A fungus ( : fungi or funguses) is any member of the group of
eukaryotic
Eukaryotes () are organisms whose Cell (biology), cells have a cell nucleus, nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the ...
organisms that includes microorganisms such as
yeast
Yeasts are eukaryotic, single-celled microorganisms classified as members of the fungus kingdom. The first yeast originated hundreds of millions of years ago, and at least 1,500 species are currently recognized. They are estimated to constit ...
s and
mold
A mold () or mould () is one of the structures 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. Not ...
s, as well as the more familiar
mushroom
A mushroom or toadstool is the fleshy, spore-bearing fruiting body of a fungus, typically produced above ground, on soil, or on its food source. ''Toadstool'' generally denotes one poisonous to humans.
The standard for the name "mushroom" is ...
s. These organisms are classified as a
kingdom
Kingdom commonly refers to:
* A monarchy ruled by a king or queen
* Kingdom (biology), a category in biological taxonomy
Kingdom may also refer to:
Arts and media Television
* ''Kingdom'' (British TV series), a 2007 British television drama s ...
, separately from the other eukaryotic kingdoms, which by one traditional classification include
Plant
Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae exclu ...
ae,
Animal
Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...
Chromista
Chromista is a biological kingdom consisting of single-celled and multicellular eukaryotic species that share similar features in their photosynthetic organelles ( plastids). It includes all protists whose plastids contain chlorophyll ''c'', ...
.
A characteristic that places fungi in a different kingdom from plants, bacteria, and some protists is chitin in their cell walls. Fungi, like animals, are
heterotroph
A heterotroph (; ) is an organism that cannot produce its own food, instead taking nutrition from other sources of organic carbon, mainly plant or animal matter. In the food chain, heterotrophs are primary, secondary and tertiary consumers, but ...
s; they acquire their food by absorbing dissolved molecules, typically by secreting
digestive enzyme
Digestive enzymes are a group of enzymes that break down polymeric macromolecules into their smaller building blocks, in order to facilitate their absorption into the cells of the body. Digestive enzymes are found in the digestive tracts of anima ...
s into their environment. Fungi do not
photosynthesize
Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in ...
. Growth is their means of
mobility
Mobility may refer to:
Social sciences and humanities
* Economic mobility, ability of individuals or families to improve their economic status
* Geographic mobility, the measure of how populations and goods move over time
* Mobilities, a conte ...
, except for spores (a few of which are flagellated), which may travel through the air or water. Fungi are the principal decomposers in ecological systems. These and other differences place fungi in a single group of related organisms, named the ''Eumycota'' (''true fungi'' or ''Eumycetes''), that share a
common ancestor
Common descent is a concept in evolutionary biology applicable when one species is the ancestor of two or more species later in time. All living beings are in fact descendants of a unique ancestor commonly referred to as the last universal comm ...
(i.e. they form a '' monophyletic group''), an interpretation that is also strongly supported by
molecular phylogenetics
Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
. This fungal group is distinct from the structurally similar myxomycetes (slime molds) and
oomycete
Oomycota forms a distinct phylogenetic lineage of fungus-like eukaryotic microorganisms, called oomycetes (). They are filamentous and heterotrophic, and can reproduce both sexually and asexually. Sexual reproduction of an oospore is the resul ...
s (water molds). The discipline of
biology
Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells that process hereditary i ...
devoted to the study of fungi is known as
mycology
Mycology is the branch of biology concerned with the study of fungi, including their genetic and biochemical properties, their taxonomy and their use to humans, including as a source for tinder, traditional medicine, food, and entheogen ...
(from the
Greek
Greek may refer to:
Greece
Anything of, from, or related to Greece, a country in Southern Europe:
*Greeks, an ethnic group.
*Greek language, a branch of the Indo-European language family.
**Proto-Greek language, the assumed last common ancestor ...
', mushroom). In the past, mycology was regarded as a branch of
botany
Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytologist is a scientist who specialises in this field. The term "botany" comes from the Ancient Greek w ...
, although it is now known fungi are genetically more closely related to animals than to plants.
Abundant worldwide, most fungi are inconspicuous because of the small size of their structures, and their cryptic lifestyles in soil or on dead matter. Fungi include symbionts of plants, animals, or other fungi and also
parasites
Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson ha ...
. They may become noticeable when
fruiting
In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering.
Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in particula ...
, either as mushrooms or as molds. Fungi perform an essential role in the decomposition of organic matter and have fundamental roles in nutrient
cycling
Cycling, also, when on a two-wheeled bicycle, called bicycling or biking, is the use of cycles for transport, recreation, exercise or sport. People engaged in cycling are referred to as "cyclists", "bicyclists", or "bikers". Apart from t ...
and exchange in the environment. They have long been used as a direct source of human food, in the form of mushrooms and truffles; as a
leavening agent
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 batters that cause a foaming action (gas bubbles) that lightens and softens the mixture. An altern ...
for bread; and in the fermentation of various food products, such as
wine
Wine is an alcoholic drink typically made from fermented grapes. Yeast consumes the sugar in the grapes and converts it to ethanol and carbon dioxide, releasing heat in the process. Different varieties of grapes and strains of yeasts are m ...
,
beer
Beer is one of the oldest and the most widely consumed type of alcoholic drink in the world, and the third most popular drink overall after water and tea. It is produced by the brewing and fermentation of starches, mainly derived from ce ...
, and soy sauce. Since the 1940s, fungi have been used for the production of antibiotics, and, more recently, various
enzyme
Enzymes () are proteins that act as biological catalysts 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 products ...
mycotoxin
A mycotoxin (from the Greek μύκης , "fungus" and τοξίνη , "toxin") is a toxic secondary metabolite produced by organisms of kingdom Fungi and is capable of causing disease and death in both humans and other animals. The term 'mycotoxin' ...
s, such as
alkaloid
Alkaloids are a class of basic, naturally occurring organic compounds that contain at least one nitrogen atom. This group also includes some related compounds with neutral and even weakly acidic properties. Some synthetic compounds of similar ...
s and
polyketide
Polyketides are a class of natural products derived from a precursor molecule consisting of a chain of alternating ketone (or reduced forms of a ketone) and methylene groups: (-CO-CH2-). First studied in the early 20th century, discovery, biosynth ...
s, that are toxic to animals, including humans. The fruiting structures of a few species contain
psychotropic
A psychoactive drug, psychopharmaceutical, psychoactive agent or psychotropic drug is a chemical substance, that changes functions of the nervous system, and results in alterations in perception, mood, consciousness, cognition or behavior.
Th ...
compounds and are consumed recreationally or in traditional spiritual ceremonies. Fungi can break down manufactured materials and buildings, and become significant
pathogens
In biology, a pathogen ( el, πάθος, "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 ger ...
of humans and other animals. Losses of crops due to fungal diseases (e.g., rice blast disease) or
food spoilage
Food spoilage is the process where a food product becomes unsuitable to ingest by the consumer. The cause of such a process is due to many outside factors as a side-effect of the type of product it is, as well as how the product is packaged and s ...
can have a large impact on human food supplies and local economies.
The fungus kingdom encompasses an enormous diversity of
taxa
In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
with varied ecologies,
life cycle
Life cycle, life-cycle, or lifecycle may refer to:
Science and academia
*Biological life cycle, the sequence of life stages that an organism undergoes from birth to reproduction ending with the production of the offspring
* Life-cycle hypothesis ...
strategies, and morphologies ranging from unicellular aquatic chytrids to large mushrooms. However, little is known of the true
biodiversity
Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic (''genetic variability''), species (''species diversity''), and ecosystem (''ecosystem diversity'') l ...
of the fungus kingdom, which has been estimated at 2.2 million to 3.8 million species. Of these, only about 148,000 have been described, with over 8,000 species known to be detrimental to plants and at least 300 that can be pathogenic to humans. Ever since the pioneering 18th and 19th century
taxonomical
In biology, taxonomy () is the scientific study of naming, defining ( circumscribing) and classifying groups of biological organisms based on shared characteristics. Organisms are grouped into taxa (singular: taxon) and these groups are given ...
works of
Carl Linnaeus
Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his Nobility#Ennoblement, ennoblement in 1761 as Carl von Linné#Blunt, Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalise ...
, Christiaan Hendrik Persoon, and Elias Magnus Fries, fungi have been classified according to their morphology (e.g., characteristics such as spore color or microscopic features) or
physiology
Physiology (; ) is the scientific study of functions and mechanisms in a living system. As a sub-discipline of biology, physiology focuses on how organisms, organ systems, individual organs, cells, and biomolecules carry out the chemical ...
. Advances in
molecular genetics
Molecular genetics is a sub-field of biology that addresses how differences in the structures or expression of DNA molecules manifests as variation among organisms. Molecular genetics often applies an "investigative approach" to determine the ...
have opened the way for DNA analysis to be incorporated into taxonomy, which has sometimes challenged the historical groupings based on morphology and other traits.
Phylogenetic
In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
studies published in the first decade of the 21st century have helped reshape the classification within the fungi kingdom, which is divided into one
subkingdom
In biology, a kingdom is the second highest taxonomic rank, just below domain. Kingdoms are divided into smaller groups called phyla.
Traditionally, some textbooks from the United States and Canada used a system of six kingdoms (Animalia, Plant ...
The English word ''fungus'' is directly adopted from the
Latin
Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
''fungus'' (mushroom), used in the writings of Horace and
Pliny
Pliny may refer to:
People
* Pliny the Elder (23–79 CE), ancient Roman nobleman, scientist, historian, and author of ''Naturalis Historia'' (''Pliny's Natural History'')
* Pliny the Younger (died 113), ancient Roman statesman, orator, w ...
. This in turn is derived from the
Greek
Greek may refer to:
Greece
Anything of, from, or related to Greece, a country in Southern Europe:
*Greeks, an ethnic group.
*Greek language, a branch of the Indo-European language family.
**Proto-Greek language, the assumed last common ancestor ...
word ''sphongos'' (σφόγγος 'sponge'), which refers to the
macroscopic
The macroscopic scale is the length scale on which objects or phenomena are large enough to be visible with the naked eye, without magnifying optical instruments. It is the opposite of microscopic.
Overview
When applied to physical phenomena a ...
structures and morphology of mushrooms and molds;Ainsworth, p. 2. the
root
In vascular plants, the roots are the 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 below the su ...
is also used in other languages, such as the German '' Schwamm'' ('sponge') and '' Schimmel'' ('mold').
The word ''
mycology
Mycology is the branch of biology concerned with the study of fungi, including their genetic and biochemical properties, their taxonomy and their use to humans, including as a source for tinder, traditional medicine, food, and entheogen ...
'' is derived from the Greek (μύκης 'mushroom') and ''logos'' (λόγος 'discourse'). It denotes the scientific study of fungi. The Latin adjectival form of "mycology" (''mycologicæ'') appeared as early as 1796 in a book on the subject by Christiaan Hendrik Persoon. The word appeared in English as early as 1824 in a book by
Robert Kaye Greville
Dr. Robert Kaye Greville FRSE FLS LLD (13 December 1794 – 4 June 1866) was an English mycologist, bryologist, and botanist. He was an accomplished artist and illustrator of natural history. In addition to art and science he was interest ...
. In 1836 the English naturalist
Miles Joseph Berkeley
Miles Joseph Berkeley (1 April 1803 – 30 July 1889) was an English cryptogamist and clergyman, and one of the founders of the science of plant pathology.
Life
Berkeley was born at Biggin Hall, Benefield, Northamptonshire, and educated at R ...
's publication ''The English Flora of Sir James Edward Smith, Vol. 5.'' also refers to mycology as the study of fungi.
A group of all the fungi present in a particular region is known as ''
mycobiota
Mycobiota (plural noun, no singular) are a group of all the fungi present in a particular geographic region (e.g. "the mycobiota of Ireland") or habitat type (e.g. "the mycobiota of cocoa").
Human mycobiota
Mycobiota exist on the surface and in th ...
'' (plural noun, no singular). The term ''mycota'' is often used for this purpose, but many authors use it as a synonym of Fungi. The word '' funga'' has been proposed as a less ambiguous term morphologically similar to
fauna
Fauna is all of the animal life present in a particular region or time. The corresponding term for plants is ''flora'', and for fungi, it is ''funga''. Flora, fauna, funga and other forms of life are collectively referred to as ''Biota (ecology ...
and
flora
Flora is all the plant life present in a particular region or time, generally the naturally occurring (indigenous (ecology), indigenous) native plant, native plants. Sometimes bacteria and fungi are also referred to as flora, as in the terms '' ...
International Union for Conservation of Nature
The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of nat ...
(IUCN) in August 2021 asked that the phrase ''fauna and flora'' be replaced by ''fauna, flora, and funga''.
