organism An organism is any life, living thing that functions as an individual. Such a definition raises more problems than it solves, not least because the concept of an individual is also difficult. Many criteria, few of them widely accepted, have be ...

microscopic The microscopic scale () is the scale of objects and events smaller than those that can easily be seen by the naked eye, requiring a lens or microscope to see them clearly. In physics, the microscopic scale is sometimes regarded as the scale betwe ...

single-celled A unicellular organism, also known as a single-celled organism, is an organism that consists of a single cell (biology), cell, unlike a multicellular organism that consists of multiple cells. Organisms fall into two general categories: prokaryotic ...

Jain literature Jain literature () refers to the literature of the Jain religion. It is a vast and ancient literary tradition, which was initially transmitted orally. The oldest surviving material is contained in the canonical ''Jain Agamas'', which are wri ...

microscope A microscope () is a laboratory equipment, laboratory instrument used to examine objects that are too small to be seen by the naked eye. Microscopy is the science of investigating small objects and structures using a microscope. Microscopic ...

Anton van Leeuwenhoek Antonie Philips van Leeuwenhoek ( ; ; 24 October 1632 – 26 August 1723) was a Dutch microbiologist and microscopist in the Golden Age of Dutch art, science and technology. A largely self-taught man in science, he is commonly known as " ...

Louis Pasteur Louis Pasteur (, ; 27 December 1822 – 28 September 1895) was a French chemist, pharmacist, and microbiologist renowned for his discoveries of the principles of vaccination, Fermentation, microbial fermentation, and pasteurization, the la ...

food spoilage Food spoilage is the process whereby food becomes unsuitable to ingest by a person; it is a matter of food safety. Bacteria and various fungi are the causes of spoilage, and can create serious consequences for consumers, but there are preventive ...

spontaneous generation Spontaneous generation is a superseded scientific theory that held that living creatures could arise from non-living matter and that such processes were commonplace and regular. It was hypothesized that certain forms, such as fleas, could ...

Robert Koch Heinrich Hermann Robert Koch ( ; ; 11 December 1843 – 27 May 1910) was a German physician and microbiologist. As the discoverer of the specific causative agents of deadly infectious diseases including tuberculosis, cholera and anthrax, he i ...

tuberculosis Tuberculosis (TB), also known colloquially as the "white death", or historically as consumption, is a contagious disease usually caused by ''Mycobacterium tuberculosis'' (MTB) bacteria. Tuberculosis generally affects the lungs, but it can al ...

cholera Cholera () is an infection of the small intestine by some Strain (biology), strains of the Bacteria, bacterium ''Vibrio cholerae''. Symptoms may range from none, to mild, to severe. The classic symptom is large amounts of watery diarrhea last ...

diphtheria Diphtheria is an infection caused by the bacteria, bacterium ''Corynebacterium diphtheriae''. Most infections are asymptomatic or have a mild Course (medicine), clinical course, but in some outbreaks, the mortality rate approaches 10%. Signs a ...

anthrax Anthrax is an infection caused by the bacterium '' Bacillus anthracis'' or ''Bacillus cereus'' biovar ''anthracis''. Infection typically occurs by contact with the skin, inhalation, or intestinal absorption. Symptom onset occurs between one ...

unicellular organism A unicellular organism, also known as a single-celled organism, is an organism that consists of a single cell, unlike a multicellular organism that consists of multiple cells. Organisms fall into two general categories: prokaryotic organisms and ...

three domains of life The three-domain system is a taxonomic classification system that groups all cellular life into three domains, namely Archaea, Bacteria and Eukarya, introduced by Carl Woese, Otto Kandler and Mark Wheelis in 1990. The key difference from earli ...

Archaea Archaea ( ) is a Domain (biology), domain of organisms. Traditionally, Archaea only included its Prokaryote, prokaryotic members, but this has since been found to be paraphyletic, as eukaryotes are known to have evolved from archaea. Even thou ...

Bacteria Bacteria (; : bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of Prokaryote, prokaryotic microorganisms. Typically a few micr ...

Eukaryota The eukaryotes ( ) constitute the Domain (biology), domain of Eukaryota or Eukarya, organisms whose Cell (biology), cells have a membrane-bound cell nucleus, nucleus. All animals, plants, Fungus, fungi, seaweeds, and many unicellular organisms ...

multicellular organism A multicellular organism is an organism that consists of more than one cell (biology), cell, unlike unicellular organisms. All species of animals, Embryophyte, land plants and most fungi are multicellular, as are many algae, whereas a few organism ...

protist A protist ( ) or protoctist is any eukaryotic organism that is not an animal, land plant, or fungus. Protists do not form a natural group, or clade, but are a paraphyletic grouping of all descendants of the last eukaryotic common ancest ...

protozoa Protozoa (: protozoan or protozoon; alternative plural: protozoans) are a polyphyletic group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic debris. Historically ...

animals Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia (). With few exceptions, animals consume organic material, breathe oxygen, have myocytes and are able to move, can reproduce sexually, and grow from a ...

green plants Viridiplantae (; Kingdom (biology), kingdom Plantae ''Sensu#Common qualifiers, sensu stricto'') is a clade of around 450,000–500,000 species of eukaryote, eukaryotic organisms, most of which obtain their energy by photosynthesis. The green pla ...

micro-animal Microfauna ( and ) are microscopic animals and organisms that exhibit animal-like qualities and have body sizes that are usually <0.1 mm. Microfauna are represented in the animal kingdom (e.g.

s, some

fungi A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...

, and some

algae Algae ( , ; : alga ) is an informal term for any organisms of a large and diverse group of photosynthesis, photosynthetic organisms that are not plants, and includes species from multiple distinct clades. Such organisms range from unicellular ...

. Microorganisms can have very different

habitat In ecology, habitat refers to the array of resources, biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species' habitat can be seen as the physical manifestation of its ...

s, and live everywhere from the

poles Pole or poles may refer to: People *Poles (people), another term for Polish people, from the country of Poland * Pole (surname), including a list of people with the name * Pole (musician) (Stefan Betke, born 1967), German electronic music artist ...

to the

equator The equator is the circle of latitude that divides Earth into the Northern Hemisphere, Northern and Southern Hemisphere, Southern Hemispheres of Earth, hemispheres. It is an imaginary line located at 0 degrees latitude, about in circumferen ...

, in

desert A desert is a landscape where little precipitation occurs and, consequently, living conditions create unique biomes and ecosystems. The lack of vegetation exposes the unprotected surface of the ground to denudation. About one-third of the la ...

s,

geyser A geyser (, ) is a spring with an intermittent water discharge ejected turbulently and accompanied by steam. The formation of geysers is fairly rare and is caused by particular hydrogeological conditions that exist only in a few places on Ea ...

s, rocks, and the

deep sea The deep sea is broadly defined as the ocean depth where light begins to fade, at an approximate depth of or the point of transition from continental shelves to continental slopes. Conditions within the deep sea are a combination of low tempe ...

. Some are adapted to extremes such as very hot or very cold conditions, others to

high pressure In science and engineering the study of high pressure examines its effects on materials and the design and construction of devices, such as a diamond anvil cell, which can create high pressure. ''High pressure'' usually means pressures of thousan ...

, and a few, such as '' Deinococcus radiodurans'', to high radiation environments. Microorganisms also make up the

microbiota Microbiota are the range of microorganisms that may be commensal, mutualistic, or pathogenic found in and on all multicellular organisms, including plants. Microbiota include bacteria, archaea, protists, fungi, and viruses, and have been found ...

found in and on all

multicellular organisms A multicellular organism is an organism that consists of more than one cell, unlike unicellular organisms. All species of animals, land plants and most fungi are multicellular, as are many algae, whereas a few organisms are partially uni- and pa ...

. There is evidence that 3.45-billion-year-old Australian rocks once contained microorganisms, the earliest direct evidence of life on Earth. Microbes are important in human culture and

health Health has a variety of definitions, which have been used for different purposes over time. In general, it refers to physical and emotional well-being, especially that associated with normal functioning of the human body, absent of disease, p ...

in many ways, serving to ferment foods and treat sewage, and to produce fuel,

enzyme An enzyme () is a protein that acts as a biological catalyst by accelerating chemical reactions. The molecules upon which enzymes may act are called substrate (chemistry), substrates, and the enzyme converts the substrates into different mol ...

s, and other

bioactive compound A bioactive compound is a compound that has an effect on a living organism, tissue or cell, usually demonstrated by basic research in vitro or in vivo in the laboratory. While dietary nutrients are essential to life, bioactive compounds have not ...

s. Microbes are essential tools in

biology Biology is the scientific study of life and living organisms. It is a broad natural science that encompasses a wide range of fields and unifying principles that explain the structure, function, growth, History of life, origin, evolution, and ...

as

model organism A model organism is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the workings of other organisms. Mo ...

s and have been put to use in

biological warfare Biological warfare, also known as germ warfare, is the use of biological toxins or Pathogen, infectious agents such as bacteria, viruses, insects, and Fungus, fungi with the intent to kill, harm or incapacitate humans, animals or plants as an ...

and

bioterrorism Bioterrorism is terrorism involving the intentional release or dissemination of biological agents. These agents include bacteria, viruses, insects, fungi, and/or their toxins, and may be in a naturally occurring or a human-modified form, in mu ...

. Microbes are a vital component of fertile soil. In the

human body The human body is the entire structure of a Human, human being. It is composed of many different types of Cell (biology), cells that together create Tissue (biology), tissues and subsequently Organ (biology), organs and then Organ system, org ...

, microorganisms make up the

human microbiota Human microbiota are microorganisms (bacteria, viruses, fungi and archaea) found in a specific environment. They can be found in the stomach, intestines, skin, genitals and other parts of the body. Various body parts have diverse microorganisms. S ...

, including the essential

gut flora Gut microbiota, gut microbiome, or gut flora are the microorganisms, including bacteria, archaea, fungi, and viruses, that live in the digestive tracts of animals. The gastrointestinal metagenome is the aggregate of all the genomes of the g ...

. The

pathogen In biology, a pathogen (, "suffering", "passion" and , "producer of"), in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a Germ theory of d ...

s responsible for many

infectious disease An infection is the invasion of tissue (biology), tissues by pathogens, their multiplication, and the reaction of host (biology), host tissues to the infectious agent and the toxins they produce. An infectious disease, also known as a transmis ...

s are microbes and, as such, are the target of hygiene measures.

Discovery

Ancient precursors

The possible existence of microscopic organisms was discussed for many centuries before their discovery in the 17th century. By the 6th century BC, the Jains of present-day India postulated the existence of tiny organisms called ''

nigoda In Jain cosmology, the Nigoda is a realm existing in which the lowest forms of invisible life reside in endless numbers, and without any hope of release by self-effort. Jain scriptures describe nigodas which are microorganisms Kanti V. Mardia, A ...

s''. These nigodas are said to be born in clusters; they live everywhere, including the bodies of plants, animals, and people; and their life lasts only for a fraction of a second. According to

Mahavira Mahavira (Devanagari: महावीर, ), also known as Vardhamana (Devanagari: वर्धमान, ), was the 24th ''Tirthankara'' (Supreme Preacher and Ford Maker) of Jainism. Although the dates and most historical details of his lif ...

