. doi:10.1016/S1526-0542(04)90065-6. PMID 14980298.
Jump up ^ Yonath A, Bashan A (2004). "Ribosomal crystallography: initiation, peptide bond formation, and amino acid polymerization are hampered by antibiotics". Annu Rev Microbiol. 58: 233–51. doi:10.1146/annurev.micro.58.030603.123822. PMID 15487937.
Jump up ^ Khachatourians GG (1998). "Agricultural use of antibiotics and the evolution and transfer of antibiotic-resistant bacteria". CMAJ. 159 (9): 1129–36. PMC 1229782  . PMID 9835883.
Jump up ^ Johnson ME, Lucey JA (2006). "Major technological advances and trends in cheese". J Dairy Sci. 89 (4): 1174–8. doi:10.3168/jds.S0022-0302(06)72186-5. PMID 16537950.
Jump up ^ Hagedorn S, Kaphammer B (1994). "Microbial biocatalysis in the generation of flavor and fragrance chemicals". Annu. Rev. Microbiol. 48: 773–800. doi:10.1146/annurev.mi.48.100194.004013. PMID 7826026.
Jump up ^ Cohen Y (2002). "Bioremediation of oil by marine microbial mats". Int Microbiol. 5 (4): 189–93. doi:10.1007/s10123-002-0089-5. PMID 12497184.
Jump up ^ Neves LC, Miyamura TT, Moraes DA, Penna TC, Converti A (2006). "Biofiltration methods for the removal of phenolic residues". Appl. Biochem. Biotechnol. 129–132: 130–52. doi:10.1385/ABAB:129:1:130. PMID 16915636.
Jump up ^ Liese A, Filho MV (1999). "Production of fine chemicals using biocatalysis". Current Opinion in Biotechnology. 10 (6): 595–603. doi:10.1016/S0958-1669(99)00040-3. PMID 10600695.
Jump up ^ Aronson AI, Shai Y (2001). "Why Bacillus thuringiensis insecticidal toxins are so effective: unique features of their mode of action". FEMS Microbiol. Lett. 195 (1): 1–8. doi:10.1111/j.1574-6968.2001.tb10489.x. PMID 11166987.
Jump up ^ Bozsik A (2006). "Susceptibility of adult Coccinella septempunctata (Coleoptera: Coccinellidae) to insecticides with different modes of action". Pest Manag Sci. 62 (7): 651–4. doi:10.1002/ps.1221. PMID 16649191.
Jump up ^ Chattopadhyay A, Bhatnagar NB, Bhatnagar R (2004). "Bacterial insecticidal toxins". Crit Rev Microbiol. 30 (1): 33–54. doi:10.1080/10408410490270712. PMID 15116762.
Jump up ^ Serres MH, Gopal S, Nahum LA, Liang P, Gaasterland T, Riley M (2001). "A functional update of the Escherichia coli K-12 genome". Genome Biology. 2 (9): research0035.1–research0035.7. doi:10.1186/gb-2001-2-9-research0035. PMC 56896  . PMID 11574054.
Jump up ^ Almaas E, Kovács B, Vicsek T, Oltvai ZN, Barabási AL (2004). "Global organization of metabolic fluxes in the bacterium Escherichia coli". Nature. 427 (6977): 839–43. arXiv:q-bio/0403001  . Bibcode:2004Natur.427..839A. doi:10.1038/nature02289. PMID 14985762.
Jump up ^ Reed JL, Vo TD, Schilling CH, Palsson BO (2003). "An expanded genome-scale model of Escherichia coli K-12 (iJR904 GSM/GPR)". Genome Biol. 4 (9): R54. doi:10.1186/gb-2003-4-9-r54. PMC 193654  . PMID 12952533.
Jump up ^ Walsh G (2005). "Therapeutic insulins and their large-scale manufacture". Appl Microbiol Biotechnol. 67 (2): 151–9. doi:10.1007/s00253-004-1809-x. PMID 15580495.
Jump up ^ Graumann K, Premstaller A (2006). "Manufacturing of recombinant therapeutic proteins in microbial systems". Biotechnol J. 1 (2): 164–86. doi:10.1002/biot.200500051. PMID 16892246.
Jump up ^ Porter JR (1976). "Antony van Leeuwenhoek: tercentenary of his discovery of bacteria". Bacteriological Reviews. 40 (2): 260–9. PMC 413956  . PMID 786250.
Jump up ^ van Leeuwenhoek A (1684). "An abstract of a letter from Mr. Anthony Leevvenhoek at Delft, dated Sep. 17, 1683, Containing Some Microscopical Observations, about Animals in the Scurf of the Teeth, the Substance Call'd Worms in the Nose, the Cuticula Consisting of Scales". Philosophical Transactions (1683–1775). 14 (155–166): 568–574. doi:10.1098/rstl.1684.0030.
