document_title: Fungus

question_text: a fungus that lives as a single cell and does not form a hyphae is called

question_tokens: a fungus that lives as a single cell and does not form a hyphae is called

text: Fungus

text: Etymology

text: The English word fungus is directly adopted from the Latin fungus (mushroom), used in the writings of Horace and Pliny.[6] This in turn is derived from the Greek word sphongos (σφογγος "sponge"), which refers to the macroscopic structures and morphology of mushrooms and molds;[7] the root is also used in other languages, such as the German Schwamm ("sponge") and Schimmel ("mold").[8] The use of the word mycology, which is derived from the Greek mykes (μύκης "mushroom") and logos (λόγος "discourse"),[9] to denote the scientific study of fungi is thought to have originated in 1836 with English naturalist Miles Joseph Berkeley's publication The English Flora of Sir James Edward Smith, Vol. 5.[7] A group of all the fungi present in a particular area or geographic region is known as mycobiota (plural noun, no singular), e.g., "the mycobiota of Ireland".[10]

text: Characteristics

text:
Hyphal wall
Septum
Mitochondrion
Vacuole
Ergosterol crystal
Ribosome
Nucleus
Endoplasmic reticulum
Lipid body
Plasma membrane
Spitzenkörper
Golgi apparatus

tag: ol

text: 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. 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.[11][12] 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:

text: Shared features:

text:
With other eukaryotes: Fungal cells contain membrane-bound nuclei with chromosomes that contain DNA with noncoding regions called introns and coding regions called exons. Fungi have membrane-bound cytoplasmic organelles such as mitochondria, sterol-containing membranes, and ribosomes of the 80S type.[13] They have a characteristic range of soluble carbohydrates and storage compounds, including sugar alcohols (e.g., mannitol), disaccharides, (e.g., trehalose), and polysaccharides (e.g., glycogen, which is also found in animals[14]).
With animals: Fungi lack chloroplasts and are heterotrophic organisms and so require preformed organic compounds as energy sources.[15]
With plants: Fungi have a cell wall[16] and vacuoles.[17] They reproduce by both sexual and asexual means, and like basal plant groups (such as ferns and mosses) produce spores. Similar to mosses and algae, fungi typically have haploid nuclei.[18]
With euglenoids and bacteria: Higher fungi, euglenoids, and some bacteria produce the amino acid L-lysine in specific biosynthesis steps, called the α-aminoadipate pathway.[19][20]
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 proteins, lipids, and other organic molecules—called the Spitzenkörper.[21] Both fungi and oomycetes grow as filamentous hyphal cells.[22] In contrast, similar-looking organisms, such as filamentous green algae, grow by repeated cell division within a chain of cells.[14] There are also single-celled fungi (yeasts) that do not form hyphae, and some fungi have both hyphal and yeast forms.[23]
In common with some plant and animal species, more than 70 fungal species display bioluminescence.[24]

text: Unique features:

text:
Some species grow as unicellular yeasts that reproduce by budding or binary fission. Dimorphic fungi can switch between a yeast phase and a hyphal phase in response to environmental conditions.[23]
The fungal cell wall is composed of glucans and chitin; while glucans are also found in plants and chitin in the exoskeleton of arthropods,[25][26] 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.[27]

text: 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,[28] which resemble and perform functions similar to the roots of plants. As eukaryotes, fungi possess a biosynthetic pathway for producing terpenes that uses mevalonic acid and pyrophosphate as chemical building blocks.[29] Plants and some other organisms have an additional terpene biosynthesis pathway in their chloroplasts, a structure fungi and animals do not have.[30] Fungi produce several secondary metabolites that are similar or identical in structure to those made by plants.[29] Many of 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.[29][31]

text: Diversity

text: 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[32] or ionizing radiation,[33] as well as in deep sea sediments.[34] Some can survive the intense UV and cosmic radiation encountered during space travel.[35] Most grow in terrestrial environments, though several species live partly or solely in aquatic habitats, such as the chytrid fungus Batrachochytrium dendrobatidis, a parasite that has been responsible for a worldwide decline in amphibian populations. This organism spends part of its life cycle as a motile zoospore, enabling it to propel itself through water and enter its amphibian host.[36] Other examples of aquatic fungi include those living in hydrothermal areas of the ocean.[37]

text: Around 120,000 species of fungi have been described by taxonomists,[38] but the global biodiversity of the fungus kingdom is not fully understood.[38] A 2017 estimate suggests there may be between 2.2 and 3.8 million species.[39] 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.[40] 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 the resolution and added robustness to estimates of genetic diversity within various taxonomic groups.[41]

text: Mycology

text: 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, the study of plant diseases, is closely related because many plant pathogens are fungi.[42]

text: 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).[43] 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.[44]

text: History

text: Mycology is a relatively new science that became systematic 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 is considered to be the publication of Pier Antonio Micheli's 1729 work Nova plantarum genera.[45] 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.[46] Extending the use of the binomial system of nomenclature introduced by Carl Linnaeus in his Species plantarum (1753), the Dutch Christian Hendrik Persoon (1761–1836) established the first classification of mushrooms with such skill so 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 various 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, the brothers Louis René and Charles Tulasne, Arthur H. R. Buller, Curtis G. Lloyd, and Pier Andrea Saccardo. The 20th century has seen a modernization of mycology that has come from advances in biochemistry, genetics, molecular biology, and biotechnology. The use of DNA sequencing technologies and phylogenetic analysis has provided new insights into fungal relationships and biodiversity, and has challenged traditional morphology-based groupings in fungal taxonomy.[47]