, the 24th preacher of Jainism, the humans destroy these nigodas on a massive scale, when they eat, breathe, sit, and move. Many modern Jains assert that Mahavira's teachings presage the existence of microorganisms as discovered by modern science. The earliest known idea to indicate the possibility of diseases spreading by yet unseen organisms was that of the Roman scholar

Marcus Terentius Varro Marcus Terentius Varro (116–27 BCE) was a Roman polymath and a prolific author. He is regarded as ancient Rome's greatest scholar, and was described by Petrarch as "the third great light of Rome" (after Virgil and Cicero). He is sometimes call ...

in a 1st-century BC book entitled ''On Agriculture'' in which he called the unseen creatures animalia minuta, and warns against locating a homestead near a swamp:''Varro on Agriculture'' 1, xii Loeb In ''

The Canon of Medicine ''The Canon of Medicine'' () is an encyclopedia of medicine in five books compiled by Avicenna (, ibn Sina) and completed in 1025. It is among the most influential works of its time. It presents an overview of the contemporary medical knowle ...

'' (1020),

Avicenna Ibn Sina ( – 22 June 1037), commonly known in the West as Avicenna ( ), was a preeminent philosopher and physician of the Muslim world, flourishing during the Islamic Golden Age, serving in the courts of various Iranian peoples, Iranian ...

suggested that

tuberculosis Tuberculosis (TB), also known colloquially as the "white death", or historically as consumption, is a contagious disease usually caused by ''Mycobacterium tuberculosis'' (MTB) bacteria. Tuberculosis generally affects the lungs, but it can al ...

and other diseases might be contagious.

Early modern

Turkish scientist Akshamsaddin mentioned the microbe in his work (The Material of Life) about two centuries prior to Leeuwenhoek's experimental discovery: In 1546, Girolamo Fracastoro proposed that epidemic diseases were caused by transferable seedlike entities that could transmit infection by direct or indirect contact, or even without contact over long distances.

Antonie van Leeuwenhoek Antonie Philips van Leeuwenhoek ( ; ; 24 October 1632 – 26 August 1723) was a Dutch microbiologist and microscopist in the Golden Age of Dutch art, science and technology. A largely self-taught man in science, he is commonly known as " ...

is considered to be one of the fathers of microbiology. He was the first in 1673 to discover and conduct scientific experiments with microorganisms, using simple single-lensed

microscopes A microscope () is a laboratory instrument used to examine objects that are too small to be seen by the naked eye. Microscopy is the science of investigating small objects and structures using a microscope. Microscopic means being invisibl ...

of his own design.

Robert Hooke Robert Hooke (; 18 July 16353 March 1703) was an English polymath who was active as a physicist ("natural philosopher"), astronomer, geologist, meteorologist, and architect. He is credited as one of the first scientists to investigate living ...

, a contemporary of Leeuwenhoek, also used

microscopy Microscopy is the technical field of using microscopes to view subjects too small to be seen with the naked eye (objects that are not within the resolution range of the normal eye). There are three well-known branches of microscopy: optical mic ...

to observe microbial life in the form of the fruiting bodies of

mould A mold () or mould () is one of the structures that certain fungi can form. The dust-like, colored appearance of molds is due to the formation of spores containing fungal secondary metabolites. The spores are the dispersal units of the fungi ...

s. In his

1665 Events January–March * January 5 – The ''Journal des sçavans'' begins publication of the first scientific journal in France. * February 15 – Molière's comedy '' Dom Juan ou le Festin de pierre'', based on the Spanis ...

book ''

Micrographia ''Micrographia: or Some Physiological Descriptions of Minute Bodies Made by Magnifying Glasses. With Observations and Inquiries Thereupon'' is a historically significant book by Robert Hooke about his observations through various lenses. It wa ...

'', he made drawings of studies, and he coined the term '' cell''.

19th century

Louis Pasteur Louis Pasteur (, ; 27 December 1822 – 28 September 1895) was a French chemist, pharmacist, and microbiologist renowned for his discoveries of the principles of vaccination, Fermentation, microbial fermentation, and pasteurization, the la ...

(1822–1895) exposed boiled broths to the air, in vessels that contained a filter to prevent particles from passing through to the

growth medium A growth medium or culture medium is a solid, liquid, or semi-solid designed to support the growth of a population of microorganisms or cells via the process of cell proliferation or small plants like the moss ''Physcomitrella patens''. Differe ...

, and also in vessels without a filter, but with air allowed in via a curved tube so dust particles would settle and not come in contact with the broth. By boiling the broth beforehand, Pasteur ensured that no microorganisms survived within the broths at the beginning of his experiment. Nothing grew in the broths in the course of Pasteur's experiment. This meant that the living organisms that grew in such broths came from outside, as

spore In biology, a spore is a unit of sexual reproduction, sexual (in fungi) or asexual reproduction that may be adapted for biological dispersal, dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores fo ...

s on dust, rather than spontaneously generated within the broth. Thus, Pasteur refuted the theory of

spontaneous generation Spontaneous generation is a superseded scientific theory that held that living creatures could arise from non-living matter and that such processes were commonplace and regular. It was hypothesized that certain forms, such as fleas, could ...

and supported the

germ theory of disease The germ theory of disease is the currently accepted scientific theory for many diseases. It states that microorganisms known as pathogens or "germs" can cause disease. These small organisms, which are too small to be seen without magnification, ...

.

In 1876,

Robert Koch Heinrich Hermann Robert Koch ( ; ; 11 December 1843 – 27 May 1910) was a German physician and microbiologist. As the discoverer of the specific causative agents of deadly infectious diseases including tuberculosis, cholera and anthrax, he i ...

(1843–1910) established that microorganisms can cause disease. He found that the blood of cattle that were infected with

anthrax Anthrax is an infection caused by the bacterium '' Bacillus anthracis'' or ''Bacillus cereus'' biovar ''anthracis''. Infection typically occurs by contact with the skin, inhalation, or intestinal absorption. Symptom onset occurs between one ...

always had large numbers of ''

Bacillus anthracis ''Bacillus anthracis'' is a gram-positive and rod-shaped bacterium that causes anthrax, a deadly disease to livestock and, occasionally, to humans. It is the only permanent (obligate) pathogen within the genus ''Bacillus''. Its infection is a ty ...

''. Koch found that he could transmit anthrax from one animal to another by taking a small sample of blood from the infected animal and injecting it into a healthy one, and this caused the healthy animal to become sick. He also found that he could grow the bacteria in a nutrient broth, then inject it into a healthy animal, and cause illness. Based on these experiments, he devised criteria for establishing a causal link between a microorganism and a disease and these are now known as

Koch's postulates Koch's postulates ( ) are four criteria designed to establish a causality, causal relationship between a microbe and a disease. The postulates were formulated by Robert Koch and Friedrich Loeffler in 1884, based on earlier concepts described by ...

. Although these postulates cannot be applied in all cases, they do retain historical importance to the development of scientific thought and are still being used today. The discovery of microorganisms such as ''

Euglena ''Euglena'' is a genus of Unicellular organism, single-celled, flagellate eukaryotes. It is the best-known and most widely studied member of the class Euglenoidea, a diverse group containing some 54 genera and at least 200 species. Species of '' ...

'' that did not fit into either the

animal Animals are multicellular, eukaryotic organisms in the Biology, biological Kingdom (biology), kingdom Animalia (). With few exceptions, animals heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, ...

or

plant Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they are predominantly Photosynthesis, photosynthetic. This means that they obtain their energy from sunlight, using chloroplasts derived from endosymbiosis with c ...

kingdoms, since they were

photosynthetic Photosynthesis ( ) is a Biological system, system of biological processes by which Photoautotrophism, photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical ener ...

like plants, but

motile Motility is the ability of an organism to move independently using metabolic energy. This biological concept encompasses movement at various levels, from whole organisms to cells and subcellular components. Motility is observed in animals, mi ...

like animals, led to the naming of a third kingdom in the 1860s. In 1860 John Hogg called this the Protoctista, and in 1866

Ernst Haeckel Ernst Heinrich Philipp August Haeckel (; ; 16 February 1834 – 9 August 1919) was a German zoologist, natural history, naturalist, eugenics, eugenicist, Philosophy, philosopher, physician, professor, marine biology, marine biologist and artist ...

named it the

Protista A protist ( ) or protoctist is any Eukaryote, eukaryotic organism that is not an animal, Embryophyte, land plant, or fungus. Protists do not form a Clade, natural group, or clade, but are a Paraphyly, paraphyletic grouping of all descendants o ...

. The work of Pasteur and Koch did not accurately reflect the true diversity of the microbial world because of their exclusive focus on microorganisms having direct medical relevance. It was not until the work of

Martinus Beijerinck Martinus Willem Beijerinck (, 16 March 1851 – 1 January 1931) was a Dutch microbiologist and botanist who was one of the founders of virology and environmental microbiology. He is credited with the co-discovery of viruses A virus i ...

and

Sergei Winogradsky Sergei Nikolaevich Winogradsky (; ; , Kyiv – 24 February 1953, Brie-Comte-Robert), also published under the name Sergius Winogradsky, was a Ukrainian and Russian microbiologist, ecologist and soil science, soil scientist who pioneered the Biog ...

in the late 19th century that the true breadth of microbiology was revealed. Beijerinck made two major contributions to microbiology: the discovery of

virus A virus is a submicroscopic infectious agent that replicates only inside the living Cell (biology), cells of an organism. Viruses infect all life forms, from animals and plants to microorganisms, including bacteria and archaea. Viruses are ...

es and the development of enrichment culture techniques. While his work on the

tobacco mosaic virus Tobacco mosaic virus (TMV) is a positive-sense single-stranded RNA virus species in the genus '' Tobamovirus'' that infects a wide range of plants, especially tobacco and other members of the family Solanaceae. The infection causes characteris ...

established the basic principles of virology, it was his development of enrichment culturing that had the most immediate impact on microbiology by allowing for the cultivation of a wide range of microbes with wildly different physiologies. Winogradsky was the first to develop the concept of chemolithotrophy and to thereby reveal the essential role played by microorganisms in geochemical processes. He was responsible for the first isolation and description of both nitrifying and nitrogen-fixing bacteria. French-Canadian microbiologist Félix d'Hérelle co-discovered

bacteriophage A bacteriophage (), also known informally as a phage (), is a virus that infects and replicates within bacteria. The term is derived . Bacteriophages are composed of proteins that Capsid, encapsulate a DNA or RNA genome, and may have structu ...

s and was one of the earliest applied microbiologists.

Classification and structure

Microorganisms can be found almost anywhere on

Earth Earth is the third planet from the Sun and the only astronomical object known to Planetary habitability, harbor life. This is enabled by Earth being an ocean world, the only one in the Solar System sustaining liquid surface water. Almost all ...

.