Jump up ^ van Leeuwenhoek A (1700). "Part of a Letter from Mr Antony van Leeuwenhoek, concerning the Worms in Sheeps Livers, Gnats, and Animalcula in the Excrements of Frogs". Philosophical Transactions (1683–1775). 22 (260–276): 509–518. doi:10.1098/rstl.1700.0013.
Jump up ^ van Leeuwenhoek A (1702). "Part of a Letter from Mr Antony van Leeuwenhoek, F. R. S. concerning Green Weeds Growing in Water, and Some Animalcula Found about Them". Philosophical Transactions (1683–1775). 23 (277–288): 1304–11. doi:10.1098/rstl.1702.0042.
Jump up ^ Asimov, Isaac (1982), Asimov's Biographical Encyclopedia of Science and Technology, 2nd edition, Garden City, New York: Doubleday and Company, pg 143.
Jump up ^ Ehrenberg's Symbolae Physioe. Animalia evertebrata. Decas prima. Berlin, 1828.
Jump up ^ Breed RS, Conn HJ (1936). "The Status of the Generic Term Bacterium Ehrenberg 1828". Journal of Bacteriology. 31 (5): 517–518. PMC 543738  . PMID 16559906.
Jump up ^ EHRENBERG (C.G.): Dritter Beitrag zur Erkenntniss grosser Organisation in der Richtung des kleinsten Raumes. Physikalische Abhandlungen der Koeniglichen Akademie der Wissenschaften zu Berlin aus den Jahren 1833–1835, 1835, pp. 143–336.
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Jump up ^ Chung, King-Thom. Ferdinand Julius Cohn (1828-1898): Pioneer of Bacteriology. Department of Microbiology and Molecular Cell Sciences, The University of Memphis.
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Jump up ^ "Biography of Paul Ehrlich". Nobelprize.org. Archived from the original on 28 November 2006. Retrieved 26 November 2006.
Further reading
Library resources about
Bacteria
Online books
Resources in your library
Resources in other libraries
Alcamo IE (2001). Fundamentals of microbiology. Boston: Jones and Bartlett. ISBN 0-7637-1067-9.
Atlas RM (1995). Principles of microbiology. St. Louis: Mosby. ISBN 0-8016-7790-4.
Martinko JM, Madigan MT (2005). Brock Biology of Microorganisms (11th ed.). Englewood Cliffs, N.J: Prentice Hall. ISBN 0-13-144329-1.
Holt JC, Bergey DH (1994). Bergey's manual of determinative bacteriology (9th ed.). Baltimore: Williams & Wilkins. ISBN 0-683-00603-7.
Hugenholtz P, Goebel BM, Pace NR (15 September 1998). "Impact of culture-independent studies on the emerging phylogenetic view of bacterial diversity". J Bacteriol. 180 (18): 4765–74. PMC 107498  . PMID 9733676.
Funke BR, Tortora GJ, Case CL (2004). Microbiology: an introduction (8th ed.). San Francisco: Benjamin Cummings. ISBN 0-8053-7614-3.
Ogunseitan OA (2005). Microbial Diversity: Form and Function in Prokaryotes. Wiley-Blackwell. ISBN 978-1-4051-4448-3.
Shively JM (2006). Complex Intracellular Structures in Prokaryotes (Microbiology Monographs). Berlin: Springer. ISBN 3-540-32524-7.
External links
Find more aboutBacteriaat Wikipedia's sister projects
Definitions from Wiktionary
Media from Wikimedia Commons
News from Wikinews
Quotations from Wikiquote
Texts from Wikisource
Textbooks from Wikibooks
Learning resources from Wikiversity
Taxonomy from Wikispecies
MicrobeWiki, an extensive wiki about bacteria and viruses
Bacteria that affect crops and other plants
Bacterial Nomenclature Up-To-Date from DSMZ
Genera of the domain Bacteria—list of Prokaryotic names with Standing in Nomenclature
The largest bacteria
Tree of Life: Eubacteria
Videos of bacteria swimming and tumbling, use of optical tweezers and other videos.
Planet of the Bacteria by Stephen Jay Gould
On-line text book on bacteriology
Animated guide to bacterial cell structure.
Bacteria Make Major Evolutionary Shift in the Lab
Online collaboration for bacterial taxonomy.