text: Morphology

text: Microscopic structures

text:
hypha
conidiophore
phialide
conidia
septa

text: 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.[48] Hyphae also sometimes fuse when they come into contact, a process called hyphal fusion (or anastamosis). These growth processes lead to the development of a mycelium, an interconnected network of hyphae.[23] 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 the cell wall giving the hypha its shape), with each compartment containing one or more nuclei; coenocytic hyphae are not compartmentalized.[49] 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.[50] Coenocytic hyphae are in essence multinucleate supercells.[51]

text: 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 the host cells to consume nutrients.[52]

text: 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.[53] 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.[54]

text: Macroscopic structures

text: 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 the identification of species or groups.[55] 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.[56]

text: The apothecium—a specialized structure important in sexual reproduction 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.[57] The fruit bodies of the basidiomycetes (basidiocarps) and some ascomycetes can sometimes grow very large, and many are well known as mushrooms.

text: Growth and physiology

text: 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.[58] Hyphae are specifically adapted for growth on solid surfaces, and to invade substrates and tissues.[59] 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.[60] The pressure generated by the appressorium, directed against the plant epidermis, can exceed 8 megapascals (1,200 psi).[60] The filamentous fungus Paecilomyces lilacinus uses a similar structure to penetrate the eggs of nematodes.[61]

text: The mechanical pressure exerted by the appressorium is generated from physiological processes that increase intracellular turgor by producing osmolytes such as glycerol.[62] Adaptations such as these are complemented by hydrolytic enzymes secreted into the environment to digest large organic molecules—such as polysaccharides, proteins, and lipids—into smaller molecules that may then be absorbed as nutrients.[63][64][65] The vast majority of filamentous fungi grow in a polar fashion (extending in one direction) by elongation at the tip (apex) of the hypha.[66] 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,[67] or growth by volume expansion during the development of mushroom stipes and other large organs.[68] Growth of fungi as multicellular structures consisting of somatic and reproductive cells—a feature independently evolved in animals and plants[69]—has several functions, including the development of fruit bodies for dissemination of sexual spores (see above) and biofilms for substrate colonization and intercellular communication.[70]

text: The fungi are traditionally considered heterotrophs, organisms that rely solely on carbon fixed by other organisms for metabolism. Fungi have evolved 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.[71][72] 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, the effects on growth rates are small, and the underlying biophysical and biochemical processes are not well known.[33] This process might bear similarity to CO2 fixation via visible light, but instead uses ionizing radiation as a source of energy.[73]

text: Reproduction

text: Fungal reproduction is complex, reflecting the differences in lifestyles and genetic makeup within this diverse kingdom of organisms.[74] 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 of a species, the teleomorph and the anamorph.[75] 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.

text: Asexual reproduction

text: 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.[76] The "Fungi imperfecti" (fungi lacking the perfect or sexual stage) or Deuteromycota comprise all the species that lack an observable sexual cycle.[77] Deuteromycota is not an accepted taxonomic clade, and is now taken to mean simply fungi that lack a known sexual stage.

text: Sexual reproduction

text: Sexual reproduction with meiosis has been directly observed in all fungal phyla except Glomeromycota [78] (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.[79][80] Mating experiments between fungal isolates may identify species on the basis of biological species concepts.[80] 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. Some species may allow mating only between individuals of opposite mating type, whereas others can mate and sexually reproduce with any other individual or itself. Species of the former mating system are called heterothallic, and of the latter homothallic.[81]

text: 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).[82]

text: In ascomycetes, dikaryotic hyphae of the hymenium (the spore-bearing tissue layer) form a characteristic hook at the hyphal septum. During cell division, 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.[83]

text: 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 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.[84] A basidiocarp is formed in which club-like structures known as basidia generate haploid basidiospores after karyogamy and meiosis.[85] The most commonly known basidiocarps are mushrooms, but they may also take other forms (see Morphology section).

text: In glomeromycetes (formerly zygomycetes), haploid hyphae of two individuals fuse, forming 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.[86]

text: Spore dispersal

text: 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.