Bacteria Bacteria (; : bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of Prokaryote, prokaryotic microorganisms. Typically a few micr ...

and

archaea Archaea ( ) is a Domain (biology), domain of organisms. Traditionally, Archaea only included its Prokaryote, prokaryotic members, but this has since been found to be paraphyletic, as eukaryotes are known to have evolved from archaea. Even thou ...

are almost always microscopic, while a number of

eukaryote The eukaryotes ( ) constitute the Domain (biology), domain of Eukaryota or Eukarya, organisms whose Cell (biology), cells have a membrane-bound cell nucleus, nucleus. All animals, plants, Fungus, fungi, seaweeds, and many unicellular organisms ...

s are also microscopic, including most

protists A protist ( ) or protoctist is any Eukaryote, eukaryotic organism that is not an animal, Embryophyte, land plant, or fungus. Protists do not form a Clade, natural group, or clade, but are a Paraphyly, paraphyletic grouping of all descendants o ...

, some

fungi A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...

, as well as some

micro-animal Microfauna ( and ) are microscopic animals and organisms that exhibit animal-like qualities and have body sizes that are usually <0.1 mm. Microfauna are represented in the animal kingdom (e.g.

s and plants.

Virus A virus is a submicroscopic infectious agent that replicates only inside the living Cell (biology), cells of an organism. Viruses infect all life forms, from animals and plants to microorganisms, including bacteria and archaea. Viruses are ...

es are generally regarded as not living and therefore not considered to be microorganisms, although a subfield of

microbiology Microbiology () is the branches of science, scientific study of microorganisms, those being of unicellular organism, unicellular (single-celled), multicellular organism, multicellular (consisting of complex cells), or non-cellular life, acellula ...

is

virology Virology is the Scientific method, scientific study of biological viruses. It is a subfield of microbiology that focuses on their detection, structure, classification and evolution, their methods of infection and exploitation of host (biology), ...

, the study of viruses.

Evolution

Single-celled microorganisms were the first forms of life to develop on Earth, approximately 3.5 billion years ago. Further evolution was slow, and for about 3 billion years in the

Precambrian The Precambrian ( ; or pre-Cambrian, sometimes abbreviated pC, or Cryptozoic) is the earliest part of Earth's history, set before the current Phanerozoic Eon. The Precambrian is so named because it preceded the Cambrian, the first period of t ...

eon, (much of the history of life on Earth), all

organism An organism is any life, living thing that functions as an individual. Such a definition raises more problems than it solves, not least because the concept of an individual is also difficult. Many criteria, few of them widely accepted, have be ...

s were microorganisms. Bacteria, algae and fungi have been identified in

amber Amber is fossilized tree resin. Examples of it have been appreciated for its color and natural beauty since the Neolithic times, and worked as a gemstone since antiquity."Amber" (2004). In Maxine N. Lurie and Marc Mappen (eds.) ''Encyclopedia ...

that is 220 million years old, which shows that the

morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...

of microorganisms has changed little since at least the

Triassic The Triassic ( ; sometimes symbolized 🝈) is a geologic period and system which spans 50.5 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.4 Mya. The Triassic is t ...

period. The newly discovered biological role played by nickel, however – especially that brought about by

volcanic eruptions A volcanic eruption occurs when material is expelled from a volcanic vent or fissure. Several types of volcanic eruptions have been distinguished by volcanologists. These are often named after famous volcanoes where that type of behavior h ...

from the

Siberian Traps The Siberian Traps () are a large region of volcanic rock, known as a large igneous province, in Siberia, Russia. The massive eruptive event that formed the trap rock, traps is one of the largest known Volcano, volcanic events in the last years ...

– may have accelerated the evolution of

methanogen Methanogens are anaerobic archaea that produce methane as a byproduct of their energy metabolism, i.e., catabolism. Methane production, or methanogenesis, is the only biochemical pathway for Adenosine triphosphate, ATP generation in methanogens. A ...

s towards the end of the

Permian–Triassic extinction event The Permian–Triassic extinction event (also known as the P–T extinction event, the Late Permian extinction event, the Latest Permian extinction event, the End-Permian extinction event, and colloquially as the Great Dying,) was an extinction ...

. Microorganisms tend to have a relatively fast rate of evolution. Most microorganisms can reproduce rapidly, and bacteria are also able to freely exchange genes through

conjugation Conjugation or conjugate may refer to: Linguistics *Grammatical conjugation, the modification of a verb from its basic form *Emotive conjugation or Russell's conjugation, the use of loaded language Mathematics *Complex conjugation, the change o ...

, transformation and transduction, even between widely divergent species. This

horizontal gene transfer Horizontal gene transfer (HGT) or lateral gene transfer (LGT) is the movement of genetic material between organisms other than by the ("vertical") transmission of DNA from parent to offspring (reproduction). HGT is an important factor in the e ...

, coupled with a high

mutation In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, ...

rate and other means of transformation, allows microorganisms to swiftly evolve (via

natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the Heredity, heritable traits characteristic of a population over generation ...

) to survive in new environments and respond to environmental stresses. This rapid evolution is important in medicine, as it has led to the development of multidrug resistant

pathogenic bacteria Pathogenic bacteria are bacteria that can cause disease. This article focuses on the bacteria that are pathogenic to humans. Most species of bacteria are harmless and many are Probiotic, beneficial but others can cause infectious diseases. The nu ...

, ''superbugs'', that are resistant to antibiotics. A possible transitional form of microorganism between a prokaryote and a eukaryote was discovered in 2012 by Japanese scientists. ''

Parakaryon myojinensis ''Parakaryon myojinensis'', also known as the Myojin parakaryote, is a highly unusual species of single-celled organism known only from a single specimen, described in 2012. It has features of both prokaryotes and eukaryotes but is apparently ...

'' is a unique microorganism larger than a typical prokaryote, but with nuclear material enclosed in a membrane as in a eukaryote, and the presence of endosymbionts. This is seen to be the first plausible evolutionary form of microorganism, showing a stage of development from the prokaryote to the eukaryote.

Archaea

Archaea are

prokaryotic A prokaryote (; less commonly spelled procaryote) is a single-celled organism whose cell lacks a nucleus and other membrane-bound organelles. The word ''prokaryote'' comes from the Ancient Greek (), meaning 'before', and (), meaning 'nut' ...

unicellular organisms, and form the first domain of life in

Carl Woese Carl Richard Woese ( ; July 15, 1928 – December 30, 2012) was an American microbiologist and biophysicist. Woese is famous for defining the Archaea (a new domain of life) in 1977 through a pioneering phylogenetic taxonomy of 16S ribosomal ...

's

three-domain system The three-domain system is a taxonomic classification system that groups all cellular life into three domains, namely Archaea, Bacteria and Eukarya, introduced by Carl Woese, Otto Kandler and Mark Wheelis in 1990. The key difference from ea ...

. A prokaryote is defined as having no

cell nucleus The cell nucleus (; : nuclei) is a membrane-bound organelle found in eukaryote, eukaryotic cell (biology), cells. Eukaryotic cells usually have a single nucleus, but a few cell types, such as mammalian red blood cells, have #Anucleated_cells, ...

or other membrane bound-

organelle In cell biology, an organelle is a specialized subunit, usually within a cell (biology), cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts of cells, as Organ (anatomy), organs are to th ...

. Archaea share this defining feature with the bacteria with which they were once grouped. In 1990 the microbiologist Woese proposed the three-domain system that divided living things into bacteria, archaea and eukaryotes, and thereby split the prokaryote domain. Archaea differ from bacteria in both their genetics and biochemistry. For example, while bacterial

cell membrane The cell membrane (also known as the plasma membrane or cytoplasmic membrane, and historically referred to as the plasmalemma) is a biological membrane that separates and protects the interior of a cell from the outside environment (the extr ...

s are made from phosphoglycerides with

ester In chemistry, an ester is a compound derived from an acid (either organic or inorganic) in which the hydrogen atom (H) of at least one acidic hydroxyl group () of that acid is replaced by an organyl group (R). These compounds contain a distin ...

bonds, Achaean membranes are made of

ether lipid In biochemistry, an ether lipid refers to any lipid in which the lipid "tail" group is attached to the glycerol backbone via an ether, ether bond at any position. In contrast, conventional glycerophospholipids and triglycerides are triesters. St ...

s. Archaea were originally described as

extremophile An extremophile () is an organism that is able to live (or in some cases thrive) in extreme environments, i.e., environments with conditions approaching or stretching the limits of what known life can adapt to, such as extreme temperature, press ...

s living in extreme environments, such as

hot spring A hot spring, hydrothermal spring, or geothermal spring is a Spring (hydrology), spring produced by the emergence of Geothermal activity, geothermally heated groundwater onto the surface of the Earth. The groundwater is heated either by shallow ...

s, but have since been found in all types of

habitat In ecology, habitat refers to the array of resources, biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species' habitat can be seen as the physical manifestation of its ...

s. Only now are scientists beginning to realize how common archaea are in the environment, with

Thermoproteota The Thermoproteota are prokaryotes that have been classified as a phylum (biology), phylum of the domain Archaea. Initially, the Thermoproteota were thought to be sulfur-dependent extremophiles but recent studies have identified characteristic T ...

(formerly Crenarchaeota) being the most common form of life in the ocean, dominating ecosystems below in depth. These organisms are also common in soil and play a vital role in

ammonia Ammonia is an inorganic chemical compound of nitrogen and hydrogen with the chemical formula, formula . A Binary compounds of hydrogen, stable binary hydride and the simplest pnictogen hydride, ammonia is a colourless gas with a distinctive pu ...

oxidation. The combined domains of archaea and bacteria make up the most diverse and abundant group of

organism An organism is any life, living thing that functions as an individual. Such a definition raises more problems than it solves, not least because the concept of an individual is also difficult. Many criteria, few of them widely accepted, have be ...

s on Earth and inhabit practically all environments where the temperature is below +. They are found in

water Water is an inorganic compound with the chemical formula . It is a transparent, tasteless, odorless, and Color of water, nearly colorless chemical substance. It is the main constituent of Earth's hydrosphere and the fluids of all known liv ...

,

soil Soil, also commonly referred to as earth, is a mixture of organic matter, minerals, gases, water, and organisms that together support the life of plants and soil organisms. Some scientific definitions distinguish dirt from ''soil'' by re ...

,

air An atmosphere () is a layer of gases that envelop an astronomical object, held in place by the gravity of the object. A planet retains an atmosphere when the gravity is great and the temperature of the atmosphere is low. A stellar atmosph ...