PATRIC, a Bioinformatics Resource Center for bacterial pathogens, funded by NIAID
Bacterial Chemotaxis Interactive Simulator—A web-app that uses several simple algorithms to simulate bacterial chemotaxis.
Cell-Cell Communication in Bacteria on-line lecture by Bonnie Bassler, and TED: Discovering bacteria's amazing communication system
Sulfur-cycling fossil bacteria from the 1.8-Ga Duck Creek Formation provide promising evidence of evolution's null hypothesis, Proceedings of the National Academy of Sciences of the United States of America. Summarised in: Scientists discover bacteria that haven't evolved in more than 2 billion years, LiveScience and BusinessInsider
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Prokaryotes: Bacteria classification (phyla and orders)
Domain
Archaea
Bacteria
Eukaryota
(Supergroup
Plant
Hacrobia
Heterokont
Alveolata
Rhizaria
Excavata
Amoebozoa
Opisthokonta
Animal
Fungi)
G-/
OM
Terra-/
Glidobacteria
(BV1)
Eobacteria
Deinococcus–Thermus
Deinococcales
Thermales
Chloroflexi
Anaerolineales
Caldilineales
Chloroflexales
Herpetosiphonales
Dehalococcoidales
Ktedonobacterales
Thermogemmatisporales
Thermomicrobiales
Sphaerobacterales
other glidobacteria
Thermodesulfobacteria
thermophiles
Aquificae
Thermotogae
Cyanobacteria
Proteobacteria
(BV2)
Alpha
Caulobacterales
Kiloniellales
Kordiimonadales
Magnetococcales
Parvularculales
Rhizobiales
Rhodobacterales
Rhodospirillales
Rickettsiales
Sneathiellales
Sphingomonadales
Beta
Burkholderiales
Hydrogenophilales
Methylophilales
Neisseriales
Nitrosomonadales
Procabacteriales
Rhodocyclales
Gamma
Acidithiobacillales
Aeromonadales
Alteromonadales
Cardiobacteriales
Chromatiales
Enterobacteriales
Legionellales
Methylococcales
Oceanospirillales
Orbales
Pasteurellales
Pseudomonadales
Salinisphaerales
Thiotrichales
Vibrionales
Xanthomonadales
Delta
Bdellovibrionales
Desulfarculales
Desulfobacterales
Desulfovibrionales
Desulfurellales
Desulfuromonadales
Myxococcales
Syntrophobacterales
Syntrophorhabdales
Epsilon
Campylobacterales
Nautiliales
Zeta
Mariprofundales
BV4
Spirochaetes
Spirochaetes
Sphingobacteria
(FCB group)
Fibrobacteres
Chlorobi
Chlorobiales
Ignavibacteriales
Bacteroidetes
Bacteroidales
Cytophagales
Flavobacteriales
Sphingobacteriales
Planctobacteria/
(PVC group)
Chlamydiae
Lentisphaerae
Lentisphaerales
Oligosphaerales
Victivallales
Planctomycetes
Phycisphaerales
Planctomycetales
Verrucomicrobia
Puniceicoccales
Opitutales
Chthoniobacterales
Verrucomicrobiales
"Poribacteria"
Other GN
Acidobacteria
Acidobacteriales
Acanthopleuribacterales
Holophagales
Solibacterales
Armatimonadetes
Armatimonadales
Chthonomonadales
Fimbriimonadales
Caldiserica
Chrysiogenetes
Deferribacteres
Dictyoglomi
Elusimicrobia
Fusobacteria
Gemmatimonadetes
Nitrospirae
Synergistetes
G+/
no OM
Firmicutes
(BV3)
Bacilli
Bacillales
Lactobacillales
Clostridia
Clostridiales
Halanaerobiales
Thermoanaerobacteriales
Natranaerobiales
Erysipelotrichia
Erysipelotrichiales
Thermolithobacteria
Thermolithobacterales
Tenericutes/
Mollicutes
Mycoplasmatales
Entomoplasmatales
Anaeroplasmatales
Acholeplasmatales
Haloplasmatales
Negativicutes
Selenomonadales
Actinobacteria
(BV5)
Actinobacteria
Actinomycetales
Bifidobacteriales
Acidimicrobiia
Acidimicrobiales
Coriobacteriidae
Coriobacteriales
Nitriliruptoria
Euzebyales
Nitriliruptorales
Rubrobacteria
Gaiellales
Rubrobacterales
Thermoleophilales
Solirubrobacterales
Incertae
sedis
†Archaeosphaeroides
†Eobacterium
†Leptotrichites
Source: Bergey's Manual (2001–2012). Alternative views: Wikispecies.
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Microbiology: Bacteria
Medical
microbiology
infection
Coley's toxins
Exotoxin
Lysogenic cycle
Pathogenic bacteria