Characteristics
Before the introduction of molecular methods for phylogenetic analysis, taxonomists considered fungi to be members of the plant kingdom because of similarities in lifestyle: both fungi and plants are mainly immobile, and have similarities in general morphology and growth habitat. Although inaccurate, the common misconception that fungi are plants persists among the general public due to their historical classification, as well as several similarities. Like plants, fungi often grow in soil and, in the case of
mushroom
A mushroom or toadstool is the fleshy, spore-bearing fruiting body of a fungus, typically produced above ground, on soil, or on its food source. ''Toadstool'' generally denotes one poisonous to humans.
The standard for the name "mushroom" is ...
s, form conspicuous fruit bodies, which sometimes resemble plants such as mosses. The fungi are now considered a separate kingdom, distinct from both plants and animals, from which they appear to have diverged around one billion years ago (around the start of the
Neoproterozoic
The Neoproterozoic Era is the unit of geologic time from 1 billion to 538.8 million years ago.
It is the last era of the Precambrian Supereon and the Proterozoic Eon; it is subdivided into the Tonian, Cryogenian, and Ediacaran periods. It is prec ...
Era). Some morphological, biochemical, and genetic features are shared with other organisms, while others are unique to the fungi, clearly separating them from the other kingdoms:
Shared features:
* With other eukaryotes: Fungal cells contain membrane-boundnuclei with
chromosomes
A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins are ...
80S
Ribosomes are a large and complex molecular machine that catalyzes the synthesis of proteins, referred to as translation. The ribosome selects aminoacylated transfer RNAs (tRNAs) based on the sequence of a protein-encoding messenger RNA (mRN ...
type. They have a characteristic range of soluble carbohydrates and storage compounds, including
sugar alcohol
Sugar alcohols (also called polyhydric alcohols, polyalcohols, alditols or glycitols) are organic compounds, typically derived from sugars, containing one hydroxyl group (–OH) attached to each carbon atom. They are white, water-soluble solids ...
s (e.g.,
mannitol
Mannitol is a type of sugar alcohol used as a sweetener and medication. It is used as a low calorie sweetener as it is poorly absorbed by the intestines. As a medication, it is used to decrease pressure in the eyes, as in glaucoma, and to lo ...
),
disaccharide
A disaccharide (also called a double sugar or ''biose'') is the sugar formed when two monosaccharides are joined by glycosidic linkage. Like monosaccharides, disaccharides are simple sugars soluble in water. Three common examples are sucrose, la ...
s, (e.g.,
trehalose
Trehalose (from Turkish '' tıgala'' – a sugar derived from insect cocoons + -ose) is a sugar consisting of two molecules of glucose. It is also known as mycose or tremalose. Some bacteria, fungi, plants and invertebrate animals synthesize it ...
heterotroph
A heterotroph (; ) is an organism that cannot produce its own food, instead taking nutrition from other sources of organic carbon, mainly plant or animal matter. In the food chain, heterotrophs are primary, secondary and tertiary consumers, but ...
ic organisms and so require preformed
organic compound
In chemistry, organic compounds are generally any chemical compounds that contain carbon-hydrogen or carbon-carbon bonds. Due to carbon's ability to catenate (form chains with other carbon atoms), millions of organic compounds are known. T ...
s as energy sources.
* With plants: Fungi have a cell wall and vacuoles. They reproduce by both sexual and asexual means, and like basal plant groups (such as
fern
A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes exce ...
s and mosses) produce spores. Similar to mosses and algae, fungi typically have haploid nuclei.
* With euglenoids and bacteria: Higher fungi, euglenoids, and some bacteria produce the
amino acid
Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha a ...
L-lysine in specific biosynthesis steps, called the α-aminoadipate pathway.
* The cells of most fungi grow as tubular, elongated, and thread-like (filamentous) structures called hyphae, which may contain multiple nuclei and extend by growing at their tips. Each tip contains a set of aggregated vesicles—cellular structures consisting of
protein
Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, res ...
s,
lipid
Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids includ ...
s, and other organic molecules—called the
Spitzenkörper
The Spitzenkörper (German for pointed body, SPK) is a structure found in fungal hyphae that is the organizing center for hyphal growth and morphogenesis. It consists of many small vesicles and is present in growing hyphal tips, during spore ger ...
. Both fungi and
oomycete
Oomycota forms a distinct phylogenetic lineage of fungus-like eukaryotic microorganisms, called oomycetes (). They are filamentous and heterotrophic, and can reproduce both sexually and asexually. Sexual reproduction of an oospore is the resul ...
s grow as filamentous hyphal cells. In contrast, similar-looking organisms, such as filamentous green algae, grow by repeated cell division within a chain of cells. There are also single-celled fungi (
yeast
Yeasts are eukaryotic, single-celled microorganisms classified as members of the fungus kingdom. The first yeast originated hundreds of millions of years ago, and at least 1,500 species are currently recognized. They are estimated to constit ...
s) that do not form hyphae, and some fungi have both hyphal and yeast forms.
* In common with some plant and animal species, more than 70 fungal species display bioluminescence.
Unique features:
* Some species grow as unicellular yeasts that reproduce 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 ...
or fission. Dimorphic fungi can switch between a yeast phase and a hyphal phase in response to environmental conditions.
* The fungal cell wall is made of a chitin-glucan complex; while glucans are also found in plants and chitin in the
exoskeleton
An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton (endoskeleton) in for example, a human. In usage, some of the ...
of arthropods, fungi are the only organisms that combine these two structural molecules in their cell wall. Unlike those of plants and oomycetes, fungal cell walls do not contain cellulose.
Most fungi lack an efficient system for the long-distance transport of water and nutrients, such as the
xylem
Xylem is one of the two types of transport tissue in vascular plants, the other being phloem. The basic function of xylem is to transport water from roots to stems and leaves, but it also transports nutrients. The word ''xylem'' is derived from ...
and
phloem
Phloem (, ) is the living tissue in vascular plants that transports the soluble organic compounds made during photosynthesis and known as ''photosynthates'', in particular the sugar sucrose, to the rest of the plant. This transport process is c ...
in many plants. To overcome this limitation, some fungi, such as ''
Armillaria
''Armillaria'' is a genus of fungi that includes the ''Armillaria mellea, A. mellea'' species known as honey fungi that live on trees and woody shrubs. It includes about 10 species formerly categorized summarily as ''A. mellea''. ''Armill ...
'', form
rhizomorphs
Mycelial cords are linear aggregations of parallel-oriented hyphae. The mature cords are composed of wide, empty vessel hyphae surrounded by narrower sheathing hyphae. Cords may look similar to plant roots, and also frequently have similar function ...
, which resemble and perform functions similar to the
root
In vascular plants, the roots are the 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 below the su ...
s of plants. As eukaryotes, fungi possess a
biosynthetic pathway
Metabolism (, from el, μεταβολή ''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 cell ...
for producing
terpene
Terpenes () are a class of natural products consisting of compounds with the formula (C5H8)n for n > 1. Comprising more than 30,000 compounds, these unsaturated hydrocarbons are produced predominantly by plants, particularly conifers. Terpenes ...
pyrophosphate
In chemistry, pyrophosphates are phosphorus oxyanions that contain two phosphorus atoms in a P–O–P linkage. A number of pyrophosphate salts exist, such as disodium pyrophosphate (Na2H2P2O7) and tetrasodium pyrophosphate (Na4P2O7), among othe ...
as chemical building blocks. Plants and some other organisms have an additional terpene biosynthesis pathway in their chloroplasts, a structure that fungi and animals do not have. Fungi produce several
secondary metabolite
Secondary metabolites, also called specialised metabolites, toxins, secondary products, or natural products, are organic compounds produced by any lifeform, e.g. bacteria, fungi, animals, or plants, which are not directly involved in the norma ...
s that are similar or identical in structure to those made by plants. Many of the plant and fungal enzymes that make these compounds differ from each other in
sequence
In mathematics, a sequence is an enumerated collection of objects in which repetitions are allowed and order matters. Like a set, it contains members (also called ''elements'', or ''terms''). The number of elements (possibly infinite) is calle ...
and other characteristics, which indicates separate origins and
convergent evolution
Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
of these enzymes in the fungi and plants.
Diversity
Fungi have a worldwide distribution, and grow in a wide range of habitats, including extreme environments such as
deserts
A desert is a barren area of landscape where little precipitation occurs and, consequently, living conditions are hostile for plant and animal life. The lack of vegetation exposes the unprotected surface of the ground to denudation. About on ...
or areas with high salt concentrations or ionizing radiation, as well as in
deep sea
The deep sea is broadly defined as the ocean depth where light begins to fade, at an approximate depth of 200 metres (656 feet) or the point of transition from continental shelves to continental slopes. Conditions within the deep sea are a combin ...
sediments. Some can survive the intense UV and cosmic radiation encountered during space travel. Most grow in terrestrial environments, though several species live partly or solely in aquatic habitats, such as the chytrid fungi ''
Batrachochytrium dendrobatidis
''Batrachochytrium dendrobatidis'' ( ), also known as ''Bd'' or the amphibian chytrid fungus, is a fungus that causes the disease chytridiomycosis in amphibians.
Since its discovery in 1998 by Lee Berger, the disease devastated amphibian popula ...
parasite
Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has ...
s that have been responsible for a worldwide decline in amphibian populations. These organisms spend part of their life cycle as a motile
zoospore
A zoospore is a motile asexual spore that uses a flagellum for locomotion. Also called a swarm spore, these spores are created by some protists, bacteria, and fungi to propagate themselves.
Diversity Flagella types
Zoospores may possess one or ...
, enabling them to propel itself through water and enter their amphibian host. Other examples of aquatic fungi include those living in
hydrothermal
Hydrothermal circulation in its most general sense is the circulation of hot water (Ancient Greek ὕδωρ, ''water'',Liddell, H.G. & Scott, R. (1940). ''A Greek-English Lexicon. revised and augmented throughout by Sir Henry Stuart Jones. with th ...
areas of the ocean. As of 2020, around 148,000 species of fungi have been described by taxonomists, but the global biodiversity of the fungus kingdom is not fully understood. A 2017 estimate suggests there may be between 2.2 and 3.8 million species. The number of new fungi species discovered yearly has increased from 1,000 to 1,500 per year about 10 years ago, to about 2000 with a peak of more than 2,500 species in 2016. In the year 2019, 1882 new species of fungi were described, and it was estimated that more than 90% of fungi remain unknown. The following year, 2905 new species were described—the highest annual record of new fungus names. In mycology, species have historically been distinguished by a variety of methods and concepts. Classification based on morphological characteristics, such as the size and shape of spores or fruiting structures, has traditionally dominated fungal taxonomy.Kirk ''et al''., p. 489. Species may also be distinguished by their biochemical and physiological characteristics, such as their ability to metabolize certain biochemicals, or their reaction to chemical tests. The
biological species concept
The species problem is the set of questions that arises when biologists attempt to define what a species is. Such a definition is called a species concept; there are at least 26 recognized species concepts. A species concept that works well for se ...
discriminates species based on their ability to mate. The application of molecular tools, such as DNA sequencing and phylogenetic analysis, to study diversity has greatly enhanced the resolution and added robustness to estimates of genetic diversity within various taxonomic groups.
Mycology
Mycology
Mycology is the branch of biology concerned with the study of fungi, including their genetic and biochemical properties, their taxonomy and their use to humans, including as a source for tinder, traditional medicine, food, and entheogen ...
is the branch of
biology
Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells that process hereditary i ...
concerned with the systematic study of fungi, including their genetic and biochemical properties, their taxonomy, and their use to humans as a source of medicine, food, and psychotropic substances consumed for religious purposes, as well as their dangers, such as poisoning or infection. The field of phytopathology, the study of plant diseases, is closely related because many plant pathogens are fungi.
The use of fungi by humans dates back to prehistory;
Ötzi the Iceman
Ötzi, also called the Iceman, is the natural mummy of a man who lived some time between 3350 and 3105 BC, discovered in September 1991 in the Ötztal Alps (hence the nickname "Ötzi") on the border between Austria and Italy.
Ötzi is believed to ...
, a well-preserved mummy of a 5,300-year-old
Neolithic
The Neolithic period, or New Stone Age, is an Old World archaeological period and the final division of the Stone Age. It saw the Neolithic Revolution, a wide-ranging set of developments that appear to have arisen independently in several p ...
man found frozen in the Austrian Alps, carried two species of polypore mushrooms that may have been used as tinder (''
Fomes fomentarius
''Fomes fomentarius'' (commonly known as the tinder fungus, false tinder fungus, hoof fungus, tinder conk, tinder polypore or ice man fungus) is a species of fungal plant pathogen found in Europe, Asia, Africa and North America. The species pro ...
''), or for medicinal purposes (''
Piptoporus betulinus
''Fomitopsis betulina'' (previously ''Piptoporus betulinus''), commonly known as the birch polypore, birch bracket, or razor strop, is a common bracket fungus and, as the name suggests, grows almost exclusively on birch trees. The brackets bur ...