, as the

microbiome A microbiome () is the community of microorganisms that can usually be found living together in any given habitat. It was defined more precisely in 1988 by Whipps ''et al.'' as "a characteristic microbial community occupying a reasonably wel ...

of an organism,

hot spring A hot spring, hydrothermal spring, or geothermal spring is a Spring (hydrology), spring produced by the emergence of Geothermal activity, geothermally heated groundwater onto the surface of the Earth. The groundwater is heated either by shallow ...

s and even deep beneath the Earth's crust in rocks. The number of prokaryotes is estimated to be around five nonillion, or 5 × 10³⁰, accounting for at least half the

biomass Biomass is a term used in several contexts: in the context of ecology it means living organisms, and in the context of bioenergy it means matter from recently living (but now dead) organisms. In the latter context, there are variations in how ...

on Earth. The biodiversity of the prokaryotes is unknown, but may be very large. A May 2016 estimate, based on laws of scaling from known numbers of species against the size of organism, gives an estimate of perhaps 1 trillion species on the planet, of which most would be microorganisms. Currently, only one-thousandth of one percent of that total have been described. Archael cells of some species aggregate and transfer

DNA Deoxyribonucleic acid (; DNA) is a polymer composed of two polynucleotide chains that coil around each other to form a double helix. The polymer carries genetic instructions for the development, functioning, growth and reproduction of al ...

from one cell to another through direct contact, particularly under stressful environmental conditions that cause

DNA damage DNA repair is a collection of processes by which a cell identifies and corrects damage to the DNA molecules that encode its genome. A weakened capacity for DNA repair is a risk factor for the development of cancer. DNA is constantly modified ...

.

Bacteria

Like archaea, bacteria are prokaryotic – unicellular, and having no cell nucleus or other membrane-bound organelle. Bacteria are microscopic, with a few extremely rare exceptions, such as ''

Thiomargarita namibiensis ''Thiomargarita namibiensis'' is a gram-negative, Facultative anaerobic organism, facultative anaerobic, coccus, coccoid bacterium found in South Africa's ocean sediments of the continental shelf of Namibia. The genus name ''Thiomargarita'' mean ...

''. Bacteria function and reproduce as individual cells, but they can often aggregate in multicellular

colonies A colony is a territory subject to a form of foreign rule, which rules the territory and its indigenous peoples separated from the foreign rulers, the colonizer, and their '' metropole'' (or "mother country"). This separated rule was often or ...

. Some species such as

myxobacteria The myxobacteria ("slime bacteria") are a group of bacteria that predominantly live in the soil and feed on insoluble organic substances. The myxobacteria have very large genomes relative to other bacteria, e.g. 9–10 million nucleotides except ...

can aggregate into complex swarming structures, operating as multicellular groups as part of their life cycle, or form clusters in bacterial colonies such as '' E. coli''. Their

genome A genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding genes, other functional regions of the genome such as ...

is usually a circular bacterial chromosome – a single loop of

DNA Deoxyribonucleic acid (; DNA) is a polymer composed of two polynucleotide chains that coil around each other to form a double helix. The polymer carries genetic instructions for the development, functioning, growth and reproduction of al ...

, although they can also harbor small pieces of DNA called

plasmid A plasmid is a small, extrachromosomal DNA molecule within a cell that is physically separated from chromosomal DNA and can replicate independently. They are most commonly found as small circular, double-stranded DNA molecules in bacteria and ...

s. These plasmids can be transferred between cells through

bacterial conjugation Bacterial conjugation is the transfer of genetic material between Bacteria, bacterial cells by direct cell-to-cell contact or by a bridge-like connection between two cells. This takes place through a pilus. It is a parasexual cycle, parasexual mode ...

. Bacteria have an enclosing

cell wall A cell wall is a structural layer that surrounds some Cell type, cell types, found immediately outside the cell membrane. It can be tough, flexible, and sometimes rigid. Primarily, it provides the cell with structural support, shape, protection, ...

, which provides strength and rigidity to their cells. They reproduce by

binary fission Binary may refer to: Science and technology Mathematics * Binary number, a representation of numbers using only two values (0 and 1) for each digit * Binary function, a function that takes two arguments * Binary operation, a mathematical o ...

or sometimes by

budding Budding or blastogenesis is a type of asexual reproduction in which a new organism develops from an outgrowth or bud due to cell division at one particular site. For example, the small bulb-like projection coming out from the yeast cell is kno ...

, but do not undergo

meiotic Meiosis () is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, the sperm or egg cells. It involves two rounds of division that ultimately result in four cells, each with only one copy ...

sexual reproduction Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote tha ...

. However, many bacterial species can transfer DNA between individual cells by a

horizontal gene transfer Horizontal gene transfer (HGT) or lateral gene transfer (LGT) is the movement of genetic material between organisms other than by the ("vertical") transmission of DNA from parent to offspring (reproduction). HGT is an important factor in the e ...

process referred to as natural transformation. Some species form extraordinarily resilient

spores In biology, a spore is a unit of sexual (in fungi) or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plant ...

, but for bacteria this is a mechanism for survival, not reproduction. Under optimal conditions bacteria can grow extremely rapidly and their numbers can double as quickly as every 20 minutes.

Eukaryotes

Most living things that are visible to the naked eye in their adult form are

eukaryote The eukaryotes ( ) constitute the Domain (biology), domain of Eukaryota or Eukarya, organisms whose Cell (biology), cells have a membrane-bound cell nucleus, nucleus. All animals, plants, Fungus, fungi, seaweeds, and many unicellular organisms ...

s, including

human Humans (''Homo sapiens'') or modern humans are the most common and widespread species of primate, and the last surviving species of the genus ''Homo''. They are Hominidae, great apes characterized by their Prehistory of nakedness and clothing ...

s. However, many eukaryotes are also microorganisms. Unlike

bacteria Bacteria (; : bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of Prokaryote, prokaryotic microorganisms. Typically a few micr ...

and

archaea Archaea ( ) is a Domain (biology), domain of organisms. Traditionally, Archaea only included its Prokaryote, prokaryotic members, but this has since been found to be paraphyletic, as eukaryotes are known to have evolved from archaea. Even thou ...

, eukaryotes contain

organelle In cell biology, an organelle is a specialized subunit, usually within a cell (biology), cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts of cells, as Organ (anatomy), organs are to th ...

s such as the

cell nucleus The cell nucleus (; : nuclei) is a membrane-bound organelle found in eukaryote, eukaryotic cell (biology), cells. Eukaryotic cells usually have a single nucleus, but a few cell types, such as mammalian red blood cells, have #Anucleated_cells, ...

, the

Golgi apparatus The Golgi apparatus (), also known as the Golgi complex, Golgi body, or simply the Golgi, is an organelle found in most eukaryotic Cell (biology), cells. Part of the endomembrane system in the cytoplasm, it protein targeting, packages proteins ...

and

mitochondria A mitochondrion () is an organelle found in the cells of most eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is us ...

in their cells. The nucleus is an organelle that houses the

DNA Deoxyribonucleic acid (; DNA) is a polymer composed of two polynucleotide chains that coil around each other to form a double helix. The polymer carries genetic instructions for the development, functioning, growth and reproduction of al ...

that makes up a cell's genome. DNA (Deoxyribonucleic acid) itself is arranged in complex

chromosome A chromosome is a package of DNA containing part or all of the genetic material of an organism. In most chromosomes, the very long thin DNA fibers are coated with nucleosome-forming packaging proteins; in eukaryotic cells, the most import ...

s.

Mitochondria A mitochondrion () is an organelle found in the cells of most eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is us ...

are organelles vital in

metabolism Metabolism (, from ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run cellular processes; the co ...

as they are the site of the

citric acid cycle The citric acid cycle—also known as the Krebs cycle, Szent–Györgyi–Krebs cycle, or TCA cycle (tricarboxylic acid cycle)—is a series of chemical reaction, biochemical reactions that release the energy stored in nutrients through acetyl-Co ...

and

oxidative phosphorylation Oxidative phosphorylation(UK , US : or electron transport-linked phosphorylation or terminal oxidation, is the metabolic pathway in which Cell (biology), cells use enzymes to Redox, oxidize nutrients, thereby releasing chemical energy in order ...

. They evolved from

symbiotic Symbiosis (Ancient Greek : living with, companionship < : together; and ''bíōsis'': living) is any type of a close and long-term biolo ...

bacteria and retain a remnant genome. Like bacteria,

plant cell Plant cells are the cells present in Viridiplantae, green plants, photosynthetic eukaryotes of the kingdom Plantae. Their distinctive features include primary cell walls containing cellulose, hemicelluloses and pectin, the presence of plastids ...

s have

cell wall A cell wall is a structural layer that surrounds some Cell type, cell types, found immediately outside the cell membrane. It can be tough, flexible, and sometimes rigid. Primarily, it provides the cell with structural support, shape, protection, ...

s, and contain organelles such as

chloroplast A chloroplast () is a type of membrane-bound organelle, organelle known as a plastid that conducts photosynthesis mostly in plant cell, plant and algae, algal cells. Chloroplasts have a high concentration of chlorophyll pigments which captur ...

s in addition to the organelles in other eukaryotes. Chloroplasts produce energy from light by

photosynthesis Photosynthesis ( ) is a system of biological processes by which photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical energy necessary to fuel their metabo ...

, and were also originally symbiotic bacteria. Unicellular eukaryotes consist of a single cell throughout their life cycle. This qualification is significant since most

multicellular A multicellular organism is an organism that consists of more than one cell (biology), cell, unlike unicellular organisms. All species of animals, Embryophyte, land plants and most fungi are multicellular, as are many algae, whereas a few organism ...

eukaryotes consist of a single cell called a

zygote A zygote (; , ) is a eukaryote, eukaryotic cell (biology), cell formed by a fertilization event between two gametes. The zygote's genome is a combination of the DNA in each gamete, and contains all of the genetic information of a new individ ...

only at the beginning of their life cycles. Microbial eukaryotes can be either

haploid Ploidy () is the number of complete sets of chromosomes in a cell (biology), cell, and hence the number of possible alleles for Autosome, autosomal and Pseudoautosomal region, pseudoautosomal genes. Here ''sets of chromosomes'' refers to the num ...

or

diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Here ''sets of chromosomes'' refers to the number of maternal and paternal chromosome copies, ...

, and some organisms have multiple

cell nuclei The cell nucleus (; : nuclei) is a membrane-bound organelle found in eukaryotic cells. Eukaryotic cells usually have a single nucleus, but a few cell types, such as mammalian red blood cells, have no nuclei, and a few others including osteoc ...

. Unicellular eukaryotes usually reproduce asexually by

mitosis Mitosis () is a part of the cell cycle in eukaryote, eukaryotic cells in which replicated chromosomes are separated into two new Cell nucleus, nuclei. Cell division by mitosis is an equational division which gives rise to genetically identic ...

under favorable conditions. However, under stressful conditions such as nutrient limitations and other conditions associated with DNA damage, they tend to reproduce sexually by

meiosis Meiosis () is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, the sperm or egg cells. It involves two rounds of division that ultimately result in four cells, each with only one c ...

and

syngamy Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a zygote and initiate its development into a new individual organism or of ...

.

