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109-400: A fungus ( pl. : 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 one of the traditional eukaryotic kingdoms , along with Animalia , Plantae , and either Protista or Protozoa and Chromista . A characteristic that places fungi in

218-476: A gametangium , a specialized cell structure that becomes a fertile gamete -producing cell. The gametangium develops into a zygospore , a thick-walled spore formed by the union of gametes. When the zygospore germinates, it undergoes meiosis , 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. The spores of most of

327-405: A microtubular spindle during nuclear division, in the distinctively eukaryotic process of mitosis . Eukaryotes differ from prokaryotes in multiple ways, with unique biochemical pathways such as sterane synthesis. The eukaryotic signature proteins have no homology to proteins in other domains of life, but appear to be universal among eukaryotes. They include the proteins of the cytoskeleton,

436-418: A monophyletic origin of fungi. The taxonomy 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 matings . There is no unique generally accepted system at

545-518: A "symbiosis-based phylogeny", giving the description "Eukarya (symbiosis-derived nucleated organisms)". By 2014, a rough consensus started to emerge from the phylogenomic studies of the previous two decades. The majority of eukaryotes can be placed in one of two large clades dubbed Amorphea (similar in composition to the unikont hypothesis) and the Diphoda (formerly bikonts), which includes plants and most algal lineages. A third major grouping,

654-430: A body, with its cells dividing by mitosis , and at some stage produce haploid gametes through meiosis , a division that reduces the number of chromosomes and creates genetic variability . There is considerable variation in this pattern. Plants have both haploid and diploid multicellular phases . Eukaryotes have lower metabolic rates and longer generation times than prokaryotes, because they are larger and therefore have

763-479: A bundle of microtubules arising from a centriole , characteristically arranged as nine doublets surrounding two singlets. Flagella may have hairs ( mastigonemes ), as in many stramenopiles . Their interior is continuous with the cell's cytoplasm . Centrioles are often present, even in cells and groups that do not have flagella, but conifers and flowering plants have neither. They generally occur in groups that give rise to various microtubular roots. These form

872-732: 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 ( basidiocarps ) and some ascomycetes can sometimes grow very large, and many are well known as mushrooms . 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

981-417: A different kingdom from plants , bacteria , and some protists is chitin in their cell walls . Fungi, like animals, are heterotrophs ; they acquire their food by absorbing dissolved molecules, typically by secreting digestive enzymes into their environment. Fungi do not photosynthesize . Growth is their means of mobility , except for spores (a few of which are flagellated ), which may travel through

1090-650: A direct source of human food , in the form of mushrooms and truffles ; as a leavening agent for bread; and in the fermentation of various food products, such as wine , beer , and soy sauce . Since the 1940s, fungi have been used for the production of antibiotics , and, more recently, various enzymes produced by fungi are used industrially and in detergents . Fungi are also used as biological pesticides to control weeds, plant diseases, and insect pests. Many species produce bioactive compounds called mycotoxins , such as alkaloids and polyketides , that are toxic to animals, including humans. The fruiting structures of

1199-615: A few species contain psychotropic compounds and are consumed recreationally or in traditional spiritual ceremonies . Fungi can break down manufactured materials and buildings, and become significant pathogens of humans and other animals. Losses of crops due to fungal diseases (e.g., rice blast disease ) or food spoilage can have a large impact on human food supplies and local economies. The fungus kingdom encompasses an enormous diversity of taxa with varied ecologies, life cycle strategies, and morphologies ranging from unicellular aquatic chytrids to large mushrooms. However, little

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1308-494: 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)

1417-403: 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 , acetate , or ethanol . In some species the pigment melanin 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,

1526-420: A high sugar content. The same effect is useful in the production of various types of food and beverages, such as bread , yogurt , cider , and alcoholic beverages . Signs of food spoilage may include an appearance different from the food in its fresh form, such as a change in color, a change in texture, an unpleasant odour , or an undesirable taste . The item may become softer than normal. If mold occurs, it