resistance
Biochemistry
and ecology
Oxygen
preference
Aerobic
Obligate
Anaerobic
Facultative
Obligate
Microaerophile
Nanaerobe
Aerotolerant
Other
Extremophile
Human flora
Gut
Lung
Mouth
Skin
Vaginal (In pregnancy)
Placental
Uterine
Salivary
Microbial metabolism
Nitrogen fixation
Microbial ecology
Primary nutritional groups
Substrate preference
Lipophilic
Saccharophilic
Shape
Bacterial cellular morphologies
Coccus
Diplococcus
Bacillus
Coccobacillus
Spiral
Structure
Cell
envelope
Cell membrane
Cell wall: Peptidoglycan
NAM
NAG
DAP
Gram-positive bacteria only: Teichoic acid
Lipoteichoic acid
Endospore
Gram-negative bacteria only: Bacterial outer membrane
Porin
Lipopolysaccharide
Periplasmic space
Mycobacteria only: Arabinogalactan
Mycolic acid
Outside
envelope
Bacterial capsule
Slime layer
S-layer
Glycocalyx
Pilus
Fimbria
Non-motile bacteria
Composite
Biofilm
Taxonomy
Bacteria (classifications)
Bacterial phyla
Former groupings: Schizomycetes
Monera
Prokaryota
Gracilicutes
Firmicutes
Mollicutes
Mendosicutes
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Microorganisms
Microorganisms
Archaea
Bacterium
Cyanobacteria
Fungi
Nanobacterium
Prokaryote
Protist
Protozoa
Virus
Microbiology
Microbial biogeography
Microbial genetics
Microbial intelligence
Microbial metabolism
Microbial phylogenetics
Microbial population biology
Mycology
Virology
Ecology
Microbial cooperation
Microbial ecology
Microbial food web
Microbial loop
Microbial synergy
Host microbe interactions in Caenorhabditis elegans
Techniques
Dark field microscopy
Impedance microbiology
Microbial cytology
Microbiological culture
Staining
Marine
Marine microorganism
Antarctic
Marine microbial symbiosis
International Census of Marine Microbes
Human related
Microbes in human culture
Food microbiology
Microbial oil
Microbial symbiosis and immunity
Nylon-eating
Human microbiota
asthma
dysbiosis
fecal
Human Microbiome Project
gut
lung
mouth
skin
vagina
in pregnancy
placenta
uterus
Protein production
Other
Lines on the Antiquity of Microbes
Microbially induced sedimentary structure
Microbial dark matter
Physical factors affecting microbial life
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Antonie van Leeuwenhoek
Microscopic organisms first observed and
described by Antonie van Leeuwenhoek 1
Bacteria
Ciliate
Giardia
Infusoria
Protist
Protozoa
Rotifer
Spermatozoon
Volvox
General topics
Animalcule
Bacteriology
Protozoology
Protistology
Spontaneous generation
Preformationism
Microscopic discovery of microorganisms
Microscopic scale
History of biology
History of microbiology
Natural history
Optical microscopy
History of microscopy
History of the microscope
Invention of the optical microscope
Timeline of microscope technology
Golden Age of Dutch science and technology
Science and technology in the Dutch Republic
Royal Society
Scientific Revolution
Age of Reason
Related people
Clifford Dobell (Leeuwenhoek scholar)
Brian J. Ford (Leeuwenhoek scholar)
Galileo Galilei
Regnier de Graaf
Robert Hooke (author of Micrographia)
Nicolaas Hartsoeker
Nicolas Steno
Jan Swammerdam
Johannes Vermeer
Recognitions
Antoni van Leeuwenhoek Ziekenhuis
Leeuwenhoek (crater)
Leeuwenhoek Lecture
Leeuwenhoek Medal
Leeuwenhoeckia
Levenhookia
Leeuwenhoekiella
List of people considered father or mother of a scientific field
1 Generally considered to be the first acknowledged microscopist and microbiologist in history.
Taxon identifiers
Wd: Q10876
EoL: 288
GBIF: 3
ITIS: 50
NCBI: 2
Authority control
LCCN: sh85010813
GND: 4004296-0
NDL: 00570000
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https://ace.wikipedia.org/wiki/Bakteri
Bakteri – Achinese
ace
ace
interlanguage-link-target