Protists

Of

eukaryotic The eukaryotes ( ) constitute the Domain (biology), domain of Eukaryota or Eukarya, organisms whose Cell (biology), cells have a membrane-bound cell nucleus, nucleus. All animals, plants, Fungus, fungi, seaweeds, and many unicellular organisms ...

groups, the

protists A protist ( ) or protoctist is any Eukaryote, eukaryotic organism that is not an animal, Embryophyte, land plant, or fungus. Protists do not form a Clade, natural group, or clade, but are a Paraphyly, paraphyletic grouping of all descendants o ...

are most commonly

unicellular A unicellular organism, also known as a single-celled organism, is an organism that consists of a single cell, unlike a multicellular organism that consists of multiple cells. Organisms fall into two general categories: prokaryotic organisms and ...

and microscopic. This is a highly diverse group of organisms that are not easy to classify. Several

algae Algae ( , ; : alga ) is an informal term for any organisms of a large and diverse group of photosynthesis, photosynthetic organisms that are not plants, and includes species from multiple distinct clades. Such organisms range from unicellular ...

species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...

are

multicellular A multicellular organism is an organism that consists of more than one cell (biology), cell, unlike unicellular organisms. All species of animals, Embryophyte, land plants and most fungi are multicellular, as are many algae, whereas a few organism ...

protists, and

slime mold Slime mold or slime mould is an informal name given to a polyphyletic assemblage of unrelated eukaryotic organisms in the Stramenopiles, Rhizaria, Discoba, Amoebozoa and Holomycota clades. Most are near-microscopic; those in the Myxogastria ...

s have unique life cycles that involve switching between unicellular, colonial, and multicellular forms. The number of species of protists is unknown since only a small proportion has been identified. Protist diversity is high in oceans, deep sea-vents, river sediment and an acidic river, suggesting that many eukaryotic microbial communities may yet be discovered.

Fungi

The

fungi A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...

have several unicellular species, such as baker's yeast (''

Saccharomyces cerevisiae ''Saccharomyces cerevisiae'' () (brewer's yeast or baker's yeast) is a species of yeast (single-celled fungal microorganisms). The species has been instrumental in winemaking, baking, and brewing since ancient times. It is believed to have be ...

'') and fission yeast (''

Schizosaccharomyces pombe ''Schizosaccharomyces pombe'', also called "fission yeast", is a species of yeast used in traditional brewing and as a model organism in molecular and cell biology. It is a unicellular eukaryote, whose cells are rod-shaped. Cells typically meas ...

''). Some fungi, such as the pathogenic yeast ''

Candida albicans ''Candida albicans'' is an opportunistic pathogenic yeast that is a common member of the human gut flora. It can also survive outside the human body. It is detected in the gastrointestinal tract and mouth in 40–60% of healthy adults. It is usu ...

'', can undergo

phenotypic switching In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology (physical form and structure), its developmental processes, its biochemical and physiological propertie ...

and grow as single cells in some environments, and filamentous hyphae in others.

Plants

The

green algae The green algae (: green alga) are a group of chlorophyll-containing autotrophic eukaryotes consisting of the phylum Prasinodermophyta and its unnamed sister group that contains the Chlorophyta and Charophyta/ Streptophyta. The land plants ...

are a large group of photosynthetic eukaryotes that include many microscopic organisms. Although some green algae are classified as

protist A protist ( ) or protoctist is any eukaryotic organism that is not an animal, land plant, or fungus. Protists do not form a natural group, or clade, but are a paraphyletic grouping of all descendants of the last eukaryotic common ancest ...

s, others such as

charophyta Charophyta () is a paraphyletic group of freshwater green algae, called charophytes (), sometimes treated as a division, yet also as a superdivision. The terrestrial plants, the Embryophyta emerged deep within Charophyta, possibly from terres ...

are classified with

embryophyte The embryophytes () are a clade of plants, also known as Embryophyta (Plantae ''sensu strictissimo'') () or land plants. They are the most familiar group of photoautotrophs that make up the vegetation on Earth's dry lands and wetlands. Embryophy ...

plants, which are the most familiar group of land plants. Algae can grow as single cells, or in long chains of cells. The green algae include unicellular and colonial

flagellate A flagellate is a cell or organism with one or more whip-like appendages called flagella. The word ''flagellate'' also describes a particular construction (or level of organization) characteristic of many prokaryotes and eukaryotes and the ...

s, usually but not always with two

flagella A flagellum (; : flagella) (Latin for 'whip' or 'scourge') is a hair-like appendage that protrudes from certain plant and animal sperm cells, from fungal spores ( zoospores), and from a wide range of microorganisms to provide motility. Many pr ...

per cell, as well as various colonial, coccoid, and filamentous forms. In the

Charales Charales is an order of freshwater green algae in the division Charophyta, class Charophyceae, commonly known as stoneworts. Depending on the treatment of the genus '' Nitellopsis'', living (extant) species are placed into either one family ( ...

, which are the algae most closely related to higher plants, cells differentiate into several distinct tissues within the organism. There are about 6000 species of green algae.

Ecology

Microorganisms are found in almost every

habitat In ecology, habitat refers to the array of resources, biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species' habitat can be seen as the physical manifestation of its ...

present in nature, including hostile environments such as the North and South poles,

desert A desert is a landscape where little precipitation occurs and, consequently, living conditions create unique biomes and ecosystems. The lack of vegetation exposes the unprotected surface of the ground to denudation. About one-third of the la ...

s,

geyser A geyser (, ) is a spring with an intermittent water discharge ejected turbulently and accompanied by steam. The formation of geysers is fairly rare and is caused by particular hydrogeological conditions that exist only in a few places on Ea ...

s, and rocks. They also include all the

marine microorganism Marine microorganisms are defined by their habitat as microorganisms living in a marine habitat, marine environment, that is, in the saline water, saltwater of a sea or ocean or the brackish water of a coastal estuary. A microorganism (or mic ...

s of the

oceans The ocean is the body of salt water that covers approximately 70.8% of Earth. The ocean is conventionally divided into large bodies of water, which are also referred to as ''oceans'' (the Pacific, Atlantic, Indian, Antarctic/Southern, and ...

and

deep sea The deep sea is broadly defined as the ocean depth where light begins to fade, at an approximate depth of or the point of transition from continental shelves to continental slopes. Conditions within the deep sea are a combination of low tempe ...

. Some types of microorganisms have adapted to extreme environments and sustained colonies; these organisms are known as

extremophiles An extremophile () is an organism that is able to live (or in some cases thrive) in extreme environments, i.e., environments with conditions approaching or stretching the limits of what known life can adapt to, such as extreme temperature, pres ...

. Extremophiles have been isolated from rocks as much as 7 kilometres below the Earth's surface, and it has been suggested that the amount of organisms living below the Earth's surface is comparable with the amount of life on or above the surface. Extremophiles have been known to survive for a prolonged time in a

vacuum A vacuum (: vacuums or vacua) is space devoid of matter. The word is derived from the Latin adjective (neuter ) meaning "vacant" or "void". An approximation to such vacuum is a region with a gaseous pressure much less than atmospheric pressur ...

, and can be highly resistant to

radiation In physics, radiation is the emission or transmission of energy in the form of waves or particles through space or a material medium. This includes: * ''electromagnetic radiation'' consisting of photons, such as radio waves, microwaves, infr ...

, which may even allow them to survive in space. Many types of microorganisms have intimate symbiotic relationships with other larger organisms; some of which are mutually beneficial ( mutualism), while others can be damaging to the

host A host is a person responsible for guests at an event or for providing hospitality during it. Host may also refer to: Places * Host, Pennsylvania, a village in Berks County * Host Island, in the Wilhelm Archipelago, Antarctica People * ...

organism (

parasitism Parasitism is a close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The en ...

). If microorganisms can cause

disease A disease is a particular abnormal condition that adversely affects the structure or function (biology), function of all or part of an organism and is not immediately due to any external injury. Diseases are often known to be medical condi ...

in a host they are known as ''

pathogen In biology, a pathogen (, "suffering", "passion" and , "producer of"), in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a Germ theory of d ...

s''. Microorganisms play critical roles in Earth's

biogeochemical cycle A biogeochemical cycle, or more generally a cycle of matter, is the movement and transformation of chemical elements and compounds between living organisms, the atmosphere, and the Earth's crust. Major biogeochemical cycles include the carbon cyc ...

s as they are responsible for

decomposition Decomposition is the process by which dead organic substances are broken down into simpler organic or inorganic matter such as carbon dioxide, water, simple sugars and mineral salts. The process is a part of the nutrient cycle and is ess ...

and

nitrogen fixation Nitrogen fixation is a chemical process by which molecular dinitrogen () is converted into ammonia (). It occurs both biologically and abiological nitrogen fixation, abiologically in chemical industry, chemical industries. Biological nitrogen ...

. Bacteria use regulatory networks that allow them to adapt to almost every environmental niche on earth. A network of interactions among diverse types of molecules including DNA, RNA, proteins and metabolites, is utilised by the bacteria to achieve

regulation of gene expression Regulation of gene expression, or gene regulation, includes a wide range of mechanisms that are used by cells to increase or decrease the production of specific gene products (protein or RNA). Sophisticated programs of gene expression are wide ...

. In bacteria, the principal function of regulatory networks is to control the response to environmental changes, for example nutritional status and environmental stress. A complex organization of networks permits the microorganism to coordinate and integrate multiple environmental signals.

Extremophiles

upA tetrad of '' Deinococcus radiodurans'', a radioresistant

extremophile An extremophile () is an organism that is able to live (or in some cases thrive) in extreme environments, i.e., environments with conditions approaching or stretching the limits of what known life can adapt to, such as extreme temperature, press ...

bacterium Extremophiles are microorganisms that have adapted so that they can survive and even thrive in extreme environments that are normally fatal to most life-forms.

Thermophile A thermophile is a type of extremophile that thrives at relatively high temperatures, between . Many thermophiles are archaea, though some of them are bacteria and fungi. Thermophilic eubacteria are suggested to have been among the earliest bacte ...

s and

hyperthermophiles A hyperthermophile is an organism that thrives in extremely hot environments—from 60 °C (140 °F) upward. An optimal temperature for the existence of hyperthermophiles is often above 80 °C (176 °F). Hyperthermophiles are of ...

thrive in high

temperature Temperature is a physical quantity that quantitatively expresses the attribute of hotness or coldness. Temperature is measurement, measured with a thermometer. It reflects the average kinetic energy of the vibrating and colliding atoms making ...

s.

Psychrophile Psychrophiles or cryophiles (adj. ''psychrophilic'' or ''cryophilic'') are extremophile, extremophilic organisms that are capable of cell growth, growth and reproduction in low temperatures, ranging from to . They are found in places that are pe ...

s thrive in extremely low temperatures. – Temperatures as high as , as low as

Halophile A halophile (from the Greek word for 'salt-loving') is an extremophile that thrives in high salt In common usage, salt is a mineral composed primarily of sodium chloride (NaCl). When used in food, especially in granulated form, it is more ...

s such as ''

Halobacterium salinarum ''Halobacterium salinarum'', formerly known as ''Halobacterium cutirubrum'' or ''Halobacterium halobium'', is an extremely halophile, halophilic ocean, marine obligate aerobic archaeon. Despite its name, this is not a bacteria, bacterium, but a mem ...

'' (an archaean) thrive in high salt conditions, up to saturation.