1635-567: A motile zoospore , enabling them to propel themselves through water and enter their amphibian host. Other examples of aquatic fungi include those living in hydrothermal 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 2,000 with

1744-401: 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 , is formed at each hyphal septum. As with

1853-406: A particularly long period of time, whether done at home or commercially. Canned food is vacuum packed in order to keep oxygen, which is needed by bacteria in aerobic spoilage, out of the can. Canning does have limitations, and does not preserve the food indefinitely. Lactic acid fermentation also preserves food and prevents spoilage. Food like meat, poultry, milk and cream should be kept out of

1962-429: A peak of more than 2,500 species in 2016. In the year 2019, 1,882 new species of fungi were described, and it was estimated that more than 90% of fungi remain unknown. The following year, 2,905 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

2071-406: A primary component of the cytoskeleton, and are often assembled over the course of several cell divisions, with one flagellum retained from the parent and the other derived from it. Centrioles produce the spindle during nuclear division. The cells of plants, algae, fungi and most chromalveolates , but not animals, are surrounded by a cell wall. This is a layer outside the cell membrane , providing

2180-514: 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 or coenocytic . Septate hyphae are divided into compartments separated by cross walls (internal cell walls, called septa, that are formed at right angles to

2289-564: 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 0.01–0.02   cm, sufficient distance for it to fall through the gills 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

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2398-510: A smaller surface area to volume ratio. The evolution of sexual reproduction may be a primordial characteristic of eukaryotes. Based on a phylogenetic analysis, Dacks and Roger have proposed that facultative sex was present in the group's common ancestor. A core set of genes that function in meiosis is present in both Trichomonas vaginalis and Giardia intestinalis , two organisms previously thought to be asexual. Since these two species are descendants of lineages that diverged early from

2507-577: A structure called an appressorium that evolved to puncture plant tissues. The pressure generated by the appressorium, directed against the plant epidermis , can exceed 8 megapascals (1,200 psi). The filamentous fungus Paecilomyces lilacinus 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 by producing osmolytes such as glycerol . Adaptations such as these are complemented by hydrolytic enzymes secreted into

2616-417: A system of domains rather than kingdoms as top level rank being put forward by Carl Woese , Otto Kandler , and Mark Wheelis in 1990, uniting all the eukaryote kingdoms in the domain "Eucarya", stating, however, that " 'eukaryotes' will continue to be an acceptable common synonym". In 1996, the evolutionary biologist Lynn Margulis proposed to replace kingdoms and domains with "inclusive" names to create

2725-486: 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

2834-667: 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 of a species, the teleomorph (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 occurs via vegetative spores ( conidia ) or through mycelial fragmentation . Mycelial fragmentation occurs when

2943-599: Is 415   Ma; 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 from the Pennsylvanian. Rare in the fossil record are the Homobasidiomycetes (a taxon roughly equivalent to

3052-472: Is a type of fungus, but the two terms are not reciprocal of each other; they have their own defining features and perform their own tasks. Very well known types of mold are Aspergillus and Penicillium , and, like regular fungi, create a fuzz, powder and slime of various colors. Yeast is also a type of fungus that grows vegetatively via single cells that either bud or divide by way of fission, allowing for yeast to multiply in liquid environments favoring

3161-610: Is closer in structure to bacterial RNA than to eukaryote RNA. Some eukaryotes, such as the metamonads Giardia and Trichomonas , and the amoebozoan Pelomyxa , appear to lack mitochondria, but all contain mitochondrion-derived organelles, like hydrogenosomes or mitosomes , having lost their mitochondria secondarily. They obtain energy by enzymatic action in the cytoplasm. Plants and various groups of algae have plastids as well as mitochondria. Plastids, like mitochondria, have their own DNA and are developed from endosymbionts , in this case cyanobacteria . They usually take