text: Specialized mechanical and physiological mechanisms, as well as spore surface structures (such as hydrophobins), enable efficient spore ejection.[87] 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.[88] 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;[89] 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.[90] Other fungi, like the puffballs, rely on alternative mechanisms for spore release, such as external mechanical forces. The bird's nest fungi use the force of falling water drops to liberate the spores from cup-shaped fruiting bodies.[91] Another strategy is seen in the stinkhorns, a group of fungi with lively colors and putrid odor that attract insects to disperse their spores.[92]

text: Other sexual processes

text: 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.[93] 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[94] and is likely required for hybridization between species, which has been associated with major events in fungal evolution.[95]

text: Evolution

text: 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 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.[96] 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.[97] Researchers study compression fossils by dissolving the surrounding matrix with acid and then using light or scanning electron microscopy to examine surface details.[98]

text: 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.[99] Other studies (2009) estimate the arrival of fungal organisms at about 760–1060 Ma on the basis of comparisons of the rate of evolution in closely related groups.[100] 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.[101] 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.[102] Recent (2009) studies suggest that the ancestral ecological state of the Ascomycota was saprobism, and that independent lichenization events have occurred multiple times.[103]

text: It is presumed that the fungi colonized the land during the Cambrian (542–488.3 Ma), long before land plants.[104] Fossilized hyphae and spores recovered from 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.[105] Prototaxites, which was probably a fungus or lichen, would have been the tallest organism of the late Silurian. Fungal fossils do not become common and uncontroversial until the early Devonian (416–359.2 Ma), when they occur abundantly in the Rhynie chert, mostly as Zygomycota and Chytridiomycota.[104][106][107] At about this same time, approximately 400 Ma, the Ascomycota and Basidiomycota diverged,[108] and all modern classes of fungi were present by the Late Carboniferous (Pennsylvanian, 318.1–299 Ma).[109]

text: Lichen-like fossils have been found in the Doushantuo Formation in southern China dating back to 635–551 Ma.[110] Lichens formed a component of the early terrestrial ecosystems, and the estimated age of the oldest terrestrial lichen fossil is 400 Ma;[111] this date corresponds to the age of the oldest known sporocarp fossil, a Paleopyrenomycites species found in the Rhynie Chert.[112] The oldest fossil with microscopic features resembling modern-day basidiomycetes is Palaeoancistrus, found permineralized with a fern from the Pennsylvanian.[113] Rare in the fossil record are the Homobasidiomycetes (a taxon roughly equivalent to 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.[114][115]

text: 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 the dominant life form at this time, representing nearly 100% of the available fossil record for this period.[116] However, the relative proportion of fungal spores relative to spores formed by algal species is difficult to assess,[117] the spike did not appear worldwide,[118][119] and in many places it did not fall on the Permian–Triassic boundary.[120]

text: Taxonomy

text: 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.[121] Analyses using molecular phylogenetics support a monophyletic origin of the Fungi.[41] The taxonomy of the Fungi is in a state of constant flux, especially due to recent 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.[122]

text: 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.[41][123] Fungal species can also have multiple scientific names depending on their life cycle and mode (sexual or asexual) of reproduction. Web sites such as Index Fungorum and ITIS list current names of fungal species (with cross-references to older synonyms).

text: 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.[41] 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 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[124] and "The Mycota: A Comprehensive Treatise on Fungi as Experimental Systems for Basic and Applied Research".[125] The lengths of the branches are not proportional to evolutionary distances.

text:
Holomycota

Cristidiscoidea

Opisthosporidia

Aphelida

Cryptomycota

Microsporidia

Metchnikovellea

Microsporea

Eumycota

Chytridiomycota

Neocallimastigomycetes

Chytridiomycetes

Hyaloraphidiomycetes

Monoblepharidomycetes

Blastocladiomycota

Blastocladiomycetes

Olpidiales

Entomophthoromycota

Basidiobolomycetes

Neozygitomycetes

Entomophthoromycetes

Kickxellomycota

Zoopagomycetes

Kickxellomycetes

Mucoromycota

Mortierellomycetes

Mucoromycetes

Symbiomycota

Glomeromycota

Glomeromycetes

Dikarya

Entorrhizomycota

Entorrhizomycetes

Basidiomycota

Ascomycota

text:
Basidiomycota

Pucciniomycotina

Tritirachiomycetes

Mixiomycetes

Agaricostilbomycetes

Cystobasidiomycetes

Classiculaceae

Microbotryomycetes

Cryptomycocolacomycetes

Atractiellomycetes

Pucciniomycetes

Orthomycotina

Ustilaginomycotina

Monilielliomycetes

Malasseziomycetes

Ustilaginomycetes

Exobasidiomycetes

Agaricomycotina

Wallemiomycetes

Bartheletiomycetes

Tremellomycetes

Dacrymycetes

Agaricomycetes

text:
Ascomycota

Taphrinomycotina

Neolectomycetes

Taphrinomycetes

Schizosaccharomyceta

Archaeorhizomycetes

Pneumocystidomycetes

Schizosaccharomycetes

Saccharomyceta

Saccharomycotina

Saccharomycetes

Pezizomycotina

?Thelocarpales

?Vezdaeales

?Lahmiales

?Triblidiales

Orbiliomycetes

Pezizomycetes

Leotiomyceta

Sordariomyceta

Xylonomycetes

Geoglossomycetes

Leotiomycetes

Laboulbeniomycetes

Sordariomycetes

Dothideomyceta

Coniocybomycetes

Lichinomycetes

Eurotiomycetes

Lecanoromycetes

Collemopsidiales

Arthoniomycetes

Dothideomycetes

text: Taxonomic groups

text: The major phyla (sometimes called divisions) of fungi have been classified mainly on the basis of characteristics of their sexual reproductive structures. Currently, seven phyla are proposed: Microsporidia, Chytridiomycota, Blastocladiomycota, Neocallimastigomycota, Glomeromycota, Ascomycota, and Basidiomycota.[41]