Alkaliphile Alkaliphiles are a class of extremophilic microbes capable of survival in alkaline ( pH roughly 8.5–11) environments, growing optimally around a pH of 10. These bacteria can be further categorized as obligate alkaliphiles (those that require hig ...

s thrive in an

alkaline In chemistry, an alkali (; from the Arabic word , ) is a basic salt of an alkali metal or an alkaline earth metal. An alkali can also be defined as a base that dissolves in water. A solution of a soluble base has a pH greater than 7.0. The ...

pH of about 8.5–11.

Acidophile Acidophiles or acidophilic organisms are those that thrive under highly acidic conditions (usually at pH 5.0 or below). These organisms can be found in different branches of the Tree of life (biology), tree of life, including Archaea, Bacteria,Bec ...

s can thrive in a pH of 2.0 or less.

Piezophile A piezophile (from Greek "piezo-" for pressure and "-phile" for loving) is an organism with optimal growth under high hydrostatic pressure, i.e., an organism that has its maximum rate of growth at a hydrostatic pressure equal to or above , when tes ...

s thrive at very

high pressure In science and engineering the study of high pressure examines its effects on materials and the design and construction of devices, such as a diamond anvil cell, which can create high pressure. ''High pressure'' usually means pressures of thousan ...

s: up to 1,000–2,000 atm, down to 0 atm as in a

vacuum A vacuum (: vacuums or vacua) is space devoid of matter. The word is derived from the Latin adjective (neuter ) meaning "vacant" or "void". An approximation to such vacuum is a region with a gaseous pressure much less than atmospheric pressur ...

of

space Space is a three-dimensional continuum containing positions and directions. In classical physics, physical space is often conceived in three linear dimensions. Modern physicists usually consider it, with time, to be part of a boundless ...

. A few extremophiles such as '' Deinococcus radiodurans'' are radioresistant, resisting

radiation In physics, radiation is the emission or transmission of energy in the form of waves or particles through space or a material medium. This includes: * ''electromagnetic radiation'' consisting of photons, such as radio waves, microwaves, infr ...

exposure of up to 5k Gy. Extremophiles are significant in different ways. They extend terrestrial life into much of the Earth's

hydrosphere The hydrosphere () is the combined mass of water found on, under, and above the Planetary surface, surface of a planet, minor planet, or natural satellite. Although Earth's hydrosphere has been around for about 4 billion years, it continues to ch ...

, crust and

atmosphere An atmosphere () is a layer of gases that envelop an astronomical object, held in place by the gravity of the object. A planet retains an atmosphere when the gravity is great and the temperature of the atmosphere is low. A stellar atmosph ...

, their specific evolutionary adaptation mechanisms to their extreme environment can be exploited in

biotechnology Biotechnology is a multidisciplinary field that involves the integration of natural sciences and Engineering Science, engineering sciences in order to achieve the application of organisms and parts thereof for products and services. Specialists ...

, and their very existence under such extreme conditions increases the potential for

extraterrestrial life Extraterrestrial life, or alien life (colloquially, aliens), is life that originates from another world rather than on Earth. No extraterrestrial life has yet been scientifically conclusively detected. Such life might range from simple forms ...

.

Plants and soil

The

nitrogen cycle The nitrogen cycle is the biogeochemical cycle by which nitrogen is converted into multiple chemical forms as it circulates among atmosphere, atmospheric, terrestrial ecosystem, terrestrial, and marine ecosystems. The conversion of nitrogen can ...

in soils depends on the fixation of atmospheric nitrogen. This is achieved by a number of

diazotroph Diazotrophs are organisms capable of nitrogen fixation, i.e. converting the relatively inert diatomic nitrogen (N2) in Earth's atmosphere into bioavailable compound forms such as ammonia. Diazotrophs are typically microorganisms such as bacteria ...

s. One way this can occur is in the

root nodule Root nodules are found on the roots of plants, primarily legumes, that form a symbiosis with nitrogen-fixing bacteria. Under nitrogen-limiting conditions, capable plants form a symbiotic relationship with a host-specific strain of bacteria known ...

s of

legumes Legumes are plants in the pea family Fabaceae (or Leguminosae), or the fruit or seeds of such plants. When used as a dry grain for human consumption, the seeds are also called pulses. Legumes are grown agriculturally, primarily for human consu ...

that contain symbiotic

bacteria Bacteria (; : bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of Prokaryote, prokaryotic microorganisms. Typically a few micr ...

of the genera ''

Rhizobium ''Rhizobium'' is a genus of Gram-negative soil bacteria that fix nitrogen. ''Rhizobium'' species form an endosymbiotic nitrogen-fixing association with roots of (primarily) legumes and other flowering plants. The bacteria colonize plant ce ...

'', ''

Mesorhizobium ''Mesorhizobium'' is a genus of Gram-negative soil bacteria. At least one, the nitrogen fixing species, ''Mesorhizobium loti'', forms symbiotic root nodule Root nodules are found on the roots of plants, primarily legumes, that form a symbiosi ...

'', ''

Sinorhizobium ''Ensifer'' (often referred to in literature by its synonym ''Sinorhizobium'') is a genus of nitrogen-fixing bacteria (rhizobia), three of which ('' Ensifer meliloti'', '' Ensifer medicae'' and '' Ensifer fredii'') have been sequenced. Etymology ...

'', ''

Bradyrhizobium ''Bradyrhizobium'' is a genus of Gram-negative bacteria, Gram-negative soil bacteria, many of which fix nitrogen. Nitrogen fixation is an important part of the nitrogen cycle. Plants cannot use atmospheric nitrogen (N2); they must use nitrogen co ...

'', and '' Azorhizobium''. The

root In vascular plants, the roots are the plant organ, organs of a plant that are modified to provide anchorage for the plant and take in water and nutrients into the plant body, which allows plants to grow taller and faster. They are most often bel ...

s of plants create a narrow region known as the

rhizosphere The rhizosphere is the narrow region of soil or Substrate (biology), substrate that is directly influenced by root secretions and associated soil microorganisms known as the root microbiome. Pore space in soil, Soil pores in the rhizosphere can ...

that supports many microorganisms known as the root microbiome. These microorganisms in the root microbiome are able to interact with each other and surrounding plants through signals and cues. For example,

mycorrhizal fungi A mycorrhiza (; , mycorrhiza, or mycorrhizas) is a symbiotic association between a fungus and a plant. The term mycorrhiza refers to the role of the fungus in the plant's rhizosphere, the plant root system and its surroundings. Mycorrhizae play ...

are able to communicate with the root systems of many plants through chemical signals between both the plant and fungi. This results in a mutualistic

symbiosis Symbiosis (Ancient Greek : living with, companionship < : together; and ''bíōsis'': living) is any type of a close and long-term biological interaction, between two organisms of different species. The two organisms, termed symbionts, can fo ...

between the two. However, these signals can be eavesdropped by other microorganisms, such as the soil bacteria, ''Myxococcus xanthus'', which preys on other bacteria. Eavesdropping, or the interception of signals from unintended receivers, such as plants and microorganisms, can lead to large-scale, evolutionary consequences. For example, signaler-receiver pairs, like plant-microorganism pairs, may lose the ability to communicate with neighboring populations because of variability in eavesdroppers. In adapting to avoid local eavesdroppers, signal divergence could occur and thus, lead to the isolation of plants and microorganisms from the inability to communicate with other populations.

Symbiosis

A

lichen A lichen ( , ) is a hybrid colony (biology), colony of algae or cyanobacteria living symbiotically among hypha, filaments of multiple fungus species, along with yeasts and bacteria embedded in the cortex or "skin", in a mutualism (biology), m ...

is a

symbiosis Symbiosis (Ancient Greek : living with, companionship < : together; and ''bíōsis'': living) is any type of a close and long-term biological interaction, between two organisms of different species. The two organisms, termed symbionts, can fo ...

of a macroscopic fungus with

photosynthetic Photosynthesis ( ) is a Biological system, system of biological processes by which Photoautotrophism, photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical ener ...

microbial

alga Algae ( , ; : alga ) is an informal term for any organisms of a large and diverse group of photosynthetic organisms that are not plants, and includes species from multiple distinct clades. Such organisms range from unicellular microalgae, suc ...

e or

cyanobacteria Cyanobacteria ( ) are a group of autotrophic gram-negative bacteria that can obtain biological energy via oxygenic photosynthesis. The name "cyanobacteria" () refers to their bluish green (cyan) color, which forms the basis of cyanobacteri ...

.

Applications

Microorganisms are useful in producing foods, treating waste water, creating biofuels and a wide range of chemicals and enzymes. They are invaluable in research as

model organism A model organism is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the workings of other organisms. Mo ...

s. They have been weaponised and sometimes used in

warfare War is an armed conflict between the armed forces of State (polity), states, or between governmental forces and armed groups that are organized under a certain command structure and have the capacity to sustain military operations, or betwe ...

and

bioterrorism Bioterrorism is terrorism involving the intentional release or dissemination of biological agents. These agents include bacteria, viruses, insects, fungi, and/or their toxins, and may be in a naturally occurring or a human-modified form, in mu ...

. They are vital to agriculture through their roles in maintaining

soil fertility Soil fertility refers to the ability of soil to sustain agricultural plant growth, i.e. to provide plant habitat and result in sustained and consistent yields of high quality.

and in decomposing organic matter. They also have applications in aquaculture, such as in

biofloc technology Biofloc technology (BFT) is a system of aquaculture that uses "microbial biotechnology to increase the efficacy and utilization of fish feeds, where toxic materials such as nitrogen components are treated and converted to a useful product, like ...

.

Food production

Microorganisms are used in a

fermentation Fermentation is a type of anaerobic metabolism which harnesses the redox potential of the reactants to make adenosine triphosphate (ATP) and organic end products. Organic molecules, such as glucose or other sugars, are catabolized and reduce ...

process to make

yoghurt Yogurt (; , from , ; also spelled yoghurt, yogourt or yoghourt) is a food produced by bacterial fermentation of milk. Fermentation of sugars in the milk by these bacteria produces lactic acid, which acts on milk protein to give yogurt its tex ...

,

cheese Cheese is a type of dairy product produced in a range of flavors, textures, and forms by coagulation of the milk protein casein. It comprises proteins and fat from milk (usually the milk of cows, buffalo, goats or sheep). During prod ...

,

curd Curd is obtained by Denaturation (biochemistry), coagulating milk in a sequential process called curdling. It can be a final dairy product or the first stage in cheesemaking. The coagulation can be caused by adding rennet, a Kefir cheese, ...

,

kefir Kefir ( ; alternative spellings: kephir or kefier; ; ; ) is a Fermented milk products, fermented milk drink similar to a thin yogurt or ayran that is made from kefir grains, a specific type of mesophilic SCOBY, symbiotic culture. It is prep ...

,

ayran Ayran ( ) is a cold savory yogurt-based beverage that is consumed across Central Asia, and the Balkans, in Turkey and Iran. The principal ingredients are yogurt, water and salt. Herbs such as mint may be optionally added. Some varieties are ...