''). Ancient peoples have used fungi as food sources—often unknowingly—for millennia, in the preparation of leavened bread and fermented juices. Some of the oldest written records contain references to the destruction of crops that were probably caused by pathogenic fungi.
History
Mycology became a systematic science after the development of the
microscope
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 invisi ...
in the 17th century. Although fungal spores were first observed by
Giambattista della Porta
Giambattista della Porta (; 1535 – 4 February 1615), also known as Giovanni Battista Della Porta, was an Italian scholar, polymath and playwright who lived in Naples at the time of the Renaissance, Scientific Revolution and Reformation.
Giamb ...
in 1588, the seminal work in the development of mycology is considered to be the publication of
Pier Antonio Micheli
Pier Antonio Micheli (December 11, 1679 – January 1, 1737) was a noted Italian botanist, professor of botany in Pisa, curator of the Orto Botanico di Firenze, author of ''Nova plantarum genera iuxta Tournefortii methodum disposita''. He discove ...
's 1729 work ''Nova plantarum genera''. Micheli not only observed spores but also showed that, under the proper conditions, they could be induced into growing into the same species of fungi from which they originated. Extending the use of the binomial system of nomenclature introduced by
Carl Linnaeus
Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his Nobility#Ennoblement, ennoblement in 1761 as Carl von Linné#Blunt, Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalise ...
in his '' Species plantarum'' (1753), the Dutch Christiaan Hendrik Persoon (1761–1836) established the first classification of mushrooms with such skill as to be considered a founder of modern mycology. Later, Elias Magnus Fries (1794–1878) further elaborated the classification of fungi, using spore color and microscopic characteristics, methods still used by taxonomists today. Other notable early contributors to mycology in the 17th–19th and early 20th centuries include
Miles Joseph Berkeley
Miles Joseph Berkeley (1 April 1803 – 30 July 1889) was an English cryptogamist and clergyman, and one of the founders of the science of plant pathology.
Life
Berkeley was born at Biggin Hall, Benefield, Northamptonshire, and educated at R ...
Anton de Bary
Heinrich Anton de Bary (26 January 183119 January 1888) was a German surgeon, botanist, microbiologist, and mycologist (fungal systematics and physiology).
He is considered a founding father of plant pathology (phytopathology) as well as the fou ...
Charles Tulasne
Charles Tulasne (5 September 1816 – 28 August 1884) was a French physician, mycologist and illustrator born in Langeais in the département of Indre-et-Loire.
He received his medical doctorate in 1840 and practiced medicine in Paris until 18 ...
Pier Andrea Saccardo
Pier Andrea Saccardo (23 April 1845 in Treviso, Treviso – 12 February 1920 in Padua) was an Italian botanist and mycologist.
Life
Saccardo studied at the Lyceum in Venice, and then at the Technical Institute of the University of Padua wher ...
. In the 20th and 21st centuries, advances in
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 and ...
,
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 Augustinian friar wor ...
,
molecular biology
Molecular biology is the branch of biology that seeks to understand the molecular basis of biological activity in and between cells, including biomolecular synthesis, modification, mechanisms, and interactions. The study of chemical and physi ...
,
biotechnology
Biotechnology is the integration of natural sciences and engineering sciences in order to achieve the application of organisms, cells, parts thereof and molecular analogues for products and services. The term ''biotechnology'' was first used ...
, DNA sequencing and phylogenetic analysis has provided new insights into fungal relationships and
biodiversity
Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic (''genetic variability''), species (''species diversity''), and ecosystem (''ecosystem diversity'') l ...
, and has challenged traditional morphology-based groupings in fungal
taxonomy
Taxonomy is the practice and science of categorization or classification.
A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
.
Morphology
Microscopic structures
Most fungi grow as hyphae, which are cylindrical, thread-like structures 2–10 µm in diameter and up to several centimeters in length. Hyphae grow at their tips (apices); new hyphae are typically formed by emergence of new tips along existing hyphae by a process called ''branching'', or occasionally growing hyphal tips fork, giving rise to two parallel-growing hyphae. Hyphae also sometimes fuse when they come into contact, a process called hyphal fusion (or anastomosis). These growth processes lead to the development of a mycelium, an interconnected network of hyphae. Hyphae can be either
septate
In biology, a septum (Latin for ''something that encloses''; plural septa) is a wall, dividing a cavity or structure into smaller ones. A cavity or structure divided in this way may be referred to as septate.
Examples
Human anatomy
* Interatr ...
or coenocytic. Septate hyphae are divided into compartments separated by cross walls (internal cell walls, called septa, that are formed at right angles to the cell wall giving the hypha its shape), with each compartment containing one or more nuclei; coenocytic hyphae are not compartmentalized. Septa have pores that allow
cytoplasm
In cell biology, the cytoplasm is all of the material within a eukaryotic cell, enclosed by the cell membrane, except for the cell nucleus. The material inside the nucleus and contained within the nuclear membrane is termed the nucleoplasm. ...
, organelles, and sometimes nuclei to pass through; an example is the dolipore septum in fungi of the phylum Basidiomycota. Coenocytic hyphae are in essence
multinucleate
Multinucleate cells (also known as multinucleated or polynuclear cells) are eukaryotic cells that have more than one nucleus per cell, i.e., multiple nuclei share one common cytoplasm. Mitosis in multinucleate cells can occur either in a coordina ...
supercells.
Many species have developed specialized hyphal structures for nutrient uptake from living hosts; examples include
haustoria
In botany and mycology, a haustorium (plural haustoria) is a rootlike structure that grows into or around another structure to absorb water or nutrients. For example, in mistletoe or members of the broomrape family, the structure penetrates th ...
in plant-parasitic species of most fungal phyla, and arbuscules of several mycorrhizal fungi, which penetrate into the host cells to consume nutrients.
Although fungi are
opisthokont
The opisthokonts () are a broad group of eukaryotes, including both the animal and fungus kingdoms. The opisthokonts, previously called the "Fungi/Metazoa group", are generally recognized as a clade. Opisthokonts together with Apusomonadida and ...
s—a grouping of evolutionarily related organisms broadly characterized by a single posterior
flagellum
A flagellum (; ) is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates.
A microorganism may have f ...
—all phyla except for the chytrids have lost their posterior flagella. Fungi are unusual among the eukaryotes in having a cell wall that, in addition to glucans (e.g., β-1,3-glucan) and other typical components, also contains the biopolymer chitin.
Macroscopic structures
Fungal mycelia can become visible to the naked eye, for example, on various surfaces and substrates, such as damp walls and spoiled food, where they are commonly called
mold
A mold () or mould () is one of the structures 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. Not ...
s. Mycelia grown on solid agar media in laboratory petri dishes are usually referred to as colonies. These colonies can exhibit growth shapes and colors (due to spores or
pigmentation
A pigment is a colored material that is completely or nearly insoluble in water. In contrast, dyes are typically soluble, at least at some stage in their use. Generally dyes are often organic compounds whereas pigments are often inorganic compoun ...
) that can be used as diagnostic features in the identification of species or groups. Some individual fungal colonies can reach extraordinary dimensions and ages as in the case of a clonal colony of '' Armillaria solidipes'', which extends over an area of more than 900 ha (3.5 square miles), with an estimated age of nearly 9,000years.
The
apothecium
An ascocarp, or ascoma (), is the fruiting body ( sporocarp) of an ascomycete phylum fungus. It consists of very tightly interwoven hyphae and millions of embedded asci, each of which typically contains four to eight ascospores. Ascocarps are mos ...
—a specialized structure important in
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 th ...
in the ascomycetes—is a cup-shaped fruit body that is often macroscopic and holds the hymenium, a layer of tissue containing the spore-bearing cells. The fruit bodies of the basidiomycetes (
basidiocarp
In fungi, a basidiocarp, basidiome, or basidioma () is the sporocarp of a basidiomycete, the multicellular structure on which the spore-producing hymenium is borne. Basidiocarps are characteristic of the hymenomycetes; rusts and smuts do ...
s) and some ascomycetes can sometimes grow very large, and many are well known as
mushroom
A mushroom or toadstool is the fleshy, spore-bearing fruiting body of a fungus, typically produced above ground, on soil, or on its food source. ''Toadstool'' generally denotes one poisonous to humans.
The standard for the name "mushroom" is ...
s.
Growth and physiology
The growth of fungi as hyphae on or in solid substrates or as single cells in aquatic environments is adapted for the efficient extraction of nutrients, because these growth forms have high surface area to volume ratios. Hyphae are specifically adapted for growth on solid surfaces, and to invade substrates and tissues. They can exert large penetrative mechanical forces; for example, many plant pathogens, including '' Magnaporthe grisea'', form a structure called an appressorium that evolved to puncture plant tissues. The pressure generated by the appressorium, directed against the plant epidermis, can exceed . The filamentous fungus ''
Paecilomyces lilacinus
''Purpureocillium lilacinum'' is a species of filamentous fungus in the family Ophiocordycipitaceae. It has been isolated from a wide range of habitats, including cultivated and uncultivated soils, forests, grassland, deserts, estuarine sedime ...
'' uses a similar structure to penetrate the eggs of nematodes.
The mechanical pressure exerted by the appressorium is generated from physiological processes that increase intracellular
turgor
Turgor pressure is the force within the cell that pushes the plasma membrane against the cell wall.
It is also called ''hydrostatic pressure'', and is defined as the pressure in a fluid measured at a certain point within itself when at equilibri ...
by producing
osmolyte
Osmolytes are low-molecular weight organic compounds that influence the properties of biological fluids. Their primary role is to maintain the integrity of cells by affecting the viscosity, melting point, and ionic strength of the aqueous solution. ...
s such as
glycerol
Glycerol (), also called glycerine in British English and glycerin in American English, is a simple triol compound. It is a colorless, odorless, viscous liquid that is sweet-tasting and non-toxic. The glycerol backbone is found in lipids known ...
. Adaptations such as these are complemented by hydrolytic enzymes secreted into the environment to digest large organic molecules—such as polysaccharides,
protein
Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, res ...
s, and
lipid
Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids includ ...
s—into smaller molecules that may then be absorbed as nutrients. The vast majority of filamentous fungi grow in a polar fashion (extending in one direction) by elongation at the tip (apex) of the hypha. Other forms of fungal growth include intercalary extension (longitudinal expansion of hyphal compartments that are below the apex) as in the case of some endophytic fungi, or growth by volume expansion during the development of mushroom stipes and other large organs. Growth of fungi as multicellular structures consisting of somatic and reproductive cells—a feature independently evolved in animals and plants—has several functions, including the development of fruit bodies for dissemination of sexual spores (see above) and
biofilm
A biofilm comprises any syntrophic consortium of microorganisms in which cells stick to each other and often also to a surface. These adherent cells become embedded within a slimy extracellular matrix that is composed of extracellular ...
heterotroph
A heterotroph (; ) is an organism that cannot produce its own food, instead taking nutrition from other sources of organic carbon, mainly plant or animal matter. In the food chain, heterotrophs are primary, secondary and tertiary consumers, but ...
s, organisms that rely solely on carbon fixed by other organisms for
metabolism
Metabolism (, from el, μεταβολή ''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 c ...
. Fungi have
evolved
Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
a high degree of metabolic versatility that allows them to use a diverse range of organic substrates for growth, including simple compounds such as nitrate,
ammonia
Ammonia is an inorganic compound of nitrogen and hydrogen with the formula . A stable binary hydride, and the simplest pnictogen hydride, ammonia is a colourless gas with a distinct pungent smell. Biologically, it is a common nitrogenous wa ...
ethanol
Ethanol (abbr. EtOH; also called ethyl alcohol, grain alcohol, drinking alcohol, or simply alcohol) is an organic compound. It is an alcohol with the chemical formula . Its formula can be also written as or (an ethyl group linked to a ...
. In some species the pigment
melanin
Melanin (; from el, μέλας, melas, black, dark) is a broad term for a group of natural pigments found in most organisms. Eumelanin is produced through a multistage chemical process known as melanogenesis, where the oxidation of the amino ...
may play a role in extracting energy from ionizing radiation, such as gamma radiation. This form of "radiotrophic" growth has been described for only a few species, the effects on growth rates are small, and the underlying
biophysical
Biophysics is an interdisciplinary science that applies approaches and methods traditionally used in physics to study biological phenomena. Biophysics covers all scales of biological organization, from molecular to organismic and populations. Bi ...
and biochemical processes are not well known. This process might bear similarity to CO2 fixation via
visible light
Light or visible light is electromagnetic radiation that can be perceived by the human eye. Visible light is usually defined as having wavelengths in the range of 400–700 nanometres (nm), corresponding to frequencies of 750–420 te ...
, but instead uses ionizing radiation as a source of energy.