text: Phylogenetic analysis has demonstrated that the Microsporidia, unicellular parasites of animals and protists, are fairly recent and highly derived endobiotic fungi (living within the tissue of another species).[101][126] One 2006 study concludes that the Microsporidia are a sister group to the true fungi; that is, they are each other's closest evolutionary relative.[127] Hibbett and colleagues suggest that this analysis does not clash with their classification of the Fungi, and although the Microsporidia are elevated to phylum status, it is acknowledged that further analysis is required to clarify evolutionary relationships within this group.[41]

text: The Chytridiomycota are commonly known as chytrids. These fungi are distributed worldwide. Chytrids and their close relatives Neocallimastigomycota and Blastocladiomycota (below) are the only fungi with active motility, producing zoospores that are capable of active movement through aqueous phases with a single flagellum, leading early taxonomists to classify them as protists. Molecular phylogenies, inferred from rRNA sequences in ribosomes, suggest that the Chytrids are a basal group divergent from the other fungal phyla, consisting of four major clades with suggestive evidence for paraphyly or possibly polyphyly.[128]

text: The Blastocladiomycota were previously considered a taxonomic clade within the Chytridiomycota. Recent molecular data and ultrastructural characteristics, however, place the Blastocladiomycota as a sister clade to the Zygomycota, Glomeromycota, and Dikarya (Ascomycota and Basidiomycota). The blastocladiomycetes are saprotrophs, feeding on decomposing organic matter, and they are parasites of all eukaryotic groups. Unlike their close relatives, the chytrids, most of which exhibit zygotic meiosis, the blastocladiomycetes undergo sporic meiosis.[101]

text: The Neocallimastigomycota were earlier placed in the phylum Chytridomycota. Members of this small phylum are anaerobic organisms, living in the digestive system of larger herbivorous mammals and in other terrestrial and aquatic environments enriched in cellulose (e.g., domestic waste landfill sites).[129] They lack mitochondria but contain hydrogenosomes of mitochondrial origin. As in the related chrytrids, neocallimastigomycetes form zoospores that are posteriorly uniflagellate or polyflagellate.[41]

text: Members of the Glomeromycota form arbuscular mycorrhizae, a form of mutualist symbiosis wherein fungal hyphae invade plant root cells and both species benefit from the resulting increased supply of nutrients. All known Glomeromycota species reproduce asexually.[78] The symbiotic association between the Glomeromycota and plants is ancient, with evidence dating to 400 million years ago.[130] Formerly part of the Zygomycota (commonly known as 'sugar' and 'pin' molds), the Glomeromycota were elevated to phylum status in 2001 and now replace the older phylum Zygomycota.[131] Fungi that were placed in the Zygomycota are now being reassigned to the Glomeromycota, or the subphyla incertae sedis Mucoromycotina, Kickxellomycotina, the Zoopagomycotina and the Entomophthoromycotina.[41] Some well-known examples of fungi formerly in the Zygomycota include black bread mold (Rhizopus stolonifer), and Pilobolus species, capable of ejecting spores several meters through the air.[132] Medically relevant genera include Mucor, Rhizomucor, and Rhizopus.

text: The Ascomycota, commonly known as sac fungi or ascomycetes, constitute the largest taxonomic group within the Eumycota.[40] These fungi form meiotic spores called ascospores, which are enclosed in a special sac-like structure called an ascus. This phylum includes morels, a few mushrooms and truffles, unicellular yeasts (e.g., of the genera Saccharomyces, Kluyveromyces, Pichia, and Candida), and many filamentous fungi living as saprotrophs, parasites, and mutualistic symbionts (e.g. lichens). Prominent and important genera of filamentous ascomycetes include Aspergillus, Penicillium, Fusarium, and Claviceps. Many ascomycete species have only been observed undergoing asexual reproduction (called anamorphic species), but analysis of molecular data has often been able to identify their closest teleomorphs in the Ascomycota.[133] Because the products of meiosis are retained within the sac-like ascus, ascomycetes have been used for elucidating principles of genetics and heredity (e.g., Neurospora crassa).[134]

text: Members of the Basidiomycota, commonly known as the club fungi or basidiomycetes, produce meiospores called basidiospores on club-like stalks called basidia. Most common mushrooms belong to this group, as well as rust and smut fungi, which are major pathogens of grains. Other important basidiomycetes include the maize pathogen Ustilago maydis,[135] human commensal species of the genus Malassezia,[136] and the opportunistic human pathogen, Cryptococcus neoformans.[137]

text: Fungus-like organisms

text: Because of similarities in morphology and lifestyle, the slime molds (mycetozoans, plasmodiophorids, acrasids, Fonticula and labyrinthulids, now in Amoebozoa, Rhizaria, Excavata, Opisthokonta and Stramenopiles, respectively), water molds (oomycetes) and hyphochytrids (both Stramenopiles) were formerly classified in the kingdom Fungi, in groups like Mastigomycotina, Gymnomycota and Phycomycetes. The slime molds were studied also as protozoans, leading to a ambiregnal, duplicated taxonomy.