3270-402: Is considered to be the publication of Pier Antonio Micheli '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 in his Species plantarum (1753),

3379-622: Is known of the true biodiversity 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 works of Carl Linnaeus , 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 . Advances in molecular genetics have opened

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3488-643: Is much larger than that of prokaryotes. The eukaryotes seemingly emerged within the Asgard archaea , and are closely related to the Heimdallarchaeia . This implies that there are only two domains of life , Bacteria and Archaea, with eukaryotes incorporated among the Archaea. Eukaryotes first emerged during the Paleoproterozoic , likely as flagellated cells. The leading evolutionary theory

3597-641: Is not an accepted taxonomic clade and is now taken to mean simply fungi that lack a known sexual stage. Sexual reproduction with meiosis 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

3706-519: Is often visible externally on the item. Spoilage bacteria do not normally cause " food poisoning "; typically, the microorganisms that cause foodborne illnesses are odorless and flavourless, and otherwise undetectable outside the lab. Eating deteriorated food could not be considered safe due to mycotoxins or microbial wastes. Some pathogenic bacteria, such as Clostridium perfringens and Bacillus cereus , are capable of causing spoilage. Issues of food spoilage do not necessarily have to do with

3815-536: Is that their cells have nuclei . This gives them their name, from the Greek εὖ ( eu , "well" or "good") and κάρυον ( karyon , "nut" or "kernel", here meaning "nucleus"). Eukaryotic cells have a variety of internal membrane-bound structures, called organelles , and a cytoskeleton which defines the cell's organization and shape. The nucleus stores the cell's DNA , which is divided into linear bundles called chromosomes ; these are separated into two matching sets by

3924-536: 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 Dikarya , the most species rich and familiar group, including all the mushrooms, most food-spoilage molds, most plant pathogenic fungi, and

4033-653: Is the use of biosensors , which are devices that use biological components to detect specific substances or changes in the environment. Biosensors can detect spoilage by identifying the byproducts produced by bacteria or fungi during the spoilage process. Another method of detecting spoilage is through the use of gas sensors . Bacterial contamination of food and drinks can generate unpleasant odours and toxic substances, and e-noses(gas sensor) can perform odour detection continually and are not subject to individual sensitivity. Lastly, colorimetric analysis can also be used to detect spoilage in food products. Biogenic amines are

4142-868: Is they were created by symbiogenesis between an anaerobic Asgard archaean and an aerobic proteobacterium , which formed the mitochondria . A second episode of symbiogenesis with a cyanobacterium created the plants, with chloroplasts . Eukaryotic cells contain membrane-bound organelles such as the nucleus , the endoplasmic reticulum , and the Golgi apparatus . Eukaryotes may be either unicellular or multicellular . In comparison, prokaryotes are typically unicellular. Unicellular eukaryotes are sometimes called protists . Eukaryotes can reproduce both asexually through mitosis and sexually through meiosis and gamete fusion ( fertilization ). Eukaryotes are organisms that range from microscopic single cells , such as picozoans under 3 micrometres across, to animals like

4251-425: 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. In contrast to plants and animals , the early fossil record of the fungi is meager. Factors that likely contribute to the under-representation of fungal species among fossils include

4360-1206: The Excavata , has been abandoned as a formal group as it is paraphyletic . The proposed phylogeny below includes only one group of excavates ( Discoba ), and incorporates the 2021 proposal that picozoans are close relatives of rhodophytes. The Provora are a group of microbial predators discovered in 2022. Ancyromonadida [REDACTED] Malawimonada [REDACTED] CRuMs [REDACTED] Amoebozoa [REDACTED] Breviatea [REDACTED] Apusomonadida [REDACTED] Holomycota (inc. fungi) [REDACTED] Holozoa (inc. animals) [REDACTED] ? Metamonada [REDACTED] Discoba [REDACTED] Cryptista [REDACTED] Rhodophyta (red algae) [REDACTED] Picozoa [REDACTED] Glaucophyta [REDACTED] Viridiplantae (plants) [REDACTED] Hemimastigophora [REDACTED] Provora [REDACTED] Haptista [REDACTED] Telonemia [REDACTED] Rhizaria [REDACTED] Alveolata [REDACTED] Stramenopiles [REDACTED] [REDACTED] Food spoilage Some bacteria are responsible for