Reproduction
Fungal reproduction is complex, reflecting the differences in lifestyles and genetic makeup within this diverse kingdom of organisms. It is estimated that a third of all fungi reproduce using more than one method of propagation; for example, reproduction may occur in two well-differentiated stages within the
life cycle
Life cycle, life-cycle, or lifecycle may refer to:
Science and academia
*Biological life cycle, the sequence of life stages that an organism undergoes from birth to reproduction ending with the production of the offspring
* Life-cycle hypothesis ...
of a species, the
teleomorph
In mycology, the terms teleomorph, anamorph, and holomorph apply to portions of the life cycles of fungi in the phyla Ascomycota and Basidiomycota:
*Teleomorph: the sexual reproductive stage (morph), typically a fruiting body.
*Anamorph: an asex ...
(sexual reproduction) and the anamorph (asexual reproduction). Environmental conditions trigger genetically determined developmental states that lead to the creation of specialized structures for sexual or asexual reproduction. These structures aid reproduction by efficiently dispersing spores or spore-containing propagules.
Asexual reproduction
Asexual reproduction occurs via vegetative spores ( conidia) or through mycelial fragmentation. Mycelial fragmentation occurs when a fungal mycelium separates into pieces, and each component grows into a separate mycelium. Mycelial fragmentation and vegetative spores maintain clonal populations adapted to a specific niche, and allow more rapid dispersal than sexual reproduction. The "Fungi imperfecti" (fungi lacking the perfect or sexual stage) or Deuteromycota comprise all the species that lack an observable sexual cycle. Deuteromycota (alternatively known as Deuteromycetes, conidial fungi, or mitosporic fungi) is not an accepted taxonomic clade and is now taken to mean simply fungi that lack a known sexual stage.
Sexual reproduction
Sexual reproduction with
meiosis
Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately r ...
has been directly observed in all fungal phyla except Glomeromycota (genetic analysis suggests meiosis in Glomeromycota as well). It differs in many aspects from sexual reproduction in animals or plants. Differences also exist between fungal groups and can be used to discriminate species by morphological differences in sexual structures and reproductive strategies. Mating experiments between fungal isolates may identify species on the basis of biological species concepts. The major fungal groupings have initially been delineated based on the morphology of their sexual structures and spores; for example, the spore-containing structures, asci and
basidia
A basidium () is a microscopic sporangium (a spore-producing structure) found on the hymenophore of fruiting bodies of basidiomycete fungi which are also called tertiary mycelium, developed from secondary mycelium. Tertiary mycelium is highly- ...
, can be used in the identification of ascomycetes and basidiomycetes, respectively. Fungi employ two
mating system
A mating system is a way in which a group is structured in relation to sexual behaviour. The precise meaning depends upon the context. With respect to animals, the term describes which males and females mate under which circumstances. Recognised ...
s:
heterothallic
Heterothallic species have sexes that reside in different individuals. The term is applied particularly to distinguish heterothallic fungi, which require two compatible partners to produce sexual spores, from homothallic ones, which are capable ...
species allow mating only between individuals of the opposite
mating type
Mating types are the microorganism equivalent to sexes in multicellular lifeforms and are thought to be the ancestor to distinct sexes. They also occur in macro-organisms such as fungi.
Definition
Mating types are the microorganism equivalent to ...
, whereas
homothallic Homothallic refers to the possession, within a single organism, of the resources to reproduce sexually; i.e., having male and female reproductive structures on the same thallus. The opposite sexual functions are performed by different cells of a si ...
species can mate, and sexually reproduce, with any other individual or itself.
Most fungi have both a haploid and a diploid stage in their life cycles. In sexually reproducing fungi, compatible individuals may combine by fusing their hyphae together into an interconnected network; this process, anastomosis, is required for the initiation of the sexual cycle. Many ascomycetes and basidiomycetes go through a dikaryotic stage, in which the nuclei inherited from the two parents do not combine immediately after cell fusion, but remain separate in the hyphal cells (see heterokaryosis).
In ascomycetes, dikaryotic hyphae of the hymenium (the spore-bearing tissue layer) form a characteristic ''hook'' (crozier) at the hyphal septum. During
cell division
Cell division is the process by which a parent cell divides into two daughter cells. Cell division usually occurs as part of a larger cell cycle in which the cell grows and replicates its chromosome(s) before dividing. In eukaryotes, there ar ...
, the formation of the hook ensures proper distribution of the newly divided nuclei into the apical and basal hyphal compartments. An ascus (plural ''asci'') is then formed, in which karyogamy (nuclear fusion) occurs. Asci are embedded in an ascocarp, or fruiting body. Karyogamy in the asci is followed immediately by meiosis and the production of ascospores. After dispersal, the ascospores may germinate and form a new haploid mycelium.
Sexual reproduction in basidiomycetes is similar to that of the ascomycetes. Compatible haploid hyphae fuse to produce a dikaryotic mycelium. However, the dikaryotic phase is more extensive in the basidiomycetes, often also present in the vegetatively growing mycelium. A specialized anatomical structure, called a
clamp connection
A clamp connection is a hook-like structure formed by growing hyphal cells of certain fungi. It is a characteristic feature of Basidiomycetes fungi. It is created to ensure that each cell, or segment of hypha separated by septa (cross walls), rec ...
, is formed at each hyphal septum. As with the structurally similar hook in the ascomycetes, the clamp connection in the basidiomycetes is required for controlled transfer of nuclei during cell division, to maintain the dikaryotic stage with two genetically different nuclei in each hyphal compartment. A
basidiocarp
In fungi, a basidiocarp, basidiome, or basidioma () is the sporocarp of a basidiomycete, the multicellular structure on which the spore-producing hymenium is borne. Basidiocarps are characteristic of the hymenomycetes; rusts and smuts do ...
is formed in which club-like structures known as
basidia
A basidium () is a microscopic sporangium (a spore-producing structure) found on the hymenophore of fruiting bodies of basidiomycete fungi which are also called tertiary mycelium, developed from secondary mycelium. Tertiary mycelium is highly- ...
generate haploid basidiospores after karyogamy and meiosis. The most commonly known basidiocarps are mushrooms, but they may also take other forms (see
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 ...
section).
In fungi formerly classified as Zygomycota, haploid hyphae of two individuals fuse, forming a
gametangium
A gametangium (plural: gametangia) is an organ or cell in which gametes are produced that is found in many multicellular protists, algae, fungi, and the gametophytes of plants. In contrast to gametogenesis in animals, a gametangium is a haploid s ...
, a specialized cell structure that becomes a fertile
gamete
A gamete (; , ultimately ) is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. In species that produce ...
-producing cell. The gametangium develops into a
zygospore A zygospore is a diploid reproductive stage in the life cycle of many fungi and protists. Zygospores are created by the nuclear fusion of haploid cells. In fungi, zygospores are formed in zygosporangia after the fusion of specialized budding struc ...
, a thick-walled spore formed by the union of gametes. When the zygospore germinates, it undergoes
meiosis
Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately r ...
, generating new haploid hyphae, which may then form asexual sporangiospores. These sporangiospores allow the fungus to rapidly disperse and germinate into new genetically identical haploid fungal mycelia.
Spore dispersal
The spores of most of the researched species of fungi are transported by wind. Such species often produce dry or
hydrophobic
In chemistry, hydrophobicity is the physical property of a molecule that is seemingly repelled from a mass of water (known as a hydrophobe). In contrast, hydrophiles are attracted to water.
Hydrophobic molecules tend to be nonpolar and, t ...
spores that do not absorb water and are readily scattered by raindrops, for example. In other species, both asexual and sexual spores or sporangiospores are often actively dispersed by forcible ejection from their reproductive structures. This ejection ensures exit of the spores from the reproductive structures as well as traveling through the air over long distances. Specialized mechanical and physiological mechanisms, as well as spore surface structures (such as hydrophobins), enable efficient spore ejection. For example, the structure of the spore-bearing cells in some ascomycete species is such that the buildup of substances affecting cell volume and fluid balance enables the explosive discharge of spores into the air. The forcible discharge of single spores termed ''ballistospores'' involves formation of a small drop of water (Buller's drop), which upon contact with the spore leads to its projectile release with an initial acceleration of more than 10,000 g; the net result is that the spore is ejected 0.01–0.02cm, sufficient distance for it to fall through the
gills
A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they are ...
or pores into the air below. Other fungi, like the puffballs, rely on alternative mechanisms for spore release, such as external mechanical forces. The hydnoid fungi (tooth fungi) produce spores on pendant, tooth-like or spine-like projections. The bird's nest fungi use the force of falling water drops to liberate the spores from cup-shaped fruiting bodies. Another strategy is seen in the
stinkhorns
Phallaceae is a family of fungi, commonly known as stinkhorns, within the order Phallales. Stinkhorns have a worldwide distribution, but are especially prevalent in tropical regions. They are known for their foul-smelling, sticky spore masses, ...
, a group of fungi with lively colors and putrid odor that attract insects to disperse their spores.
Homothallism
In
homothallic Homothallic refers to the possession, within a single organism, of the resources to reproduce sexually; i.e., having male and female reproductive structures on the same thallus. The opposite sexual functions are performed by different cells of a si ...
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 th ...
, two haploid nuclei derived from the same individual fuse to form a
zygote
A zygote (, ) is a eukaryotic 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 individual organism.
In multicell ...
that can then undergo
meiosis
Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately r ...
. Homothallic fungi include species with an ''Aspergillus''-like asexual stage (anamorphs) occurring in numerous different genera, several species of the ascomycete genus '' Cochliobolus'', and the ascomycete '' Pneumocystis jirovecii''. The earliest mode of sexual reproduction among eukaryotes was likely homothallism, that is, self-fertile unisexual reproduction.
Other sexual processes
Besides regular sexual reproduction with meiosis, certain fungi, such as those in the genera ''
Penicillium
''Penicillium'' () is a genus of ascomycetous fungi that is part of the mycobiome of many species and is of major importance in the natural environment, in food spoilage, and in food and drug production.
Some members of the genus produce pe ...
'' and ''
Aspergillus
' () is a genus consisting of several hundred mold species found in various climates worldwide.
''Aspergillus'' was first catalogued in 1729 by the Italian priest and biologist Pier Antonio Micheli. Viewing the fungi under a microscope, Mic ...
'', may exchange genetic material via parasexual processes, initiated by anastomosis between hyphae and
plasmogamy
Plasmogamy is a stage in the sexual reproduction of fungi, in which the protoplasm of two parent cells (usually from the mycelia) fuse without the fusion of nuclei, effectively bringing two haploid nuclei close together in the same cell. This sta ...
of fungal cells. The frequency and relative importance of parasexual events is unclear and may be lower than other sexual processes. It is known to play a role in intraspecific hybridization and is likely required for hybridization between species, which has been associated with major events in fungal evolution.
Evolution
In contrast to
plants
Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae exclude ...
and
animals
Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...
, the early fossil record of the fungi is meager. Factors that likely contribute to the under-representation of fungal species among fossils include the nature of fungal
fruiting bodies
The sporocarp (also known as fruiting body, fruit body or fruitbody) of fungi is a multicellular structure on which spore-producing structures, such as basidia or asci, are borne. The fruitbody is part of the sexual phase of a fungal life cyc ...
, which are soft, fleshy, and easily degradable tissues and the microscopic dimensions of most fungal structures, which therefore are not readily evident. Fungal fossils are difficult to distinguish from those of other microbes, and are most easily identified when they resemble extant fungi. Often recovered from a permineralized plant or animal host, these samples are typically studied by making thin-section preparations that can be examined with light microscopy or transmission electron microscopy. Researchers study
compression fossil
A compression fossil is a fossil preserved in sedimentary rock that has undergone physical compression. While it is uncommon to find animals preserved as good compression fossils, it is very common to find plants preserved this way. The reason f ...
s by dissolving the surrounding matrix with acid and then using light or scanning electron microscopy to examine surface details.
The earliest fossils possessing features typical of fungi date to the
Paleoproterozoic
The Paleoproterozoic Era (;, also spelled Palaeoproterozoic), spanning the time period from (2.5–1.6 Ga), is the first of the three sub-divisions ( eras) of the Proterozoic Eon. The Paleoproterozoic is also the longest era of the Earth's ...
era, some ( Ma); these multicellular benthic organisms had filamentous structures capable of anastomosis. Other studies (2009) estimate the arrival of fungal organisms at about 760–1060Ma on the basis of comparisons of the rate of evolution in closely related groups. For much of the
Paleozoic
The Paleozoic (or Palaeozoic) Era is the earliest of three geologic eras of the Phanerozoic Eon.
The name ''Paleozoic'' ( ;) was coined by the British geologist Adam Sedgwick in 1838
by combining the Greek words ''palaiós'' (, "old") and ' ...
Era (542–251Ma), the fungi appear to have been aquatic and consisted of organisms similar to the extant chytrids in having flagellum-bearing spores. The evolutionary adaptation from an aquatic to a terrestrial lifestyle necessitated a diversification of ecological strategies for obtaining nutrients, including parasitism, saprobism, and the development of mutualistic relationships such as mycorrhiza and lichenization. Studies suggest that the ancestral ecological state of the Ascomycota was saprobism, and that independent lichenization events have occurred multiple times.