text: Unlike true fungi, the cell walls of oomycetes contain cellulose and lack chitin. Hyphochytrids have both chitin and cellulose. Slime molds lack a cell wall during the assimilative phase (except labyrinthulids, which have a wall of scales), and ingest nutrients by ingestion (phagocytosis, except labyrinthulids) rather than absorption (osmotrophy, as fungi, labyrinthulids, oomycetes and hyphochytrids). Neither water molds nor slime molds are closely related to the true fungi, and, therefore, taxonomists no longer group them in the kingdom Fungi. Nonetheless, studies of the oomycetes and myxomycetes are still often included in mycology textbooks and primary research literature.[138]

text: The Eccrinales and Amoebidiales are opisthokont protists, previously thought to be zygomycete fungi. Other groups now in Opisthokonta (e.g., Corallochytrium, Ichthyosporea) were also at given time classified as fungi. The genus Blastocystis, now in Stramenopiles, was originally classified as a yeast. Ellobiopsis, now in Alveolata, was considered a chytrid. The bacteria were also included in fungi in some classifications, as the group Schizomycetes.

text: The Rozellida clade, including the "ex-chytrid" Rozella, is a genetically disparate group known mostly from environmental DNA sequences that is a sister group to fungi. Members of the group that have been isolated lack the chitinous cell wall that is characteristic of fungi.

text: The nucleariids, protists currently grouped in the Choanozoa (Opisthokonta), may be the next sister group to the eumycete clade, and as such could be included in an expanded fungal kingdom.[121]

text: Many Actinomycetales (Actinobacteria), a group with many filamentous bacteria, were also long believed to be fungi.

text: Ecology

text: Although often inconspicuous, fungi occur in every environment on Earth and play very important roles in most ecosystems. Along with bacteria, fungi are the major decomposers in most terrestrial (and some aquatic) ecosystems, and therefore play a critical role in biogeochemical cycles[139] and in many food webs. As decomposers, they play an essential role in nutrient cycling, especially as saprotrophs and symbionts, degrading organic matter to inorganic molecules, which can then re-enter anabolic metabolic pathways in plants or other organisms.[140][141]

text: Symbiosis

text: Many fungi have important symbiotic relationships with organisms from most if not all Kingdoms.[142][143][144] These interactions can be mutualistic or antagonistic in nature, or in the case of commensal fungi are of no apparent benefit or detriment to the host.[145][146][147]

text: With plants

text: Mycorrhizal symbiosis between plants and fungi is one of the most well-known plant–fungus associations and is of significant importance for plant growth and persistence in many ecosystems; over 90% of all plant species engage in mycorrhizal relationships with fungi and are dependent upon this relationship for survival.[148]

text: The mycorrhizal symbiosis is ancient, dating to at least 400 million years ago.[130] It often increases the plant's uptake of inorganic compounds, such as nitrate and phosphate from soils having low concentrations of these key plant nutrients.[140][149] The fungal partners may also mediate plant-to-plant transfer of carbohydrates and other nutrients. Such mycorrhizal communities are called "common mycorrhizal networks".[150] A special case of mycorrhiza is myco-heterotrophy, whereby the plant parasitizes the fungus, obtaining all of its nutrients from its fungal symbiont.[151] Some fungal species inhabit the tissues inside roots, stems, and leaves, in which case they are called endophytes.[152] Similar to mycorrhiza, endophytic colonization by fungi may benefit both symbionts; for example, endophytes of grasses impart to their host increased resistance to herbivores and other environmental stresses and receive food and shelter from the plant in return.[153]

text: With algae and cyanobacteria

text: Lichens are a symbiotic relationship between fungi and photosynthetic algae or cyanobacteria. The photosynthetic partner in the relationship is referred to in lichen terminology as a "photobiont". The fungal part of the relationship is composed mostly of various species of ascomycetes and a few basidiomycetes.[154] Lichens occur in every ecosystem on all continents, play a key role in soil formation and the initiation of biological succession,[155] and are prominent in some extreme environments, including polar, alpine, and semiarid desert regions.[156] They are able to grow on inhospitable surfaces, including bare soil, rocks, tree bark, wood, shells, barnacles and leaves.[157] As in mycorrhizas, the photobiont provides sugars and other carbohydrates via photosynthesis to the fungus, while the fungus provides minerals and water to the photobiont. The functions of both symbiotic organisms are so closely intertwined that they function almost as a single organism; in most cases the resulting organism differs greatly from the individual components. Lichenization is a common mode of nutrition for fungi; around 20% of fungi—between 17,500 and 20,000 described species—are lichenized.[158] Characteristics common to most lichens include obtaining organic carbon by photosynthesis, slow growth, small size, long life, long-lasting (seasonal) vegetative reproductive structures, mineral nutrition obtained largely from airborne sources, and greater tolerance of desiccation than most other photosynthetic organisms in the same habitat.[159]