4469-643: The Latin fungus (mushroom), used in the writings of Horace and Pliny . This in turn is derived from the Greek word sphongos (σφόγγος 'sponge'), which refers to the macroscopic structures and morphology of mushrooms and molds; the root is also used in other languages, such as the German Schwamm ('sponge') and Schimmel ('mold'). The word mycology is derived from the Greek mykes (μύκης 'mushroom') and logos (λόγος 'discourse'). It denotes

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4578-517: The Ordovician of Wisconsin (460   Ma) resemble modern-day Glomerales , and existed at a time when the land flora likely consisted of only non-vascular bryophyte -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 (416–359.2   Ma), when they occur abundantly in

4687-526: The Rhynie chert , mostly as Zygomycota and Chytridiomycota . At about this same time, approximately 400   Ma, the Ascomycota and Basidiomycota diverged, and all modern classes of fungi were present by the Late Carboniferous ( Pennsylvanian , 318.1–299   Ma). Lichens formed a component of the early terrestrial ecosystems, and the estimated age of the oldest terrestrial lichen fossil

4796-518: The archaea —having a volume of around 10,000 times greater. Eukaryotes represent a small minority of the number of organisms , but, as many of them are much larger, their collective global biomass (468 gigatons) is far larger than that of prokaryotes (77 gigatons), with plants alone accounting for over 81% of the total biomass of Earth . The eukaryotes are a diverse lineage, consisting mainly of microscopic organisms . Multicellularity in some form has evolved independently at least 25 times within

4905-541: 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 . Besides regular sexual reproduction with meiosis, certain fungi, such as those in the genera Penicillium and Aspergillus , may exchange genetic material via parasexual processes, initiated by anastomosis between hyphae and plasmogamy of fungal cells. The frequency and relative importance of parasexual events

5014-439: The biopolymer chitin. 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 molds . 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 ) that can be used as diagnostic features in

5123-583: The blue whale , weighing up to 190 tonnes and measuring up to 33.6 metres (110 ft) long, or plants like the coast redwood , up to 120 metres (390 ft) tall. Many eukaryotes are unicellular; the informal grouping called protists includes many of these, with some multicellular forms like the giant kelp up to 200 feet (61 m) long. The multicellular eukaryotes include the animals, plants, and fungi , but again, these groups too contain many unicellular species . Eukaryotic cells are typically much larger than those of prokaryotes —the bacteria and

5232-558: The domain of Eukaryota or Eukarya , organisms whose cells have a membrane-bound nucleus . All animals , plants , fungi , and many unicellular organisms are eukaryotes. They constitute a major group of life forms alongside the two groups of prokaryotes : the Bacteria and the Archaea . Eukaryotes represent a small minority of the number of organisms, but given their generally much larger size, their collective global biomass

5341-452: The roots of plants. As eukaryotes, fungi possess a biosynthetic pathway for producing terpenes that uses mevalonic acid and pyrophosphate 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 metabolites that are similar or identical in structure to those made by plants. Many of

5450-514: The taxonomic rank of kingdom by Linnaeus in the 18th century. Though he included the fungi with plants with some reservations, it was later realized that they are quite distinct and warrant a separate kingdom. The various single-cell eukaryotes were originally placed with plants or animals when they became known. In 1818, the German biologist Georg A. Goldfuss coined the word Protozoa to refer to organisms such as ciliates , and this group