In May 2019, scientists reported the discovery of a
fossil
A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
plant
Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae exclu ...
microfossil
A microfossil is a fossil that is generally between 0.001 mm and 1 mm in size, the visual study of which requires the use of light or electron microscopy. A fossil which can be studied with the naked eye or low-powered magnification, ...
s preserved in the basal Ediacaran Doushantuo Formation (~635 Ma) have been reported in South China. Earlier, it had been presumed that the fungi colonized the land during the Cambrian (542–488.3Ma), also long before land plants. Fossilized hyphae and spores recovered from the
Ordovician
The Ordovician ( ) is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start of the Silurian Period Mya.
T ...
of Wisconsin (460Ma) resemble modern-day
Glomerales
Glomerales is an order of symbiotic fungi within the phylum Glomeromycota.
Biology
These fungi are all biotrophic mutualists. Most employ the arbuscular mycorrhizal method of nutrient exchange with plants. They produce large (.1-.5mm) spor ...
, and existed at a time when the land flora likely consisted of only non-vascular
bryophyte
The Bryophyta s.l. are a proposed taxonomic division containing three groups of non-vascular land plants (embryophytes): the liverworts, hornworts and mosses. Bryophyta s.s. consists of the mosses only. They are characteristically limited in s ...
-like plants.Prototaxites, which was probably a fungus or lichen, would have been the tallest organism of the late Silurian and early Devonian. Fungal fossils do not become common and uncontroversial until the early Devonian (416–359.2Ma), when they occur abundantly in the Rhynie chert, mostly as Zygomycota and
Chytridiomycota
Chytridiomycota are a division of zoosporic organisms in the kingdom Fungi, informally known as chytrids. The name is derived from the Ancient Greek ('), meaning "little pot", describing the structure containing unreleased zoöspores. Chytri ...
. At about this same time, approximately 400Ma, the Ascomycota and Basidiomycota diverged, and all modern classes of fungi were present by the Late Carboniferous ( Pennsylvanian, 318.1–299Ma).
Lichens formed a component of the early terrestrial ecosystems, and the estimated age of the oldest terrestrial lichen fossil is 415Ma; this date roughly corresponds to the age of the oldest known sporocarp fossil, a '' Paleopyrenomycites'' species found in the Rhynie Chert. The oldest fossil with microscopic features resembling modern-day basidiomycetes is ''Palaeoancistrus'', found permineralized with a
fern
A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes exce ...
from the Pennsylvanian. Rare in the fossil record are the Homobasidiomycetes (a
taxon
In biology, a taxon ( back-formation from '' taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular n ...
roughly equivalent to the mushroom-producing species of the Agaricomycetes). Two
amber
Amber is fossilized tree resin that has been appreciated for its color and natural beauty since Neolithic times. Much valued from antiquity to the present as a gemstone, amber is made into a variety of decorative objects."Amber" (2004). In ...
-preserved specimens provide evidence that the earliest known mushroom-forming fungi (the extinct species '' Archaeomarasmius leggetti'') appeared during the late
Cretaceous
The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
, 90Ma.
Some time after the Permian–Triassic extinction event (251.4Ma), a fungal spike (originally thought to be an extraordinary abundance of fungal spores in
sediment
Sediment is a naturally occurring material that is broken down by processes of weathering and erosion, and is subsequently transported by the action of wind, water, or ice or by the force of gravity acting on the particles. For example, sa ...
s) formed, suggesting that fungi were the dominant life form at this time, representing nearly 100% of the available
fossil record
A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
for this period. However, the relative proportion of fungal spores relative to spores formed by algal species is difficult to assess, the spike did not appear worldwide, and in many places it did not fall on the Permian–Triassic boundary.
Sixty-five million years ago, immediately after the
Cretaceous–Paleogene extinction event
The Cretaceous–Paleogene (K–Pg) extinction event (also known as the Cretaceous–Tertiary extinction) was a sudden mass extinction of three-quarters of the plant and animal species on Earth, approximately 66 million years ago. With the ...
that famously killed off most dinosaurs, there was a dramatic increase in evidence of fungi; apparently the death of most plant and animal species led to a huge fungal bloom like "a massive compost heap".
Taxonomy
Although commonly included in botany curricula and textbooks, fungi are more closely related to
animal
Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...
s than to plants and are placed with the animals in the monophyletic group of
opisthokont
The opisthokonts () are a broad group of eukaryotes, including both the animal and fungus kingdoms. The opisthokonts, previously called the "Fungi/Metazoa group", are generally recognized as a clade. Opisthokonts together with Apusomonadida and ...
s. Analyses using
molecular phylogenetics
Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
taxonomy
Taxonomy is the practice and science of categorization or classification.
A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
of fungi is in a state of constant flux, especially due to research based on DNA comparisons. These current phylogenetic analyses often overturn classifications based on older and sometimes less discriminative methods based on morphological features and biological species concepts obtained from experimental
mating
In biology, mating is the pairing of either opposite- sex or hermaphroditic organisms for the purposes of sexual reproduction. ''Fertilization'' is the fusion of two gametes. ''Copulation'' is the union of the sex organs of two sexually reprod ...
s.
There is no unique generally accepted system at the higher taxonomic levels and there are frequent name changes at every level, from species upwards. Efforts among researchers are now underway to establish and encourage usage of a unified and more consistent
nomenclature
Nomenclature (, ) is a system of names or terms, or the rules for forming these terms in a particular field of arts or sciences. The principles of naming vary from the relatively informal conventions of everyday speech to the internationally ag ...
. Until relatively recent (2012) changes to the International Code of Nomenclature for algae, fungi and plants, fungal species could also have multiple scientific names depending on their life cycle and mode (sexual or asexual) of reproduction. Web sites such as
Index Fungorum
''Index Fungorum'' is an international project to index all formal names (scientific names) in the fungus kingdom. the project is based at the Royal Botanic Gardens, Kew, one of three partners along with Landcare Research and the Institute of M ...
and
MycoBank
MycoBank is an online database, documenting new mycological names and combinations, eventually combined with descriptions and illustrations. It is run by the Westerdijk Fungal Biodiversity Institute in Utrecht.
Each novelty, after being screened ...
are officially recognized nomenclatural repositories and list current names of fungal species (with cross-references to older synonyms).
The 2007 classification of Kingdom Fungi is the result of a large-scale collaborative research effort involving dozens of mycologists and other scientists working on fungal taxonomy. It recognizes seven phyla, two of which—the Ascomycota and the Basidiomycota—are contained within a branch representing
subkingdom
In biology, a kingdom is the second highest taxonomic rank, just below domain. Kingdoms are divided into smaller groups called phyla.
Traditionally, some textbooks from the United States and Canada used a system of six kingdoms (Animalia, Plant ...
Dikarya, the most species rich and familiar group, including all the mushrooms, most food-spoilage molds, most plant pathogenic fungi, and the beer, wine, and bread yeasts. The accompanying
cladogram
A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
depicts the major fungal
taxa
In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
and their relationship to opisthokont and unikont organisms, based on the work of Philippe Silar, "The Mycota: A Comprehensive Treatise on Fungi as Experimental Systems for Basic and Applied Research" and Tedersoo et al. 2018. The lengths of the branches are not proportional to evolutionary distances.
Taxonomic groups
The major phyla (sometimes called divisions) of fungi have been classified mainly on the basis of characteristics of their sexual reproductive structures. , nine major lineages have been identified: Opisthosporidia, Chytridiomycota, Neocallimastigomycota, Blastocladiomycota, Zoopagomycota, Mucoromycota, Glomeromycota, Ascomycota and Basidiomycota.
Phylogenetic analysis has demonstrated that the Microsporidia, unicellular parasites of animals and protists, are fairly recent and highly derived
toxic
Toxicity is the degree to which a chemical substance or a particular mixture of substances can damage an organism. Toxicity can refer to the effect on a whole organism, such as an animal, bacterium, or plant, as well as the effect on a subs ...
to animals or plants and are therefore called mycotoxins. Of particular relevance to humans are mycotoxins produced by molds causing food spoilage, and poisonous mushrooms (see above). Particularly infamous are the lethal
amatoxin Amatoxin is the collective name of a subgroup of at least nine related toxic compounds found in three genera of poisonous mushrooms (''Amanita'', '' Galerina'' and '' Lepiota'') and one species ( Conocybe filaris) of the genus '' Conocybe''. Amatoxi ...
s in some ''
Amanita
The genus ''Amanita'' contains about 600 species of agarics, including some of the most toxic known mushrooms found worldwide, as well as some well-regarded edible species. This genus is responsible for approximately 95% of the fatalities resul ...
'' mushrooms, and
ergot alkaloids
Ergot ( ) or ergot fungi refers to a group of fungi of the genus ''Claviceps''.
The most prominent member of this group is ''Claviceps purpurea'' ("rye ergot fungus"). This fungus grows on rye and related plants, and produces alkaloids that ca ...
, which have a long history of causing serious epidemics of ergotism (St Anthony's Fire) in people consuming rye or related cereals contaminated with
sclerotia
A sclerotium (; (), is a compact mass of hardened fungal mycelium containing food reserves. One role of sclerotia is to survive environmental extremes. In some higher fungi such as ergot, sclerotia become detached and remain dormant until favor ...
of the ergot fungus, ''
Claviceps purpurea
''Claviceps purpurea'' is an ergot fungus that grows on the ears of rye and related cereal and forage plants. Consumption of grains or seeds contaminated with the survival structure of this fungus, the ergot sclerotium, can cause ergotism in h ...
''. Other notable mycotoxins include the
aflatoxin
Aflatoxins are various poisonous carcinogens and mutagens that are produced by certain molds, particularly ''Aspergillus'' species. The fungi grow in soil, decaying vegetation and various staple foodstuffs and commodities such as hay, sweetcorn ...
Aspergillus
' () is a genus consisting of several hundred mold species found in various climates worldwide.
''Aspergillus'' was first catalogued in 1729 by the Italian priest and biologist Pier Antonio Micheli. Viewing the fungi under a microscope, Mic ...
'' species often growing in or on grains and nuts consumed by humans, ochratoxins,
patulin
Patulin is an organic compound classified as a polyketide. It is a white powder soluble in acidic water and in organic solvents. It is a lactone that is heat-stable, so it is not destroyed by pasteurization or thermal denaturation.http://www.s ...
T-2 mycotoxin
T-2 Mycotoxin (pronounced as 'Tee-Two') is a trichothecene mycotoxin. It is a naturally occurring mold byproduct of ''Fusarium'' spp. fungus which is toxic to humans and animals. The clinical condition it causes is ''alimentary toxic aleukia'' ...
) and
fumonisin The fumonisins are a group of mycotoxins derived from ''Fusarium'' and their Liseola section. They have strong structural similarity to sphinganine, the backbone precursor of sphingolipids.
More specifically, it can refer to:
* Fumonisin B1
* Fum ...
s, which have significant impact on human food supplies or animal
livestock
Livestock are the domesticated animals raised in an agricultural setting to provide labor and produce diversified products for consumption such as meat, eggs, milk, fur, leather, and wool. The term is sometimes used to refer solely to animal ...
.
Mycotoxins are secondary metabolites (or natural products), and research has established the existence of biochemical pathways solely for the purpose of producing mycotoxins and other natural products in fungi. Mycotoxins may provide fitness benefits in terms of physiological adaptation, competition with other microbes and fungi, and protection from consumption ( fungivory). Many fungal secondary metabolites (or derivatives) are used medically, as described under Human use below.
Pathogenic mechanisms
'' Ustilago maydis'' is a pathogenic plant fungus that causes smut disease in maize and teosinte. Plants have evolved efficient defense systems against pathogenic microbes such as ''U. maydis''. A rapid defense reaction after pathogen attack is the oxidative burst where the plant produces
reactive oxygen species
In chemistry, reactive oxygen species (ROS) are highly reactive chemicals formed from diatomic oxygen (). Examples of ROS include peroxides, superoxide, hydroxyl radical, singlet oxygen, and alpha-oxygen.
The reduction of molecular oxygen () p ...
at the site of the attempted invasion. ''U. maydis'' can respond to the oxidative burst with an oxidative stress response, regulated by the gene ''
YAP1
YAP1 (yes-associated protein 1), also known as YAP or YAP65, is a protein that acts as a transcription coregulator that promotes transcription of genes involved in cellular proliferation and suppressing apoptotic genes. YAP1 is a component in th ...
''. The response protects ''U. maydis'' from the host defense, and is necessary for the pathogen's virulence. Furthermore, ''U. maydis'' has a well-established recombinational
DNA repair
DNA repair is a collection of processes by which a cell identifies and corrects damage to the DNA molecules that encode its genome. In human cells, both normal metabolic activities and environmental factors such as radiation can cause DNA da ...
system which acts during mitosis and meiosis. The system may assist the pathogen in surviving DNA damage arising from the host plant's oxidative defensive response to infection.