text: With insects

text: Many insects also engage in mutualistic relationships with fungi. Several groups of ants cultivate fungi in the order Agaricales as their primary food source, while ambrosia beetles cultivate various species of fungi in the bark of trees that they infest.[160] Likewise, females of several wood wasp species (genus Sirex) inject their eggs together with spores of the wood-rotting fungus Amylostereum areolatum into the sapwood of pine trees; the growth of the fungus provides ideal nutritional conditions for the development of the wasp larvae.[161] At least one species of stingless bee has a relationship with a fungus in the genus Monascus, where the larvae consume and depend on fungus transferred from old to new nests.[162] Termites on the African savannah are also known to cultivate fungi,[142] and yeasts of the genera Candida and Lachancea inhabit the gut of a wide range of insects, including neuropterans, beetles, and cockroaches; it is not known whether these fungi benefit their hosts.[163] Fungi ingrowing dead wood are essential for xylophagous insects (e.g. woodboring beetles).[164][non-primary source needed] They deliver nutrients needed by xylophages to nutritionally scarce dead wood.[165][non-primary source needed] Thanks to this nutritional enrichment the larvae of woodboring insect is able to grow and develop to adulthood.[164] The larvae of many families of fungicolous flies, particularly those within the superfamily Sciaroidea such as the Mycetophilidae and some Keroplatidae feed on fungal fruiting bodies and sterile mycorrhizae.[166]

text: As pathogens and parasites

text: Many fungi are parasites on plants, animals (including humans), and other fungi. Serious pathogens of many cultivated plants causing extensive damage and losses to agriculture and forestry include the rice blast fungus Magnaporthe oryzae,[167] tree pathogens such as Ophiostoma ulmi and Ophiostoma novo-ulmi causing Dutch elm disease[168] and Cryphonectria parasitica responsible for chestnut blight,[169] and plant pathogens in the genera Fusarium, Ustilago, Alternaria, and Cochliobolus.[146] Some carnivorous fungi, like Paecilomyces lilacinus, are predators of nematodes, which they capture using an array of specialized structures such as constricting rings or adhesive nets.[170]

text: Some fungi can cause serious diseases in humans, several of which may be fatal if untreated. These include aspergillosis, candidiasis, coccidioidomycosis, cryptococcosis, histoplasmosis, mycetomas, and paracoccidioidomycosis. Furthermore, persons with immuno-deficiencies are particularly susceptible to disease by genera such as Aspergillus, Candida, Cryptoccocus,[147][171][172] Histoplasma,[173] and Pneumocystis.[174] Other fungi can attack eyes, nails, hair, and especially skin, the so-called dermatophytic and keratinophilic fungi, and cause local infections such as ringworm and athlete's foot.[175] Fungal spores are also a cause of allergies, and fungi from different taxonomic groups can evoke allergic reactions.[176]

text: As targets of mycoparasites

text: The organisms which parasitize fungi are known as mycoparasitic organisms. Certain species of the Pythium genus, which are oomycetes, have potential as biocontrol agents against certain fungi.[177] Fungi can also act as mycoparasites or antagonists of other fungi, such as Hypomyces chrysospermus, which grows on bolete mushrooms.

text: Fungi can become the target of infection by mycoviruses.

text: Mycotoxins

text: Many fungi produce biologically active compounds, several of which are toxic 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 amatoxins in some Amanita mushrooms, and ergot alkaloids, 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 of the ergot fungus, Claviceps purpurea.[178] Other notable mycotoxins include the aflatoxins, which are insidious liver toxins and highly carcinogenic metabolites produced by certain Aspergillus species often growing in or on grains and nuts consumed by humans, ochratoxins, patulin, and trichothecenes (e.g., T-2 mycotoxin) and fumonisins, which have significant impact on human food supplies or animal livestock.[179]

text: 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.[29] Mycotoxins may provide fitness benefits in terms of physiological adaptation, competition with other microbes and fungi, and protection from consumption (fungivory).[180][181] Many fungal secondary metabolites (or derivatives) are used medically, as described under Human Use below.

text: Pathogenic mechanisms

text: 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 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. The response protects U. maydis from the host defense, and is necessary for the pathogen’s virulence.[182] Furthermore, U. maydis has a well-established recombinational DNA repair system which acts during mitosis and meiosis.[183] The system may assist the pathogen in surviving DNA damage arising from the host plant’s oxidative defensive response to infection.[184]

text: 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 macrophages.[185] Some C. neoformans can survive inside 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.[185] Another mechanism involves meiosis. 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 blastospores. The diploid nuclei of blastospores can undergo meiosis, including recombination, to form haploid basidiospores that can be dispersed.[186] This process is referred to as monokaryotic fruiting. this process requires a gene called DMC1, which is a conserved homologue of genes recA 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.[184][186]

text: Human use

text: The human use of fungi for food preparation or preservation and other purposes is extensive and has a long history. Mushroom farming and mushroom gathering are large industries in many countries. The study of the historical uses and sociological impact of fungi is known as ethnomycology. Because of the capacity of this group to produce an enormous range of natural products with antimicrobial or other biological activities, many species have long been used or are being developed for industrial production of antibiotics, vitamins, and anti-cancer and cholesterol-lowering drugs. More recently, methods have been developed for genetic engineering of fungi,[187] enabling metabolic engineering of fungal species. For example, genetic modification of yeast species[188]—which are easy to grow at fast rates in large fermentation vessels—has opened up ways of pharmaceutical production that are potentially more efficient than production by the original source organisms.[189]