5559-536: The Danger Zone (between 4 and 60 °C (39 and 140 °F)). Anything between that range is considered dangerous and can cause pathogenic toxins to be emitted, resulting in severe illness in the consumer. Another way to keep food from spoiling is by following a four step system: Clean, Separate, Cook, Chill. The detection of spoiled food can be crucial in preventing Foodborne illnesses , and various methods have been developed to detect spoilage. One such method

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5668-601: 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 , August Carl Joseph Corda , Anton de Bary ,

5777-454: 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 (i.e. they form a monophyletic group ), an interpretation that is also strongly supported by molecular phylogenetics . This fungal group is distinct from

5886-485: 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

5995-641: 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 biofilms for substrate colonization and intercellular communication . Fungi are traditionally considered heterotrophs , organisms that rely solely on carbon fixed by other organisms for metabolism . Fungi have evolved

6104-413: 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 , can be used in the identification of ascomycetes and basidiomycetes, respectively. Fungi employ two mating systems : heterothallic species allow mating only between individuals of

6213-494: The basis of comparisons of the rate of evolution in closely related groups. The oldest fossilizied mycelium to be identified from its molecular composition is between 715 and 810 million years old. For much of the Paleozoic Era (542–251   Ma), 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

6322-648: The beer, wine, and bread yeasts. The accompanying cladogram depicts the major fungal taxa 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. Rozellomycetes Mitosporidium Paramicrosporidium Nucleophaga Metchnikovellea Eukaryotic The eukaryotes ( / j uː ˈ k ær i oʊ t s , - ə t s / yoo- KARR -ee-ohts, -⁠əts ) constitute

6431-691: The brothers Louis René and Charles Tulasne , Arthur H. R. Buller , Curtis G. Lloyd , and Pier Andrea Saccardo . In the 20th and 21st centuries, advances in biochemistry , genetics , molecular biology , biotechnology , DNA sequencing , and phylogenetic analysis have provided new insights into fungal relationships and biodiversity , and have challenged traditional morphology-based groupings in fungal taxonomy . 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

6540-709: The cell to move, change shape, or transport materials. The motor structures are microfilaments of actin and actin-binding proteins , including α- actinin , fimbrin , and filamin are present in submembranous cortical layers and bundles. Motor proteins of microtubules, dynein and kinesin , and myosin of actin filaments, provide dynamic character of the network. Many eukaryotes have long slender motile cytoplasmic projections, called flagella , or multiple shorter structures called cilia . These organelles are variously involved in movement, feeding, and sensation. They are composed mainly of tubulin , and are entirely distinct from prokaryotic flagella. They are supported by

6649-617: 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 , 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 supercells. Many species have developed specialized hyphal structures for nutrient uptake from living hosts; examples include haustoria in plant-parasitic species of most fungal phyla, and arbuscules of several mycorrhizal fungi, which penetrate into

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6758-404: The cell with structural support, protection, and a filtering mechanism. The cell wall also prevents over-expansion when water enters the cell. The major polysaccharides making up the primary cell wall of land plants are cellulose , hemicellulose , and pectin . The cellulose microfibrils are linked together with hemicellulose, embedded in a pectin matrix. The most common hemicellulose in

6867-427: The complex transcription machinery, the membrane-sorting systems, the nuclear pore , and some enzymes in the biochemical pathways. Eukaryote cells include a variety of membrane-bound structures, together forming the endomembrane system. Simple compartments, called vesicles and vacuoles , can form by budding off other membranes. Many cells ingest food and other materials through a process of endocytosis , where

6976-622: The cytoplasm. Mitochondria are organelles in eukaryotic cells. The mitochondrion is commonly called "the powerhouse of the cell", for its function providing energy by oxidising sugars or fats to produce the energy-storing molecule ATP . Mitochondria have two surrounding membranes , each a phospholipid bilayer , the inner of which is folded into invaginations called cristae where aerobic respiration takes place. Mitochondria contain their own DNA , which has close structural similarities to bacterial DNA , from which it originated, and which encodes rRNA and tRNA genes that produce RNA which