'' Cryptococcus neoformans'' is an encapsulated yeast that can live in both plants and animals. ''C.neoformans'' usually infects the lungs, where it is phagocytosed by
alveolar macrophage
An alveolar macrophage, pulmonary macrophage, (or dust cell) is a type of macrophage, a professional phagocyte, found in the airways and at the level of the alveoli in the lungs, but separated from their walls.
Activity of the alveolar macropha ...
s. Some ''C.neoformans'' can survive
inside
Inside may refer to:
* Insider, a member of any group of people of limited number and generally restricted access
Film
* ''Inside'' (1996 film), an American television film directed by Arthur Penn and starring Eric Stoltz
* ''Inside'' (2002 f ...
macrophages, which appears to be the basis for latency, disseminated disease, and resistance to antifungal agents. One mechanism by which ''C.neoformans'' survives the hostile macrophage environment is by up-regulating the expression of genes involved in the oxidative stress response. Another mechanism involves
meiosis
Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately r ...
. The majority of ''C.neoformans'' are mating "type a". Filaments of mating "type a" ordinarily have haploid nuclei, but they can become diploid (perhaps by endoduplication or by stimulated nuclear fusion) to form
blastospore
A blastospore is an asexual fungal spore produced by budding. Produced by fungi within the phylum Glomeromycota and others. It is also known as a blastoconidium (plural = blastoconidia).
An example of a fungus that forms blastospores is ''Ca ...
s. The diploid nuclei of blastospores can undergo meiosis, including recombination, to form haploid basidiospores that can be dispersed. This process is referred to as monokaryotic fruiting. This process requires a gene called ''
DMC1
Meiotic recombination protein DMC1/LIM15 homolog is a protein that in humans is encoded by the ''DMC1'' gene.
Meiotic recombination protein Dmc1 is a homolog of the bacterial strand exchange protein RecA. Dmc1 plays the central role in homologous ...
'', which is a conserved homologue of genes ''
recA
RecA is a 38 kilodalton protein essential for the repair and maintenance of DNA. A RecA structural and functional homolog has been found in every species in which one has been seriously sought and serves as an archetype for this class of homolog ...
'' in bacteria and '' RAD51'' in eukaryotes, that mediates homologous chromosome pairing during meiosis and repair of DNA double-strand breaks. Thus, ''C.neoformans'' can undergo a meiosis, monokaryotic fruiting, that promotes recombinational repair in the oxidative, DNA damaging environment of the host macrophage, and the repair capability may contribute to its virulence.
Human use
The human use of fungi for food preparation or preservation and other purposes is extensive and has a long history.
Mushroom farming
Fungiculture is the cultivation of fungi such as mushrooms. Cultivating fungi can yield foods (which include mostly mushrooms), medicine, construction materials and other products. A ''mushroom farm'' is involved in the business of growing fun ...
and
mushroom gathering
Mushroom hunting, mushrooming, mushroom picking, mushroom foraging, and similar terms describe the activity of gathering mushrooms in the wild, typically for culinary use. This practice is popular throughout most of Europe, Australia, Japan, ...
are large industries in many countries. The study of the historical uses and sociological impact of fungi is known as
ethnomycology
Ethnomycology is the study of the historical uses and sociological impact of fungi and can be considered a subfield of ethnobotany or ethnobiology. Although in theory the term includes fungi used for such purposes as tinder, medicine ( medicinal ...
. Because of the capacity of this group to produce an enormous range of
natural products
A natural product is a natural Chemical compound, compound or chemical substance, substance produced by a living organism—that is, found in nature. In the broadest sense, natural products include any substance produced by life. Natural product ...
with
antimicrobial
An antimicrobial is an agent that kills microorganisms or stops their growth. Antimicrobial medicines can be grouped according to the microorganisms they act primarily against. For example, antibiotics are used against bacteria, and antifungals ar ...
or other biological activities, many species have long been used or are being developed for industrial
production of antibiotics
Production of antibiotics is a naturally occurring event, that thanks to advances in science can now be replicated and improved upon in laboratory settings. Due to the discovery of penicillin by Alexander Flemming, and the efforts of Florey and ...
, vitamins, and
anti-cancer
Cancer is a group of diseases involving abnormal cell growth with the potential to invade or spread to other parts of the body. These contrast with benign tumors, which do not spread. Possible signs and symptoms include a lump, abnormal ble ...
genetic engineering
Genetic engineering, also called genetic modification or genetic manipulation, is the modification and manipulation of an organism's genes using technology. It is a set of technologies used to change the genetic makeup of cells, including t ...
of fungi, enabling
metabolic engineering
Metabolic engineering is the practice of optimizing genetic and regulatory processes within cells to increase the cell's production of a certain substance. These processes are chemical networks that use a series of biochemical reactions and enz ...
of fungal species. For example, genetic modification of yeast species—which are easy to grow at fast rates in large fermentation vessels—has opened up ways of
pharmaceutical
A medication (also called medicament, medicine, pharmaceutical drug, medicinal drug or simply drug) is a drug used to diagnose, cure, treat, or prevent disease. Drug therapy (pharmacotherapy) is an important part of the medical field and re ...
production that are potentially more efficient than production by the original source organisms. Fungi-based industries are sometimes considered to be a major part of a growing
bioeconomy
Biobased economy, bioeconomy or biotechonomy is economic activity involving the use of biotechnology and biomass in the production of goods, services, or energy. The terms are widely used by regional development agencies, national and international ...
, with applications under
research and development
Research and development (R&D or R+D), known in Europe as research and technological development (RTD), is the set of innovative activities undertaken by corporations or governments in developing new services or products, and improving existi ...
including use for textiles,
meat
Meat is animal flesh that is eaten as food. Humans have hunted, farmed, and scavenged animals for meat since prehistoric times. The establishment of settlements in the Neolithic Revolution allowed the domestication of animals such as chic ...
substitution and general fungal biotechnology.
Therapeutic uses
Modern chemotherapeutics
Many species produce metabolites that are major sources of pharmacologically active drugs.
= Antibiotics
=
Particularly important are the antibiotics, including the
penicillin
Penicillins (P, PCN or PEN) are a group of β-lactam antibiotics originally obtained from ''Penicillium'' moulds, principally '' P. chrysogenum'' and '' P. rubens''. Most penicillins in clinical use are synthesised by P. chrysogenum using ...
s, a structurally related group of
β-lactam antibiotic
β-lactam antibiotics (beta-lactam antibiotics) are antibiotics that contain a beta-lactam ring in their chemical
structure. This includes penicillin derivatives (penams), cephalosporins and cephamycins (cephems), monobactams, carbapenems and c ...
s that are synthesized from small
peptide
Peptides (, ) are short chains of amino acids linked by peptide bonds. Long chains of amino acids are called proteins. Chains of fewer than twenty amino acids are called oligopeptides, and include dipeptides, tripeptides, and tetrapeptides.
A ...
s. Although naturally occurring penicillins such as
penicillin G
Benzylpenicillin, also known as penicillin G (PenG) or BENPEN, and in military slang "Peanut Butter Shot" is an antibiotic used to treat a number of bacterial infections. This includes pneumonia, strep throat, syphilis, necrotizing enteroco ...
(produced by ''
Penicillium chrysogenum
''Penicillium chrysogenum'' (formerly known as ''Penicillium notatum'') is a species of fungus in the genus ''Penicillium''. It is common in temperate and subtropical regions and can be found on salted food products, but it is mostly found in in ...
'') have a relatively narrow spectrum of biological activity, a wide range of other penicillins can be produced by
chemical modification
Chemical modification refers to a number of various processes involving the alteration of the chemical constitution or structure of molecules.
In chemistry
Chemical modification describes the conversion of macromolecules through a chemical react ...
of the natural penicillins. Modern penicillins are
semisynthetic
Semisynthesis, or partial chemical synthesis, is a type of chemical synthesis that uses chemical compounds isolated from natural sources (such as microbial cell cultures or plant material) as the starting materials to produce novel compounds with ...
compounds, obtained initially from
fermentation
Fermentation is a metabolic process that produces chemical changes in organic substrates through the action of enzymes. In biochemistry, it is narrowly defined as the extraction of energy from carbohydrates in the absence of oxygen. In food ...
cultures, but then structurally altered for specific desirable properties. Other antibiotics produced by fungi include:
ciclosporin
Ciclosporin, also spelled cyclosporine and cyclosporin, is a calcineurin inhibitor, used as an immunosuppressant medication. It is a natural product. It is taken orally or intravenously for rheumatoid arthritis, psoriasis, Crohn's disease, ...
, commonly used as an
immunosuppressant
Immunosuppressive drugs, also known as immunosuppressive agents, immunosuppressants and antirejection medications, are drugs that inhibit or prevent activity of the immune system.
Classification
Immunosuppressive drugs can be classified in ...
during
transplant surgery
Organ transplantation is a medical procedure in which an organ is removed from one body and placed in the body of a recipient, to replace a damaged or missing organ. The donor and recipient may be at the same location, or organs may be transpor ...
tuberculosis
Tuberculosis (TB) is an infectious disease usually caused by '' Mycobacterium tuberculosis'' (MTB) bacteria. Tuberculosis generally affects the lungs, but it can also affect other parts of the body. Most infections show no symptoms, in ...
,
syphilis
Syphilis () is a sexually transmitted infection caused by the bacterium ''Treponema pallidum'' subspecies ''pallidum''. The signs and symptoms of syphilis vary depending in which of the four stages it presents (primary, secondary, latent, an ...
,
leprosy
Leprosy, also known as Hansen's disease (HD), is a long-term infection by the bacteria ''Mycobacterium leprae'' or ''Mycobacterium lepromatosis''. Infection can lead to damage of the nerves, respiratory tract, skin, and eyes. This nerve damag ...
, and others began in the early 20th century and continues to date. In nature, antibiotics of fungal or bacterial origin appear to play a dual role: at high concentrations they act as chemical defense against competition with other microorganisms in species-rich environments, such as the
rhizosphere
The rhizosphere is the narrow region of soil or substrate that is directly influenced by root secretions and associated soil microorganisms known as the root microbiome. Soil pores in the rhizosphere can contain many bacteria and other microor ...
, and at low concentrations as
quorum-sensing
In biology, quorum sensing or quorum signalling (QS) is the ability to detect and respond to cell population density by gene regulation. As one example, QS enables bacteria to restrict the expression of specific genes to the high cell densities at ...
molecules for intra- or interspecies signaling.
= Other
=
Other drugs produced by fungi include
griseofulvin
Griseofulvin is an antifungal medication used to treat a number of types of dermatophytoses (ringworm). This includes fungal infections of the nails and scalp, as well as the skin when antifungal creams have not worked. It is taken by mouth.
C ...
isolated from ''
Penicillium griseofulvum
''Penicillium griseofulvum'' is a species of the genus of ''Penicillium'' which produces patulin, penifulvin A, cyclopiazonic acid, roquefortine C, shikimic acid, griseofulvin, and 6-Methylsalicylic acid (via a polyketide synthase
Polyketides ...
'', used to treat fungal infections, and
statin
Statins, also known as HMG-CoA reductase inhibitors, are a class of lipid-lowering medications that reduce illness and mortality in those who are at high risk of cardiovascular disease. They are the most common cholesterol-lowering drugs.
Low- ...
cholesterol synthesis
The mevalonate pathway, also known as the isoprenoid pathway or HMG-CoA reductase pathway is an essential metabolic pathway present in eukaryotes, archaea, and some bacteria. The pathway produces two five-carbon building blocks called isopenten ...
. Examples of statins found in fungi include
mevastatin
Mevastatin (compactin, ML-236B) is a hypolipidemic agent that belongs to the statins class.
It was isolated from the mold ''Penicillium citrinum'' by Akira Endo in the 1970s, and he identified it as a HMG-CoA reductase inhibitor, i.e., a statin. ...
from ''
Penicillium citrinum
''Penicillium citrinum'' is an anamorph, mesophilic fungus species of the genus of ''Penicillium'' which produces tanzawaic acid A-D, ACC, Mevastatin, Quinocitrinine A, Quinocitrinine B, and nephrotoxic citrinin. ''Penicillium citrinum'' ...
'' and
lovastatin
Lovastatin, sold under the brand name Mevacor among others, is a statin medication, to treat hypercholesterolemia, high blood cholesterol and reduce the risk of cardiovascular disease. Its use is recommended together with lifestyle changes. It ...
from ''
Aspergillus terreus
''Aspergillus terreus'', also known as ''Aspergillus terrestris'', is a fungus (mold) found worldwide in soil. Although thought to be strictly asexual until recently, ''A. terreus'' is now known to be capable of sexual reproduction. This saprotr ...