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https://af.wikipedia.org/wiki/Bakterie
Bakterie – Afrikaans
af
af
interlanguage-link-target

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values:
https://ast.wikipedia.org/wiki/Bacteria
Bacteria – Asturian
ast
ast
interlanguage-link-target

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https://bs.wikipedia.org/wiki/Bakterija
Bakterija – Bosnian
bs
bs
interlanguage-link-target

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Bakteri – Breton
br
br
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https://ceb.wikipedia.org/wiki/Bacteria
Bacteria – Cebuano
ceb
ceb
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https://cs.wikipedia.org/wiki/Bakterie
Bakterie – Czech
cs
cs
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https://cy.wikipedia.org/wiki/Bacteria
Bacteria – Welsh
cy
cy
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https://da.wikipedia.org/wiki/Bakterier
Bakterier – Danish
da
da
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values:
https://de.wikipedia.org/wiki/Bakterien
Bakterien – German
de
de
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values: ބެކްޓީރިޔާ – Divehi

values:
https://et.wikipedia.org/wiki/Bakterid
Bakterid – Estonian
et
et
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https://es.wikipedia.org/wiki/Bacteria
Bacteria – Spanish
es
es
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https://eo.wikipedia.org/wiki/Bakterioj
Bakterioj – Esperanto
eo
eo
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Bakterio – Basque
eu
eu
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https://hif.wikipedia.org/wiki/Bacteria
Bacteria – Fiji Hindi
hif
hif
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https://fo.wikipedia.org/wiki/Bakteria
Bakteria – Faroese
fo
fo
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https://fy.wikipedia.org/wiki/Baktearje
Baktearje – Western Frisian
fy
fy
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https://gv.wikipedia.org/wiki/Bacteria
Bacteria – Manx
gv
gv
interlanguage-link-target

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https://gl.wikipedia.org/wiki/Bacterias
Bacterias – Galician
gl
gl
interlanguage-link-target

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https://ki.wikipedia.org/wiki/Bacteria
Bacteria – Kikuyu
ki
ki
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Bakterije – Croatian
hr
hr
interlanguage-link-target

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Bakterio – Ido
io
io
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Bakteria – Iloko
ilo
ilo
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Bakteri – Indonesian
id
id
interlanguage-link-target

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https://ia.wikipedia.org/wiki/Bacterio
Bacterio – Interlingua
ia
ia
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https://is.wikipedia.org/wiki/Gerlar
Gerlar – Icelandic
is
is
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https://it.wikipedia.org/wiki/Bacteria
Bacteria – Italian
it
it
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Bakteria – Cornish
kw
kw
interlanguage-link-target

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Bagiteri – Kinyarwanda
rw
rw
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Bakteria – Swahili
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sw
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Bakteri – Haitian Creole
ht
ht
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Bacterium – Latin
la
la
interlanguage-link-target

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Bakterien – Luxembourgish
lb
lb
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Bacteria – Ligurian
lij
lij
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Bakteria – Malay
ms
ms
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Bacteria – Neapolitan
nap
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Bakterier – Norwegian
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Bakteriar – Norwegian Nynorsk
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Bakteria – Papiamento
pap
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Bakterie – Polish
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Bacterie – Romanian
ro
ro
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Bacteria – Scots
sco
sco
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Bakteret – Albanian
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sq
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Batteriu – Sicilian
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scn
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Bakterije – Slovenian
sl
sl
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Bakteriya – Somali
so
so
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Bakterija – Serbo-Croatian
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sh
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Bakteerit – Finnish
fi
fi
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Bakterier – Swedish
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sv
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Bakterya – Tagalog
tl
tl
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Bakteri – Turkish
tr
tr
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Sigin – Zhuang
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za
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Bateri – Venetian
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vec
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Bactereye – Walloon
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wa
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Bakterya – Waray
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war
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