, xynogala, and other types of food. Fermentation cultures provide flavour and aroma, and inhibit undesirable organisms. They are used to

leaven In cooking, a leavening agent () or raising agent, also called a leaven () or leavener, is any one of a number of substances used in doughs and batter (cooking), batters that cause a Effervescence, foaming action (gas bubbles) that lightens and ...

bread Bread is a baked food product made from water, flour, and often yeast. It is a staple food across the world, particularly in Europe and the Middle East. Throughout recorded history and around the world, it has been an important part of many cu ...

, and to convert

sugar Sugar is the generic name for sweet-tasting, soluble carbohydrates, many of which are used in food. Simple sugars, also called monosaccharides, include glucose Glucose is a sugar with the Chemical formula#Molecular formula, molecul ...

s to

alcohol Alcohol may refer to: Common uses * Alcohol (chemistry), a class of compounds * Ethanol, one of several alcohols, commonly known as alcohol in everyday life ** Alcohol (drug), intoxicant found in alcoholic beverages ** Alcoholic beverage, an alco ...

in

wine Wine is an alcoholic drink made from Fermentation in winemaking, fermented fruit. Yeast in winemaking, Yeast consumes the sugar in the fruit and converts it to ethanol and carbon dioxide, releasing heat in the process. Wine is most often made f ...

and

beer Beer is an alcoholic beverage produced by the brewing and fermentation of starches from cereal grain—most commonly malted barley, although wheat, maize (corn), rice, and oats are also used. The grain is mashed to convert starch in the ...

. Microorganisms are used in

brewing Brewing is the production of beer by steeping a starch source (commonly cereal grains, the most popular of which is barley) in water and #Fermenting, fermenting the resulting sweet liquid with Yeast#Beer, yeast. It may be done in a brewery ...

,

wine making Winemaking, wine-making, or vinification is the production of wine, starting with the selection of the fruit, its fermentation into alcohol, and the bottling of the finished liquid. The history of wine-making stretches over millennia. There is ...

,

baking Baking is a method of preparing food that uses dry heat, typically in an oven, but it can also be done in hot ashes, or on hot Baking stone, stones. Bread is the most commonly baked item, but many other types of food can also be baked. Heat is ...

,

pickling Pickling is the process of food preservation, preserving or extending the shelf life of food by either Anaerobic organism, anaerobic fermentation (food), fermentation in brine or immersion in vinegar. The pickling procedure typically affects t ...

and other

food Food is any substance consumed by an organism for Nutrient, nutritional support. Food is usually of plant, animal, or Fungus, fungal origin and contains essential nutrients such as carbohydrates, fats, protein (nutrient), proteins, vitamins, ...

-making processes.

Water treatment

These depend for their ability to clean up water contaminated with organic material on microorganisms that can respire dissolved substances. Respiration may be aerobic, with a well-oxygenated filter bed such as a slow sand filter.

Anaerobic digestion Anaerobic digestion is a sequence of processes by which microorganisms break down biodegradable material in the absence of oxygen. The process is used for industrial or domestic purposes to Waste management, manage waste or to produce fuels. Mu ...

by

methanogen Methanogens are anaerobic archaea that produce methane as a byproduct of their energy metabolism, i.e., catabolism. Methane production, or methanogenesis, is the only biochemical pathway for Adenosine triphosphate, ATP generation in methanogens. A ...

s generate useful

methane Methane ( , ) is a chemical compound with the chemical formula (one carbon atom bonded to four hydrogen atoms). It is a group-14 hydride, the simplest alkane, and the main constituent of natural gas. The abundance of methane on Earth makes ...

gas as a by-product.

Energy

Microorganisms are used in fermentation to produce ethanol, and in

biogas Biogas is a gaseous renewable energy source produced from raw materials such as agricultural waste, manure, municipal waste, plant material, sewage, green waste, Wastewater treatment, wastewater, and food waste. Biogas is produced by anaerobic ...

reactors to produce

methane Methane ( , ) is a chemical compound with the chemical formula (one carbon atom bonded to four hydrogen atoms). It is a group-14 hydride, the simplest alkane, and the main constituent of natural gas. The abundance of methane on Earth makes ...

. Scientists are researching the use of algae to produce liquid fuels, and bacteria to convert various forms of agricultural and urban waste into usable fuels.

Chemicals, enzymes

Microorganisms are used to produce many commercial and industrial chemicals,

enzymes An enzyme () is a protein that acts as a biological catalyst by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as pro ...

and other bioactive molecules. Organic acids produced on a large industrial scale by microbial fermentation include

acetic acid Acetic acid , systematically named ethanoic acid , is an acidic, colourless liquid and organic compound with the chemical formula (also written as , , or ). Vinegar is at least 4% acetic acid by volume, making acetic acid the main compone ...

produced by

acetic acid bacteria Acetic acid bacteria (AAB) are a group of Gram-negative bacteria which Oxidation, oxidize sugars or ethanol and produce acetic acid during Aerobic fermentation, fermentation. The acetic acid bacteria consist of 10 genus, genera in the family Acet ...

such as ''

Acetobacter aceti ''Acetobacter aceti,'' a Gram-negative bacterium that moves using its peritrichous flagella, was discovered when Louis Pasteur proved it to be the cause of conversion of ethanol to acetic acid in 1864. Today, ''A.'' ''aceti'' is recognized as a s ...

'',

butyric acid Butyric acid (; from , meaning "butter"), also known under the systematic name butanoic acid, is a straight-chain alkyl carboxylic acid with the chemical formula . It is an oily, colorless liquid with an unpleasant odor. Isobutyric acid (2-met ...

made by the bacterium ''

Clostridium butyricum ''Clostridium butyricum'' is a strictly anaerobic endospore-forming Gram-positive butyric acid–producing bacillus subsisting by means of fermentation using an intracellularly accumulated amylopectin-like α- polyglucan (granulose) as a sub ...

'',

lactic acid Lactic acid is an organic acid. It has the molecular formula C3H6O3. It is white in the solid state and it is miscible with water. When in the dissolved state, it forms a colorless solution. Production includes both artificial synthesis as wel ...

made by ''

Lactobacillus ''Lactobacillus'' is a genus of gram-positive, aerotolerant anaerobes or microaerophilic, rod-shaped, non-spore-forming bacteria. Until 2020, the genus ''Lactobacillus'' comprised over 260 phylogenetically, ecologically, and metabolically div ...

'' and other

lactic acid bacteria Lactobacillales are an order of gram-positive, low-GC, acid-tolerant, generally nonsporulating, nonrespiring, either rod-shaped (bacilli) or spherical ( cocci) bacteria that share common metabolic and physiological characteristics. These bact ...

, and

citric acid Citric acid is an organic compound with the formula . It is a Transparency and translucency, colorless Weak acid, weak organic acid. It occurs naturally in Citrus, citrus fruits. In biochemistry, it is an intermediate in the citric acid cycle, ...

produced by the mould fungus ''

Aspergillus niger ''Aspergillus niger'' is a mold classified within the ''Nigri'' section of the ''Aspergillus'' genus. The ''Aspergillus'' genus consists of common molds found throughout the environment within soil and water, on vegetation, in fecal matter, on de ...

''. Microorganisms are used to prepare bioactive molecules such as

Streptokinase Streptokinase is a thrombolysis, thrombolytic medication activating plasminogen by nonenzymatic mechanism. As a medication it is used to thrombolysis, break down clots in some cases of myocardial infarction (heart attack), pulmonary embolism, an ...

from the bacterium ''

Streptococcus ''Streptococcus'' is a genus of gram-positive spherical bacteria that belongs to the family Streptococcaceae, within the order Lactobacillales (lactic acid bacteria), in the phylum Bacillota. Cell division in streptococci occurs along a sing ...

'', Cyclosporin A from the ascomycete fungus '' Tolypocladium inflatum'', and

statins Statins (or HMG-CoA reductase inhibitors) are a class of medications that lower cholesterol. They are prescribed typically to people who are at high risk of cardiovascular disease. Low-density lipoprotein (LDL) carriers of cholesterol play ...

produced by the yeast '' Monascus purpureus''.

Science

Microorganisms are essential tools in

biotechnology Biotechnology is a multidisciplinary field that involves the integration of natural sciences and Engineering Science, engineering sciences in order to achieve the application of organisms and parts thereof for products and services. Specialists ...

,

biochemistry Biochemistry, or biological chemistry, is the study of chemical processes within and relating to living organisms. A sub-discipline of both chemistry and biology, biochemistry may be divided into three fields: structural biology, enzymology, a ...

,

genetics Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinians, Augustinian ...

, and

molecular biology Molecular biology is a branch of biology that seeks to understand the molecule, molecular basis of biological activity in and between Cell (biology), cells, including biomolecule, biomolecular synthesis, modification, mechanisms, and interactio ...

. The

yeast Yeasts are eukaryotic, single-celled microorganisms classified as members of the fungus kingdom (biology), kingdom. The first yeast originated hundreds of millions of years ago, and at least 1,500 species are currently recognized. They are est ...

s ''

Saccharomyces cerevisiae ''Saccharomyces cerevisiae'' () (brewer's yeast or baker's yeast) is a species of yeast (single-celled fungal microorganisms). The species has been instrumental in winemaking, baking, and brewing since ancient times. It is believed to have be ...

'' and ''

Schizosaccharomyces pombe ''Schizosaccharomyces pombe'', also called "fission yeast", is a species of yeast used in traditional brewing and as a model organism in molecular and cell biology. It is a unicellular eukaryote, whose cells are rod-shaped. Cells typically meas ...

'' are important

model organism A model organism is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the workings of other organisms. Mo ...

s in science, since they are simple eukaryotes that can be grown rapidly in large numbers and are easily manipulated. They are particularly valuable in

genetics Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinians, Augustinian ...

,

genomics Genomics is an interdisciplinary field of molecular biology focusing on the structure, function, evolution, mapping, and editing of genomes. A genome is an organism's complete set of DNA, including all of its genes as well as its hierarchical, ...

and

proteomics Proteomics is the large-scale study of proteins. Proteins are vital macromolecules of all living organisms, with many functions such as the formation of structural fibers of muscle tissue, enzymatic digestion of food, or synthesis and replicatio ...

. Microorganisms can be harnessed for uses such as creating steroids and treating skin diseases. Scientists are also considering using microorganisms for living

fuel cells A fuel cell is an electrochemical cell that converts the chemical energy of a fuel (often hydrogen) and an oxidizing agent (often oxygen) into electricity through a pair of redox reactions. Fuel cells are different from most batteries in req ...

, and as a solution for pollution.

Warfare

In the

Middle Ages In the history of Europe, the Middle Ages or medieval period lasted approximately from the 5th to the late 15th centuries, similarly to the post-classical period of global history. It began with the fall of the Western Roman Empire and ...

, as an early example of

biological warfare Biological warfare, also known as germ warfare, is the use of biological toxins or Pathogen, infectious agents such as bacteria, viruses, insects, and Fungus, fungi with the intent to kill, harm or incapacitate humans, animals or plants as an ...