7085-688: The discovery of a fossilized fungus, named Ourasphaira giraldae , in the Canadian Arctic , that may have grown on land a billion years ago, well before plants were living on land. Pyritized fungus-like microfossils 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 (542–488.3   Ma), also long before land plants. Fossilized hyphae and spores recovered from

7194-412: The dissemination of single celled microorganisms. Yeast forms mainly in liquid environments and anaerobic conditions, but being single celled, it oftentimes cannot spread on or into solid surfaces where other fungus flourish. Yeast also produces at a slower rate than bacteria, therefore being at a disadvantage in environments where bacteria are. Yeasts can be responsible for the decomposition of food with

7303-597: The dominant life form at this time, representing nearly 100% of the available fossil record 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 that famously killed off most dinosaurs, there

7412-409: The effects on growth rates are small, and the underlying biophysical and biochemical processes are not well known. This process might bear similarity to CO 2 fixation via visible light , but instead uses ionizing radiation as a source of energy. Fungal reproduction is complex, reflecting the differences in lifestyles and genetic makeup within this diverse kingdom of organisms. It is estimated that

7521-423: The environment to digest large organic molecules—such as polysaccharides , proteins , and lipids —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

7630-406: The eukaryotes. Complex multicellular organisms, not counting the aggregation of amoebae to form slime molds , have evolved within only six eukaryotic lineages: animals , symbiomycotan fungi , brown algae , red algae , green algae , and land plants . Eukaryotes are grouped by genomic similarities, so that groups often lack visible shared characteristics. The defining feature of eukaryotes

7739-484: The eukaryotic evolutionary tree, core meiotic genes, and hence sex, were likely present in the common ancestor of eukaryotes. Species once thought to be asexual, such as Leishmania parasites, have a sexual cycle. Amoebae, previously regarded as asexual, may be anciently sexual; while present-day asexual groups could have arisen recently. In antiquity , the two lineages of animals and plants were recognized by Aristotle and Theophrastus . The lineages were given

7848-424: The first in first out method (FIFO), which ensures that the first item purchased is the first item consumed. Preservatives can expand the shelf life of food and can lengthen the time long enough for it to be harvested, processed, sold, and kept in the consumer's home for a reasonable length of time. One of the age old techniques for food preservation, to avoid mold and fungus growth, is the process of drying out

7957-551: The food or dehydrating it. While there is a chance of it developing a fungus targeted towards dried food products, the chances are quite low. Other than drying, other methods include salting, curing, canning, refrigeration, freezing, preservatives, irradiation, and high hydrostatic pressure: Refrigeration can increase the shelf life of certain foods and beverages, though with most items, it does not indefinitely expand it. Freezing can preserve food even longer, though even freezing has limitations. Canning of food can preserve food for

8066-532: The force of falling water drops to liberate the spores from cup-shaped fruiting bodies. Another strategy is seen in the stinkhorns , a group of fungi with lively colors and putrid odor that attract insects to disperse their spores. In homothallic sexual reproduction , two haploid nuclei derived from the same individual fuse to form a zygote that can then undergo meiosis . Homothallic fungi include species with an Aspergillus -like asexual stage (anamorphs) occurring in numerous different genera, several species of

8175-646: The form of chloroplasts which, like cyanobacteria, contain chlorophyll and produce organic compounds (such as glucose ) through photosynthesis . Others are involved in storing food. Although plastids probably had a single origin, not all plastid-containing groups are closely related. Instead, some eukaryotes have obtained them from others through secondary endosymbiosis or ingestion. The capture and sequestering of photosynthetic cells and chloroplasts, kleptoplasty , occurs in many types of modern eukaryotic organisms. The cytoskeleton provides stiffening structure and points of attachment for motor structures that enable

8284-417: 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

8393-412: 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 mushrooms , 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

8502-773: 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 . 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 and MycoBank are officially recognized nomenclatural repositories and list current names of fungal species (with cross-references to older synonyms ). The 2007 classification of Kingdom Fungi