text: Therapeutic uses

text: Modern chemotherapeutics

text: Many species produce metabolites that are major sources of pharmacologically active drugs. Particularly important are the antibiotics, including the penicillins, a structurally related group of β-lactam antibiotics that are synthesized from small peptides. Although naturally occurring penicillins such as penicillin G (produced by Penicillium chrysogenum) have a relatively narrow spectrum of biological activity, a wide range of other penicillins can be produced by chemical modification of the natural penicillins. Modern penicillins are semisynthetic compounds, obtained initially from fermentation cultures, but then structurally altered for specific desirable properties.[190] Other antibiotics produced by fungi include: ciclosporin, commonly used as an immunosuppressant during transplant surgery; and fusidic acid, used to help control infection from methicillin-resistant Staphylococcus aureus bacteria.[191] Widespread use of antibiotics for the treatment of bacterial diseases, such as tuberculosis, syphilis, leprosy, 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, and at low concentrations as quorum-sensing molecules for intra- or interspecies signaling.[192] Other drugs produced by fungi include griseofulvin isolated from Penicillium griseofulvum, used to treat fungal infections,[193] and statins (HMG-CoA reductase inhibitors), used to inhibit cholesterol synthesis. Examples of statins found in fungi include mevastatin from Penicillium citrinum and lovastatin from Aspergillus terreus and the oyster mushroom.[194]

text: Traditional and folk medicine

text: Certain mushrooms enjoy usage as therapeutics in folk medicines, such as Traditional Chinese medicine. Notable medicinal mushrooms with a well-documented history of use include Agaricus subrufescens,[195][196] Ganoderma lucidum,[197] and Ophiocordyceps sinensis.[198] Research has identified compounds produced by these and other fungi that have inhibitory biological effects against viruses[199][200] and cancer cells.[195][201] Specific metabolites, such as polysaccharide-K, ergotamine, and β-lactam antibiotics, are routinely used in clinical medicine. The shiitake mushroom is a source of lentinan, a clinical drug approved for use in cancer treatments in several countries, including Japan.[202][203] In Europe and Japan, polysaccharide-K (brand name Krestin), a chemical derived from Trametes versicolor, is an approved adjuvant for cancer therapy.[204]

text: Cultured foods

text: Baker's yeast or Saccharomyces cerevisiae, a unicellular fungus, is used to make bread and other wheat-based products, such as pizza dough and dumplings.[205] Yeast species of the genus Saccharomyces are also used to produce alcoholic beverages through fermentation.[206] Shoyu koji mold (Aspergillus oryzae) is an essential ingredient in brewing Shoyu (soy sauce) and sake, and the preparation of miso,[207] while Rhizopus species are used for making tempeh.[208] Several of these fungi are domesticated 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.[209] Quorn, a meat substitute, is made from Fusarium venenatum.[210]

text: Edible and poisonous species

text: Edible mushrooms are well-known examples of fungi. Many are commercially raised, but others must be harvested from the wild. Agaricus bisporus, sold as button mushrooms when small or Portobello mushrooms when larger, is a commonly eaten species, 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 stores and markets, including straw mushrooms (Volvariella volvacea), oyster mushrooms (Pleurotus ostreatus), shiitakes (Lentinula edodes), and enokitake (Flammulina spp.).[211]

text: There are many more mushroom species that are harvested from the wild for personal consumption or commercial sale. Milk mushrooms, morels, chanterelles, truffles, black trumpets, and porcini mushrooms (Boletus edulis) (also known as king boletes) demand a high price on the market. They are often used in gourmet dishes.[212]

text: 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 color in cheeses such as Stilton or Roquefort, which are made by inoculation with Penicillium roqueforti.[213] Molds used in cheese production are non-toxic and are thus safe for human consumption; however, mycotoxins (e.g., aflatoxins, roquefortine C, patulin, or others) may accumulate because of growth of other fungi during cheese ripening or storage.[214]

text: Many mushroom species are poisonous to humans, with toxicities ranging from slight digestive problems or allergic reactions as well as hallucinations to severe organ failures and death. Genera with mushrooms containing deadly toxins include Conocybe, Galerina, Lepiota, and, the most infamous, Amanita.[215] The latter genus includes the destroying angel (A. virosa) and the death cap (A. phalloides), the most common cause of deadly mushroom poisoning.[216] The false morel (Gyromitra esculenta) is occasionally considered a delicacy when cooked, yet can be highly toxic when eaten raw.[217] Tricholoma equestre was considered edible until it was implicated in serious poisonings causing rhabdomyolysis.[218] Fly agaric mushrooms (Amanita muscaria) also cause occasional non-fatal poisonings, mostly as a result of ingestion for its hallucinogenic properties. Historically, fly agaric was used by different peoples in Europe and Asia and its present usage for religious or shamanic purposes is reported from some ethnic groups such as the Koryak people of north-eastern Siberia.[219]

text: 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.[220][221]