, diseased corpses were thrown into castles during

siege A siege () . is a military blockade of a city, or fortress, with the intent of conquering by attrition, or by well-prepared assault. Siege warfare (also called siegecrafts or poliorcetics) is a form of constant, low-intensity conflict charact ...

s using catapults or other

siege engine A siege engine is a device that is designed to break or circumvent heavy castle doors, thick city walls and other fortifications in siege warfare. Some are immobile, constructed in place to attack enemy fortifications from a distance, while othe ...

s. Individuals near the corpses were exposed to the pathogen and were likely to spread that pathogen to others. In modern times,

bioterrorism Bioterrorism is terrorism involving the intentional release or dissemination of biological agents. These agents include bacteria, viruses, insects, fungi, and/or their toxins, and may be in a naturally occurring or a human-modified form, in mu ...

has included the

1984 Rajneeshee bioterror attack In 1984, 751 people suffered food poisoning in The Dalles, Oregon, United States, due to the deliberate contamination of salad bars at ten local restaurants with ''Salmonella''. A group of prominent followers of Rajneesh (also known as O ...

and the 1993 release of

anthrax Anthrax is an infection caused by the bacterium '' Bacillus anthracis'' or ''Bacillus cereus'' biovar ''anthracis''. Infection typically occurs by contact with the skin, inhalation, or intestinal absorption. Symptom onset occurs between one ...

by

Aum Shinrikyo , better known by their former name , is a Japanese new religions, Japanese new religious movement and doomsday cult founded by Shoko Asahara in 1987. It carried out the deadly Tokyo subway sarin attack in 1995 and was found to have been respo ...

in Tokyo.

Soil

Microbes can make

nutrient A nutrient is a substance used by an organism to survive, grow and reproduce. The requirement for dietary nutrient intake applies to animals, plants, fungi and protists. Nutrients can be incorporated into cells for metabolic purposes or excret ...

s and minerals in the soil available to plants, produce

hormones A hormone (from the Greek participle , "setting in motion") is a class of signaling molecules in multicellular organisms that are sent to distant organs or tissues by complex biological processes to regulate physiology and behavior. Hormones a ...

that spur growth, stimulate the plant

immune system The immune system is a network of biological systems that protects an organism from diseases. It detects and responds to a wide variety of pathogens, from viruses to bacteria, as well as Tumor immunology, cancer cells, Parasitic worm, parasitic ...

and trigger or dampen stress responses. In general a more diverse set of

soil Soil, also commonly referred to as earth, is a mixture of organic matter, minerals, gases, water, and organisms that together support the life of plants and soil organisms. Some scientific definitions distinguish dirt from ''soil'' by re ...

microbes results in fewer plant diseases and higher yield.

Human health

Human gut flora

Microorganisms can form an

endosymbiotic An endosymbiont or endobiont is an organism that lives within the body or cells of another organism. Typically the two organisms are in a mutualistic relationship. Examples are nitrogen-fixing bacteria (called rhizobia), which live in the root ...

relationship with other, larger organisms. For example, microbial symbiosis plays a crucial role in the immune system. The microorganisms that make up the

gut flora Gut microbiota, gut microbiome, or gut flora are the microorganisms, including bacteria, archaea, fungi, and viruses, that live in the digestive tracts of animals. The gastrointestinal metagenome is the aggregate of all the genomes of the g ...

in the

gastrointestinal tract The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the Digestion, digestive system that leads from the mouth to the anus. The tract is the largest of the body's systems, after the cardiovascula ...

contribute to gut immunity, synthesize

vitamin Vitamins are Organic compound, organic molecules (or a set of closely related molecules called vitamer, vitamers) that are essential to an organism in small quantities for proper metabolism, metabolic function. Nutrient#Essential nutrients, ...

s such as

folic acid Folate, also known as vitamin B9 and folacin, is one of the B vitamins. Manufactured folic acid, which is converted into folate by the body, is used as a dietary supplement and in food fortification as it is more stable during processing and ...

and

biotin Biotin (also known as vitamin B7 or vitamin H) is one of the B vitamins. It is involved in a wide range of metabolic processes, both in humans and in other organisms, primarily related to the utilization of fats, carbohydrates, and amino acids. ...

, and ferment complex indigestible

carbohydrate A carbohydrate () is a biomolecule composed of carbon (C), hydrogen (H), and oxygen (O) atoms. The typical hydrogen-to-oxygen atomic ratio is 2:1, analogous to that of water, and is represented by the empirical formula (where ''m'' and ''n'' ...

s. Some microorganisms that are seen to be beneficial to health are termed

probiotic Probiotics are live microorganisms that provide health benefits when consumed, generally by improving or restoring the microbiota in the gut. Probiotics are considered generally safe to consume, but may cause bacteria– host interactions ...

s and are available as

dietary supplement A dietary supplement is a manufactured product intended to supplement a person's diet by taking a pill (pharmacy), pill, capsule (pharmacy), capsule, tablet (pharmacy), tablet, powder, or liquid. A supplement can provide nutrients eithe ...

s, or

food additives Food additives are substances added to food to preserve flavor or enhance taste, appearance, or other sensory qualities. Some additives, such as vinegar ( pickling), salt ( salting), smoke (smoking) and sugar (crystallization), have been used for ...

.

Disease

Microorganisms are the causative agents (

pathogen In biology, a pathogen (, "suffering", "passion" and , "producer of"), in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a Germ theory of d ...

s) in many

infectious diseases infection is the invasion of tissues by pathogens, their multiplication, and the reaction of host tissues to the infectious agent and the toxins they produce. An infectious disease, also known as a transmissible disease or communicable dise ...

. The organisms involved include

pathogenic bacteria Pathogenic bacteria are bacteria that can cause disease. This article focuses on the bacteria that are pathogenic to humans. Most species of bacteria are harmless and many are Probiotic, beneficial but others can cause infectious diseases. The nu ...

, causing diseases such as plague,

tuberculosis Tuberculosis (TB), also known colloquially as the "white death", or historically as consumption, is a contagious disease usually caused by ''Mycobacterium tuberculosis'' (MTB) bacteria. Tuberculosis generally affects the lungs, but it can al ...

and

anthrax Anthrax is an infection caused by the bacterium '' Bacillus anthracis'' or ''Bacillus cereus'' biovar ''anthracis''. Infection typically occurs by contact with the skin, inhalation, or intestinal absorption. Symptom onset occurs between one ...

;

protozoa Protozoa (: protozoan or protozoon; alternative plural: protozoans) are a polyphyletic group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic debris. Historically ...

n

parasite Parasitism is a Symbiosis, close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the Host (biology), host, causing it some harm, and is Adaptation, adapted str ...

s, causing diseases such as

malaria Malaria is a Mosquito-borne disease, mosquito-borne infectious disease that affects vertebrates and ''Anopheles'' mosquitoes. Human malaria causes Signs and symptoms, symptoms that typically include fever, Fatigue (medical), fatigue, vomitin ...

,

sleeping sickness African trypanosomiasis is an insect-borne parasitic infection of humans and other animals. Human African trypanosomiasis (HAT), also known as African sleeping sickness or simply sleeping sickness, is caused by the species '' Trypanosoma b ...

,

dysentery Dysentery ( , ), historically known as the bloody flux, is a type of gastroenteritis that results in bloody diarrhea. Other symptoms may include fever, abdominal pain, and a feeling of incomplete defecation. Complications may include dehyd ...

and

toxoplasmosis Toxoplasmosis is a parasitic disease caused by ''Toxoplasma gondii'', an apicomplexan. Infections with toxoplasmosis are associated with a variety of neuropsychiatric and behavioral conditions. Occasionally, people may have a few weeks or month ...

; and also fungi causing diseases such as

ringworm Dermatophytosis, also known as tinea and ringworm, is a mycosis, fungal infection of the skin (a dermatomycosis), that may affect skin, hair, and nails. Typically it results in a red, itchy, scaly, circular rash. Hair loss may occur in the a ...

,

candidiasis Candidiasis is a fungal infection due to any species of the genus '' Candida'' (a yeast). When it affects the mouth, in some countries it is commonly called thrush. Signs and symptoms include white patches on the tongue or other areas of the ...

or

histoplasmosis Histoplasmosis is a fungal infection caused by ''Histoplasma capsulatum''. Symptoms of this infection vary greatly, but the disease affects primarily the lungs. Occasionally, other organs are affected; called disseminated histoplasmosis, it can ...

. However, other diseases such as

influenza Influenza, commonly known as the flu, is an infectious disease caused by influenza viruses. Symptoms range from mild to severe and often include fever, runny nose, sore throat, muscle pain, headache, coughing, and fatigue. These sympto ...

, yellow fever or AIDS are caused by pathogenic viruses, which are not usually classified as living organisms and are not, therefore, microorganisms by the strict definition. No clear examples of archaean pathogens are known, although a relationship has been proposed between the presence of some archaean methanogens and human periodontal disease. Numerous microbial pathogens are capable of sexual processes that appear to facilitate their survival in their infected host.

Hygiene

Hygiene is a set of practices to avoid infection or

food spoilage Food spoilage is the process whereby food becomes unsuitable to ingest by a person; it is a matter of food safety. Bacteria and various fungi are the causes of spoilage, and can create serious consequences for consumers, but there are preventive ...

by eliminating microorganisms from the surroundings. As microorganisms, in particular bacteria, are found virtually everywhere, pathogen, harmful microorganisms may be reduced to acceptable levels rather than actually eliminated. In food preparation, microorganisms are reduced by Food preservation, preservation methods such as cooking, cleanliness of utensils, short storage periods, or by low temperatures. If complete sterility is needed, as with surgical equipment, an autoclave is used to kill microorganisms with heat and pressure.

In fiction

*''Osmosis Jones'', a 2001 film, and its show ''Ozzy & Drix'', set in a stylized version of the human body, featured Anthropomorphism, anthropomorphic microorganisms. *''War of the Worlds (2005 film)'', when alien lifeforms attempt to conquer Earth, they are ultimately defeated by a common microbe to which humans are immune.

Notes

References

External links

Microbes.info
is a microbiology information portal containing a vast collection of resources including articles, news, frequently asked questions, and links pertaining to the field of microbiology.
Our Microbial Planet
A free poster from the National Academy of Sciences about the positive roles of micro-organisms.
"Uncharted Microbial World: Microbes and Their Activities in the Environment"
Report from the American Academy of Microbiology
Understanding Our Microbial Planet: The New Science of Metagenomics
A 20-page educational booklet providing a basic overview of metagenomics and our microbial planet.
Tree of Life Eukaryotes

Microbe News from Genome News Network

On-line textbook
Through the microscope: A look at all things small
On-line microbiology textbook by Timothy Paustian and Gary Roberts, University of Wisconsin–Madison *
Methane-spewing microbe blamed in worst mass extinction. CBCNews
{{Authority control Microorganisms,

Discovery

Ancient precursors

Early modern

19th century

Classification and structure

Evolution

Archaea

Bacteria

Eukaryotes

Protists

Fungi

Plants

Ecology

Extremophiles

Plants and soil

Symbiosis

Applications

Food production

Water treatment

Energy

Chemicals, enzymes

Science

Warfare

Soil

Human health

Human gut flora

Disease

Hygiene

In fiction

See also

Notes

References

External links