8611-406: The host cells to consume nutrients. Although fungi are opisthokonts —a grouping of evolutionarily related organisms broadly characterized by a single posterior flagellum —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

8720-409: 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 9,000   years. The apothecium —a specialized structure important in sexual reproduction in the ascomycetes—is

8829-473: The important markers for food spoilage, therefore, an on-package sensor for biogenic amine detection is crucial for food quality control. The colorimetric sensors use a chemical reaction between a substance in the food product and a reagent to produce a color change. The color change can indicate the presence of spoilage, as well as the extent of the spoilage. These methods of detecting spoilage can be used in combination to provide more accurate results, while it

8938-415: 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 and B. salamandrivorans , parasites that have been responsible for a worldwide decline in amphibian populations. These organisms spend part of their life cycle as

9047-591: The mushroom-producing species of the Agaricomycetes ). Two amber -preserved specimens provide evidence that the earliest known mushroom-forming fungi (the extinct species Archaeomarasmius leggetti ) appeared during the late Cretaceous , 90   Ma. Some time after the Permian–Triassic extinction event (251.4   Ma), a fungal spike (originally thought to be an extraordinary abundance of fungal spores in sediments ) formed, suggesting that fungi were

9156-605: The nature of fungal fruiting bodies , 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 fossils by dissolving

9265-400: The opposite mating type , whereas homothallic 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

9374-518: The organisms will begin to breed apace, releasing harmful toxins that can cause severe illness, even when cooked safely. Fungi have been seen as a method of food spoilage, causing only an undesirable appearance to food, however, there has been significant evidence of various fungi being a cause of death. Fungi are caused by acidifying, fermenting, discoloring and disintegrating processes and can create fuzz, powder and slimes of many different colors, including black, white, red, brown and green. Mold

9483-449: The outer membrane invaginates and then pinches off to form a vesicle. Some cell products can leave in a vesicle through exocytosis . The nucleus is surrounded by a double membrane known as the nuclear envelope , with nuclear pores that allow material to move in and out. Various tube- and sheet-like extensions of the nuclear membrane form the endoplasmic reticulum , which is involved in protein transport and maturation. It includes

9592-444: The phrase fauna and flora be replaced by fauna, flora, and funga . 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

9701-463: The plant and fungal enzymes that make these compounds differ from each other in sequence and other characteristics, which indicates separate origins and convergent evolution of these enzymes in the fungi and plants. Fungi have a worldwide distribution, and grow in a wide range of habitats, including extreme environments such as deserts or areas with high salt concentrations or ionizing radiation , as well as in deep sea sediments. Some can survive

9810-416: 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. Mycology became a systematic science after the development of the microscope in the 17th century. Although fungal spores were first observed by Giambattista della Porta in 1588, the seminal work in the development of mycology

9919-417: The primary cell wall is xyloglucan . Eukaryotes have a life cycle that involves sexual reproduction , alternating between a haploid phase, where only one copy of each chromosome is present in each cell, and a diploid phase, with two copies of each chromosome in each cell. The diploid phase is formed by fusion of two haploid gametes, such as eggs and spermatozoa , to form a zygote ; this may grow into

10028-530: The quality of the food, but more so with the safety of consuming said food. However, there are cases where food has been proven to contain toxic ingredients. 200 years ago, Claviceps purpurea , a type of fungus, was linked to human diseases and 100 years ago in Japan, yellow rice was found to contain toxic ingredients. A number of methods of prevention can be used that can either totally prevent, delay, or otherwise reduce food spoilage. A food rotation system uses

10137-446: The researched species of fungi are transported by wind. Such species often produce dry or hydrophobic 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

10246-465: The resolution and added robustness to estimates of genetic diversity within various taxonomic groups. Mycology is the branch of biology 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 ,