'' and the
oyster mushroom
''Pleurotus'' is a genus of gilled mushrooms which includes one of the most widely eaten mushrooms, '' P. ostreatus''. Species of ''Pleurotus'' may be called oyster, abalone, or tree mushrooms, and are some of the most commonly cultivated edib ...
.
Psilocybin
Psilocybin ( , ) is a naturally occurring psychedelic prodrug compound produced by more than 200 species of fungi. The most potent are members of the genus ''Psilocybe'', such as '' P. azurescens'', '' P. semilanceata'', and '' P.&nbs ...
from
fungi
A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...
network
Network, networking and networked may refer to:
Science and technology
* Network theory, the study of graphs as a representation of relations between discrete objects
* Network science, an academic field that studies complex networks
Mathematics
...
integration
Integration may refer to:
Biology
*Multisensory integration
*Path integration
* Pre-integration complex, viral genetic material used to insert a viral genome into a host genome
*DNA integration, by means of site-specific recombinase technology, ...
. Fungi produce compounds that inhibit
virus
A virus is a submicroscopic infectious agent that replicates only inside the living cells of an organism. Viruses infect all life forms, from animals and plants to microorganisms, including bacteria and archaea.
Since Dmitri Ivanovsky's 1 ...
es and cancer cells. Specific metabolites, such as
polysaccharide-K
Polysaccharide-K (Krestin, PSK) is a protein-bound polysaccharide isolated from the fruitbody of ''Trametes versicolor''.
Research summary
PSK has been studied in patients with gastric cancer, breast cancer, colorectal cancer, and lung cancer. T ...
,
ergotamine
Ergotamine, sold under the brand names Cafergot (with caffeine) and Ergomar among others, is an ergopeptine and part of the ergot family of alkaloids; it is structurally and biochemically closely related to ergoline. It possesses structural simil ...
, and
β-lactam antibiotics
β-lactam antibiotics (beta-lactam antibiotics) are antibiotics that contain a beta-lactam ring in their chemical
structure. This includes penicillin derivatives (penams), cephalosporins and cephamycins (cephems), monobactams, carbapenems and ...
, are routinely used in clinical medicine. The
shiitake
The shiitake (alternate form shitake) (; ''Lentinula edodes'') is an edible mushroom native to East Asia, which is now cultivated and consumed around the globe. It is considered a Medicinal fungi, medicinal mushroom in some forms of tradition ...
mushroom is a source of
lentinan
Lentinan is a polysaccharide isolated from the fruit body of shiitake mushroom (''Lentinula edodes'' mycelium).
Chemistry
Lentinan is a β-1,3 beta-glucan with β-1,6 branching. It has a molecular weight of 500,000 Da and specific rotation of +1 ...
, a clinical drug approved for use in cancer treatments in several countries, including
Japan
Japan ( ja, 日本, or , and formally , ''Nihonkoku'') is an island country in East Asia. It is situated in the northwest Pacific Ocean, and is bordered on the west by the Sea of Japan, while extending from the Sea of Okhotsk in the north ...
. In Europe and Japan,
polysaccharide-K
Polysaccharide-K (Krestin, PSK) is a protein-bound polysaccharide isolated from the fruitbody of ''Trametes versicolor''.
Research summary
PSK has been studied in patients with gastric cancer, breast cancer, colorectal cancer, and lung cancer. T ...
(brand name Krestin), a chemical derived from ''
Trametes versicolor
''Trametes versicolor''also known as ''Coriolus versicolor'' and ''Polyporus versicolor''is a common polypore mushroom found throughout the world. Meaning 'of several colors', ''versicolor'' reliably describes this fungus that displays a varie ...
'', is an approved
adjuvant In pharmacology, an adjuvant is a drug or other substance, or a combination of substances, that is used to increase the efficacy or potency of certain drugs. Specifically, the term can refer to:
* Adjuvant therapy in cancer management
* Analgesic ...
traditional Chinese medicine
Traditional Chinese medicine (TCM) is an alternative medical practice drawn from traditional medicine in China. It has been described as "fraught with pseudoscience", with the majority of its treatments having no logical mechanism of action ...
. Mushrooms with a history of such use include ''
Agaricus subrufescens
''Agaricus subrufescens'' (syn. ''Agaricus blazei'', ''Agaricus brasiliensis'' or ''Agaricus rufotegulis'') is a species of mushroom, commonly known as almond mushroom, mushroom of the sun, God's mushroom, mushroom of life, royal sun agaricus, '' ...
Ophiocordyceps sinensis
''Ophiocordyceps sinensis'' (formerly known as ''Cordyceps sinensis''), known colloquially as caterpillar fungus, is an entomopathogenic fungus (a fungus that grows on insects) in the family Ophiocordycipitaceae. It is mainly found in the mead ...
''.
Cultured foods
Baker's yeast
Baker's yeast is the common name for the strains of yeast commonly used in baking bread and other bakery products, serving as a leavening agent which causes the bread to rise (expand and become lighter and softer) by converting the fermentable ...
or ''
Saccharomyces cerevisiae
''Saccharomyces cerevisiae'' () (brewer's yeast or baker's yeast) is a species of yeast (single-celled fungus microorganisms). The species has been instrumental in winemaking, baking, and brewing since ancient times. It is believed to have been o ...
'', a unicellular fungus, is used to make
bread
Bread is a staple food prepared from a dough of flour (usually wheat) and water, usually by baking. Throughout recorded history and around the world, it has been an important part of many cultures' diet. It is one of the oldest human-made f ...
and other wheat-based products, such as
pizza
Pizza (, ) is a dish of Italian origin consisting of a usually round, flat base of leavened wheat-based dough topped with tomatoes, cheese, and often various other ingredients (such as various types of sausage, anchovies, mushrooms, onions ...
dough and
dumpling
Dumpling is a broad class of dishes that consist of pieces of dough (made from a variety of starch sources), oftentimes wrapped around a filling. The dough can be based on bread, flour, buckwheat or potatoes, and may be filled with meat, fi ...
s. Yeast species of the genus '' Saccharomyces'' are also used to produce
alcoholic beverage
An alcoholic beverage (also called an alcoholic drink, adult beverage, or a drink) is a drink that contains ethanol, a type of alcohol that acts as a drug and is produced by fermentation of grains, fruits, or other sources of sugar. The c ...
s through fermentation. Shoyu koji mold (''
Aspergillus oryzae
''Aspergillus oryzae'', also known as , is a filamentous fungus (a mold) used in East Asia to saccharify rice, sweet potato, and barley in the making of alcoholic beverages such as ''sake'' and '' shōchū'', and also to ferment soybeans for m ...
'') is an essential ingredient in brewing
Shoyu
Soy sauce (also called simply soy in American English and soya sauce in British English) is a liquid condiment of Chinese origin, traditionally made from a fermented paste of soybeans, roasted grain, brine, and '' Aspergillus oryzae'' or '' A ...
sake
Sake, also spelled saké ( ; also referred to as Japanese rice wine), is an alcoholic beverage of Japanese origin made by fermenting rice that has been polished to remove the bran. Despite the name ''Japanese rice wine'', sake, and indee ...
, and the preparation of
miso
is a traditional Japanese seasoning. It is a thick paste produced by fermenting soybeans with salt and ''kōji'' (the fungus '' Aspergillus oryzae'') and sometimes rice, barley, seaweed, or other ingredients. It is used for sauces and spre ...
, while ''
Rhizopus
''Rhizopus'' is a genus of common saprophytic fungi on plants and specialized parasites on animals. They are found in a wide variety of organic substances, including "mature fruits and vegetables", jellies, syrups, leather, bread, peanuts, and ...
'' species are used for making
tempeh
Tempeh or tempe (; jv, ꦠꦺꦩ꧀ꦥꦺ, témpé, ) is a traditional Indonesian food made from fermented soybeans. It is made by a natural culturing and controlled fermentation process that binds soybeans into a cake form. A fungus, ''Rhizopus ...
. Several of these fungi are
domesticated
Domestication is a sustained multi-generational relationship in which humans assume a significant degree of control over the reproduction and care of another group of organisms to secure a more predictable supply of resources from that group. A ...
species that were bred or selected according to their capacity to ferment food without producing harmful mycotoxins (see below), which are produced by very closely related '' Aspergilli''.
Quorn
Quorn is a brand of meat substitute products, or the company that makes them. Quorn originated in the UK and is sold primarily in Europe, but is available in 14 countries. The brand is owned by parent company Monde Nissin.
Quorn is sold as bo ...
, a
meat substitute
A meat alternative or meat substitute (also called plant-based meat or fake meat, sometimes pejoratively) is a food product made from vegetarian or vegan ingredients, eaten as a replacement for meat. Meat alternatives typically approximate qua ...
, is made from ''
Fusarium venenatum
''Fusarium venenatum'' is a microfungus of the genus ''Fusarium'' that has a high protein content. One of its strains is used commercially for the production of the single cell protein mycoprotein Quorn.
''Fusarium venenatum'' was discovered ...
''.
In food
Edible mushroom
Edible mushrooms are the fleshy and edible fruit bodies of several species of macrofungi (fungi which bear fruiting structures that are large enough to be seen with the naked eye). They can appear either below ground (hypogeous) or above ground ...
s include commercially raised and wild-harvested fungi. ''
Agaricus bisporus
''Agaricus bisporus'' is an edible basidiomycete mushroom native to grasslands in Eurasia and North America. It has two color states while immature – white and brown – both of which have various names, with additional names for the mature ...
'', sold as button mushrooms when small or Portobello mushrooms when larger, is the most widely cultivated species in the West, used in salads, soups, and many other dishes. Many Asian fungi are commercially grown and have increased in popularity in the West. They are often available fresh in
grocery store
A grocery store ( AE), grocery shop ( BE) or simply grocery is a store that primarily retails a general range of food products, which may be fresh or packaged. In everyday U.S. usage, however, "grocery store" is a synonym for supermarket, a ...
s and markets, including straw mushrooms (''
Volvariella volvacea
''Volvariella volvacea'' (also known as paddy straw mushroom or straw mushroom) is a species of edible mushroom cultivated throughout East and Southeast Asia and used extensively in Asian cuisines. They are often available fresh in regions they ...
''), oyster mushrooms (''
Pleurotus ostreatus
''Pleurotus ostreatus'', the oyster mushroom, oyster fungus, or hiratake, is a common edible mushroom. It was first cultivated in Germany as a subsistence measure during World War I and is now grown commercially around the world for food. It is ...
''), shiitakes (''
Lentinula edodes
The shiitake (alternate form shitake) (; ''Lentinula edodes'') is an edible mushroom native to East Asia, which is now cultivated and consumed around the globe. It is considered a medicinal mushroom in some forms of traditional medicine.
Ta ...
''), and
enokitake
''Flammulina filiformis'' is a species of agaric (gilled mushroom) in the family Physalacriaceae. It is well known for its role in Japanese cuisine, where it is called ''enokitake'' (榎茸, エノキタケ, )
and is widely cultivated in East A ...
(''
Flammulina
''Flammulina'' is a genus of fungi in the family Physalacriaceae. The genus, widespread in temperate regions, has been estimated to contain 10 species.
List of species
* '' Flammulina callistosporioides''
* '' Flammulina elastica''
* '' Flamm ...
chanterelle
Chanterelle is the common name of several species of fungi in the genera '' Cantharellus'', '' Craterellus'', '' Gomphus'', and ''Polyozellus''. They are among the most popular of wild edible mushrooms. They are orange, yellow or white, meaty a ...
Boletus edulis
''Boletus edulis'' (English: cep, penny bun, porcino or porcini) is a basidiomycete fungus, and the type species of the genus ''Boletus''. Widely distributed in the Northern Hemisphere across Europe, Asia, and North America, it does not occu ...
'') (also known as king boletes) demand a high price on the market. They are often used in gourmet dishes.
Certain types of cheeses require inoculation of milk curds with fungal species that impart a unique flavor and texture to the cheese. Examples include the
blue
Blue is one of the three primary colours in the RYB colour model (traditional colour theory), as well as in the RGB (additive) colour model. It lies between violet and cyan on the spectrum of visible light. The eye perceives blue when obs ...
color in cheeses such as
Stilton
Stilton is a village and civil parish in Cambridgeshire, England, about north of Huntingdon in Huntingdonshire, which is a non-metropolitan district of Cambridgeshire as well as a historic county of England.
History
There is evidence of Neo ...
or
Roquefort
Roquefort is a sheep milk cheese from Southern France, and is one of the world's best known blue cheeses. Though similar cheeses are produced elsewhere, EU law dictates that only those cheeses aged in the natural Combalou caves of Roquefort-sur ...
, which are made by inoculation with ''
Penicillium roqueforti
''Penicillium roqueforti'' is a common saprotrophic fungus in the genus ''Penicillium''. Widespread in nature, it can be isolated from soil, decaying organic matter, and plants.
The major industrial use of this fungus is the production of blue ...