10355-498: The rough endoplasmic reticulum, covered in ribosomes which synthesize proteins; these enter the interior space or lumen. Subsequently, they generally enter vesicles, which bud off from the smooth endoplasmic reticulum. In most eukaryotes, these protein-carrying vesicles are released and further modified in stacks of flattened vesicles ( cisternae ), the Golgi apparatus . Vesicles may be specialized; for instance, lysosomes contain digestive enzymes that break down biomolecules in

10464-458: 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 . In 1836 the English naturalist Miles Joseph Berkeley 's publication The English Flora of Sir James Edward Smith, Vol. 5. also refers to mycology as

10573-419: 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 ,

10682-507: The size and shape of spores or fruiting structures, has traditionally dominated fungal taxonomy. 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 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

10791-434: 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 . They may become noticeable when fruiting , either as mushrooms or as molds. Fungi perform an essential role in the decomposition of organic matter and have fundamental roles in nutrient cycling and exchange in the environment. They have long been used as

10900-538: The spoilage of food. When bacteria breaks down the food, acids and other waste products are generated in the process. While the bacteria itself may or may not be harmful, the waste products may be unpleasant to taste or may even be harmful to one's health. There are many species of pathogenic bacteria that target different categories of food. For example, Clostridium botulinum spoils food such as meat and poultry, and Bacillus cereus , which spoils almost all type of food. When stored or subjected to unruly conditions,

11009-595: The start of the Neoproterozoic 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: Unique features: Most fungi lack an efficient system for the long-distance transport of water and nutrients, such as the xylem and phloem in many plants. To overcome this limitation, some fungi, such as Armillaria , form rhizomorphs , which resemble and perform functions similar to

11118-438: The structurally similar myxomycetes (slime molds) and oomycetes (water molds). The discipline of biology devoted to the study of fungi is known as mycology (from the Greek μύκης mykes , mushroom). In the past, mycology was regarded as a branch of botany , although it is now known that fungi are genetically more closely related to animals than to plants. Abundant worldwide, most fungi are inconspicuous because of

11227-607: 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 is formed in which club-like structures known as basidia generate haploid basidiospores after karyogamy and meiosis. The most commonly known basidiocarps are mushrooms, but they may also take other forms (see Morphology section). In fungi formerly classified as Zygomycota , haploid hyphae of two individuals fuse, forming

11336-590: The study of fungi. A group of all the fungi present in a particular region is known as mycobiota (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 and flora . The Species Survival Commission (SSC) of the International Union for Conservation of Nature (IUCN) in August 2021 asked that

11445-614: 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 , a well-preserved mummy of a 5,300-year-old Neolithic man found frozen in the Austrian Alps, carried two species of polypore mushrooms that may have been used as tinder ( Fomes fomentarius ), or for medicinal purposes ( Piptoporus betulinus ). Ancient peoples have used fungi as food sources—often unknowingly—for millennia, in

11554-443: 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 era, some 2,400  million years ago ( Ma ); these multicellular benthic organisms had filamentous structures capable of anastomosis . Other studies (2009) estimate the arrival of fungal organisms at about 760–1060   Ma on

11663-420: The way for DNA analysis to be incorporated into taxonomy, which has sometimes challenged the historical groupings based on morphology and other traits. Phylogenetic 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 , seven phyla , and ten subphyla . The English word fungus is directly adopted from

11772-408: 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". Although commonly included in botany curricula and textbooks, fungi are more closely related to animals than to plants and are placed with the animals in the monophyletic group of opisthokonts . Analyses using molecular phylogenetics support

11881-503: Was expanded until Ernst Haeckel made it a kingdom encompassing all single-celled eukaryotes, the Protista , in 1866. The eukaryotes thus came to be seen as four kingdoms: The protists were at that time thought to be "primitive forms", and thus an evolutionary grade , united by their primitive unicellular nature. Understanding of the oldest branchings in the tree of life only developed substantially with DNA sequencing , leading to

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