''. Molds used in cheese production are non-toxic and are thus safe for human consumption; however, mycotoxins (e.g., aflatoxins,
roquefortine C
Roquefortine C is a mycotoxin that belongs to a class of naturally occurring 2,5-diketopiperazines produced by various fungi, particularly species from the genus ''Penicillium''. It was first isolated from a strain of ''Penicillium roqueforti'', a ...
, patulin, or others) may accumulate because of growth of other fungi during cheese ripening or storage.
Poisonous fungi
Many mushroom species are
poisonous
Poison is a chemical substance that has a detrimental effect to life. The term is used in a wide range of scientific fields and industries, where it is often specifically defined. It may also be applied colloquially or figuratively, with a broa ...
to humans and cause a range of reactions including slight digestive problems,
allergic
Allergies, also known as allergic diseases, refer a number of conditions caused by the hypersensitivity of the immune system to typically harmless substances in the environment. These diseases include hay fever, food allergies, atopic der ...
reactions,
hallucination
A hallucination is a perception in the absence of an external stimulus that has the qualities of a real perception. Hallucinations are vivid, substantial, and are perceived to be located in external objective space. Hallucination is a combinatio ...
s, severe organ failure, and death. Genera with mushrooms containing deadly toxins include ''
Conocybe
''Conocybe'' is a genus of mushrooms with ''Conocybe tenera'' as the type species and at least 243 other species. There are at least 50 different species in North America.
Most have a long, thin fragile stipe and are delicate, growing in grassl ...
'', ''
Galerina
''Galerina'' is a genus of small brown-spore saprobic fungi (colloquially often ''mushrooms''), with over 300 species found throughout the world from the far north to remote Macquarie Island in the Southern Ocean. The genus is most noted for some ...
'', ''
Lepiota
''Lepiota'' is a genus of gilled mushrooms in the family Agaricaceae. All ''Lepiota'' species are ground-dwelling saprotrophs with a preference for rich, calcareous soils. Basidiocarps (fruit bodies) are agaricoid with whitish spores, typically ...
'' and the most infamous, ''
Amanita
The genus ''Amanita'' contains about 600 species of agarics, including some of the most toxic known mushrooms found worldwide, as well as some well-regarded edible species. This genus is responsible for approximately 95% of the fatalities resul ...
''. The latter genus includes the destroying angel ''( A.virosa)'' and the death cap ''( A.phalloides)'', the most common cause of deadly mushroom poisoning. The false morel ('' Gyromitra esculenta'') is occasionally considered a delicacy when cooked, yet can be highly toxic when eaten raw. ''
Tricholoma equestre
''Tricholoma equestre'' or ''Tricholoma flavovirens'', also known as man on horseback or yellow knight is a formerly widely eaten but arguably hazardous fungus of the genus ''Tricholoma'' that forms ectomycorrhiza with pine trees.
Known as ''Gr ...
'' was considered edible until it was implicated in serious poisonings causing
rhabdomyolysis
Rhabdomyolysis (also called rhabdo) is a condition in which damaged skeletal muscle breaks down rapidly. Symptoms may include muscle pains, weakness, vomiting, and confusion. There may be tea-colored urine or an irregular heartbeat. Some of th ...
.
Fly agaric
''Amanita muscaria'', commonly known as the fly agaric or fly amanita, is a basidiomycete of the genus ''Amanita''. It is also a muscimol mushroom. Native throughout the temperate and boreal regions of the Northern Hemisphere, ''Amanita muscar ...
mushrooms (''Amanita muscaria'') also cause occasional non-fatal poisonings, mostly as a result of ingestion for its
hallucinogenic
Hallucinogens are a large, diverse class of psychoactive drugs that can produce altered states of consciousness characterized by major alterations in thought, mood, and perception as well as other changes. Most hallucinogens can be categorize ...
properties. Historically, fly agaric was used by different peoples in Europe and Asia and its present usage for religious or
shamanic
Shamanism is a religious practice that involves a practitioner (shaman) interacting with what they believe to be a spirit world through altered states of consciousness, such as trance. The goal of this is usually to direct spirits or spiri ...
purposes is reported from some ethnic groups such as the Koryak people of northeastern
Siberia
Siberia ( ; rus, Сибирь, r=Sibir', p=sʲɪˈbʲirʲ, a=Ru-Сибирь.ogg) is an extensive geographical region, constituting all of North Asia, from the Ural Mountains in the west to the Pacific Ocean in the east. It has been a part of ...
.
As it is difficult to accurately identify a safe mushroom without proper training and knowledge, it is often advised to assume that a wild mushroom is poisonous and not to consume it.Hall, p. 7.
Pest control
In agriculture, fungi may be useful if they actively compete for nutrients and space with
pathogen
In biology, a pathogen ( el, πάθος, "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 ...
parasites
Parasitism is a Symbiosis, close relationship between species, where one organism, the parasite, lives on or inside another organism, the Host (biology), host, causing it some harm, and is Adaptation, adapted structurally to this way of lif ...
of these pathogens. For example, certain species eliminate or suppress the growth of harmful plant pathogens, such as insects,
mites
Mites are small arachnids (eight-legged arthropods). Mites span two large orders of arachnids, the Acariformes and the Parasitiformes, which were historically grouped together in the subclass Acari, but genetic analysis does not show clear evid ...
,
weed
A weed is a plant considered undesirable in a particular situation, "a plant in the wrong place", or a plant growing where it is not wanted.Harlan, J. R., & deWet, J. M. (1965). Some thoughts about weeds. ''Economic botany'', ''19''(1), 16-24. ...
s,
nematodes
The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-parasitic nematodes also known as eelworms. They are a diverse animal phylum inhabiting a broa ...
, and other fungi that cause diseases of important
crop
A crop is a plant that can be grown and harvested extensively for profit or subsistence. When the plants of the same kind are cultivated at one place on a large scale, it is called a crop. Most crops are cultivated in agriculture or hydroponic ...
plants. This has generated strong interest in practical applications that use these fungi in the
biological control
Biological control or biocontrol is a method of controlling pests, such as insects, mites, weeds, and plant diseases, using other organisms. It relies on predation, parasitism, herbivory, or other natural mechanisms, but typically also invo ...
of these agricultural pests.
Entomopathogenic fungi
An entomopathogenic fungus is a fungus that can kill or seriously disable insects.
Typical life cycle
These fungi usually attach to the external body surface of insects in the form of microscopic spores (usually asexual, mitosporic spores also ...
can be used as biopesticides, as they actively kill insects. Examples that have been used as
biological insecticide
A Biopesticide is a biological substance or organism that damages, kills, or repels organisms seens as pests. Biological pest management intervention involves predatory, parasitic, or chemical relationships.
They are obtained from organisms inclu ...
s are ''
Beauveria bassiana
''Beauveria bassiana'' is a fungus that grows naturally in soils throughout the world and acts as a parasite on various arthropod species, causing white muscardine disease; it thus belongs to the entomopathogenic fungi. It is used as a biological ...
'', ''
Metarhizium
''Metarhizium'' is a genus of entomopathogenic fungi in the Clavicipitaceae family. With the advent of genetic profiling, placing these fungi in proper taxa has now become possible. Most turn out to be the asexual forms (anamorphs) of fungi in ...
'' spp., ''
Hirsutella
''Hirsutella'' is a genus of asexually reproducing fungi in the Ophiocordycipitaceae family. Originally described by French mycologist Narcisse Théophile Patouillard in 1892, this genus includes species that are pathogens of insects, mites and ne ...
'' spp., ''
Paecilomyces
''Paecilomyces'' is a genus of fungi. A number of species in this genus are plant pathogens.
Several of the entomopathogenic species, such as "''Paecilomyces fumosoroseus''" have now been placed in the genus '' Isaria'': in the order Hypocreale ...
'' (''Isaria'') spp., and ''
Lecanicillium lecanii
''Lecanicillium lecanii'' is now an approved name of an entomopathogenic fungus species, that was previously widely known as ''Verticillium lecanii'' (Zimmerman) Viegas), but is now understood to be an anamorphic form in the ''Cordyceps'' group ...
''.
Endophytic
An endophyte is an endosymbiont, often a bacterium or fungus, that lives within a plant for at least part of its life cycle without causing apparent disease. Endophytes are ubiquitous and have been found in all species of plants studied to date; h ...
fungi of grasses of the genus ''
Epichloë
''Epichloë'' is a genus of ascomycete fungi forming an endophytic symbiosis with grasses. Grass choke disease is a symptom in grasses induced by some ''Epichloë'' species, which form spore-bearing mats ( stromata) on tillers and suppress t ...
'', such as '' E. coenophiala'', produce alkaloids that are toxic to a range of invertebrate and vertebrate
herbivores
A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
. These alkaloids protect grass plants from
herbivory
A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
, but several endophyte alkaloids can poison grazing animals, such as cattle and sheep. Infecting cultivars of
pasture
Pasture (from the Latin ''pastus'', past participle of ''pascere'', "to feed") is land used for grazing. Pasture lands in the narrow sense are enclosed tracts of farmland, grazed by domesticated livestock, such as horses, cattle, sheep, or swine ...
or
forage
Forage is a plant material (mainly plant leaves and stems) eaten by grazing livestock. Historically, the term ''forage'' has meant only plants eaten by the animals directly as pasture, crop residue, or immature cereal crops, but it is also used m ...
grasses with ''Epichloë'' endophytes is one approach being used in grass breeding programs; the fungal strains are selected for producing only alkaloids that increase resistance to herbivores such as insects, while being non-toxic to livestock.
Bioremediation
Certain fungi, in particular white-rot fungi, can degrade
insecticide
Insecticides are substances used to kill insects. They include ovicides and larvicides used against insect eggs and larvae, respectively. Insecticides are used in agriculture, medicine, industry and by consumers. Insecticides are claimed to b ...
s,
herbicide
Herbicides (, ), also commonly known as weedkillers, are substances used to control undesired plants, also known as weeds.EPA. February 201Pesticides Industry. Sales and Usage 2006 and 2007: Market Estimates. Summary in press releasMain page fo ...
s,
pentachlorophenol
Pentachlorophenol (PCP) is an organochlorine compound used as a pesticide and a disinfectant. First produced in the 1930s, it is marketed under many trade names. It can be found as pure PCP, or as the sodium salt of PCP, the latter of which diss ...
,
creosote
Creosote is a category of carbonaceous chemicals formed by the distillation of various tars and pyrolysis of plant-derived material, such as wood or fossil fuel. They are typically used as preservatives or antiseptics.
Some creosote types were ...
,
coal tar
Coal tar is a thick dark liquid which is a by-product of the production of coke and coal gas from coal. It is a type of creosote. It has both medical and industrial uses. Medicinally it is a topical medication applied to skin to treat psoriasi ...
s, and heavy fuels and turn them into
carbon dioxide
Carbon dioxide (chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is transpar ...
, water, and basic elements. Fungi have been shown to biomineralization, biomineralize uranium#Oxides, uranium oxides, suggesting they may have application in the bioremediation of radioactively polluted sites.
Model organisms
Several pivotal discoveries in biology were made by researchers using fungi as model organisms, that is, fungi that grow and sexually reproduce rapidly in the laboratory. For example, the one gene-one enzyme hypothesis was formulated by scientists using the bread mold ''
Neurospora crassa
''Neurospora crassa'' is a type of red bread mold of the phylum Ascomycota. The genus name, meaning "nerve spore" in Greek, refers to the characteristic striations on the spores. The first published account of this fungus was from an infestation ...
'' to test their biochemical theories. Other important model fungi are ''Aspergillus nidulans'' and the yeasts ''
Saccharomyces cerevisiae
''Saccharomyces cerevisiae'' () (brewer's yeast or baker's yeast) is a species of yeast (single-celled fungus microorganisms). The species has been instrumental in winemaking, baking, and brewing since ancient times. It is believed to have been o ...
'' and ''Schizosaccharomyces pombe'', each of which with a long history of use to investigate issues in eukaryotic cell biology and
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 Augustinian friar wor ...
, such as cell cycle regulation, chromatin structure, and gene regulation. Other fungal models have emerged that address specific biological questions relevant to medicine, plant pathology, and industrial uses; examples include ''Candida albicans'', a dimorphic, opportunistic human pathogen, '' Magnaporthe grisea'', a plant pathogen, and ''Pichia pastoris'', a yeast widely used for eukaryotic protein production.
Others
Fungi are used extensively to produce industrial chemicals like citric acid, citric, gluconic acid, gluconic, lactic acid, lactic, and malic acid, malic acids, and industrial enzymes, such as lipases used in biological detergents, cellulases used in making cellulosic ethanol and stonewashed jeans, and amylases, invertases, proteases and xylanases.
See also
* Conservation of fungi
* Fantastic Fungi
* Glossary of fungi
* Marine fungi
* Mycosis
* Outline of fungi
References
Citations
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