text: Pest control

text: In agriculture, fungi may be useful if they actively compete for nutrients and space with pathogenic microorganisms such as bacteria or other fungi via the competitive exclusion principle,[222] or if they are parasites of these pathogens. For example, certain species may be used to eliminate or suppress the growth of harmful plant pathogens, such as insects, mites, weeds, nematodes, and other fungi that cause diseases of important crop plants.[223] This has generated strong interest in practical applications that use these fungi in the biological control of these agricultural pests. Entomopathogenic fungi can be used as biopesticides, as they actively kill insects.[224] Examples that have been used as biological insecticides are Beauveria bassiana, Metarhizium spp, Hirsutella spp, Paecilomyces (Isaria) spp, and Lecanicillium lecanii.[225][226] Endophytic fungi of grasses of the genus Neotyphodium, such as N. coenophialum, produce alkaloids that are toxic to a range of invertebrate and vertebrate herbivores. These alkaloids protect grass plants from herbivory, but several endophyte alkaloids can poison grazing animals, such as cattle and sheep.[227] Infecting cultivars of pasture or forage grasses with Neotyphodium 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.[228]

text: Bioremediation

text: Certain fungi, in particular "white rot" fungi, can degrade insecticides, herbicides, pentachlorophenol, creosote, coal tars, and heavy fuels and turn them into carbon dioxide, water, and basic elements.[229] Fungi have been shown to biomineralize uranium oxides, suggesting they may have application in the bioremediation of radioactively polluted sites.[230][231][232]

text: Model organisms

text: 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 to test their biochemical theories.[233] Other important model fungi are Aspergillus nidulans and the yeasts Saccharomyces cerevisiae and Schizosaccharomyces pombe, each of which with a long history of use to investigate issues in eukaryotic cell biology and genetics, such as cell cycle regulation, chromatin structure, and gene regulation. Other fungal models have more recently emerged that address specific biological questions relevant to medicine, plant pathology, and industrial uses; examples include Candida albicans, a dimorphic, opportunistic human pathogen,[234] Magnaporthe grisea, a plant pathogen,[235] and Pichia pastoris, a yeast widely used for eukaryotic protein production.[236]

text: Others

text: Fungi are used extensively to produce industrial chemicals like citric, gluconic, lactic, and malic acids,[237] and industrial enzymes, such as lipases used in biological detergents,[238] cellulases used in making cellulosic ethanol[239] and stonewashed jeans,[240] and amylases,[241] invertases, proteases and xylanases.[242] Several species, most notably Psilocybin mushrooms (colloquially known as magic mushrooms), are ingested for their psychedelic properties, both recreationally and religiously.


document_title: God of War (2018 video game)

question_text: who voices the kid in the new god of war

root

text: God of War (2018 video game)

text: Gameplay[edit]

text: The gameplay is vastly different from the previous installments, as it was rebuilt from the ground up.[6] Although the previous main installment, Ascension (2013), introduced multiplayer to the series, this installment is single-player-only.[4] The game features a third-person, over-the-shoulder free camera, a departure from the previous installments, which featured a third-person, fixed cinematic camera (with the exception of 2007's 2D side-scroller Betrayal).[7] Cinematographically, the game is presented in a continuous shot, with no camera cuts.[8] The game is open, but it is not open-world.[6] Due to it being open, players can fast travel to different locations through Mystic Gateways.[9] As the ability to swim was cut from the game,[10] players instead use a boat to traverse bodies of water.[9] Just like previous entries, there are puzzles for players to solve to progress through parts of the game. Enemies in the game stem from Norse mythology, such as variants of trolls, ogres, dark elves and their king, wolves, wulvers, nightmares, draugrs,[11] tatzelwurms, as well as Gullveig and the revenants, beings warped by seiðr magic, among other original creatures.[12] Valkyries appear as optional boss battles, and players can free the dragons Fáfnir, Otr, and Reginn—dwarfs that were turned into dragons—in addition to battling a dragon called Hræzlyr.[9]

tag: p

text: A major change is that Kratos no longer uses his signature double-chained blades, the Blades of Chaos, as his default weapon. Instead, he uses a magical battle axe, called the Leviathan Axe,[13] which is infused with ice elemental magic. The axe can be thrown at enemies and magically summoned back to his hand (similar to Thor's hammer Mjölnir). Larger enemies have precision targets and throwing the axe at those targets stuns the enemy. The weapon can also be thrown at environmental objects to trigger a damaging explosion and it can freeze objects and some enemies in place for puzzle solving until the axe is summoned back to Kratos' hand. The axe has standard light and heavy attacks, and over time, it can be upgraded with runes to allow for special runic attacks, with one slot being for a light runic attack and the other for heavy. This provides players with a variety of options to cater to their own play style. For example, one of the light runic attacks allows Kratos to charge the axe and let out a burst of energy and one of the heavy runic attacks allows Kratos to summon a swirling ice storm.[14] Another new weapon that Kratos utilizes is the Guardian Shield. When not in use, it folds up and appears like a vambrace on Kratos' left forearm. When summoned, the shield can be used offensively and defensively, similar to the Golden Fleece in previous games.[15] Kratos also utilizes hand-to-hand combat, a feature originally introduced in Ascension.[7] The Blades of Chaos are acquired late into the game via a plot device and perform similarly as they did in previous installments, but can also be upgraded with light and heavy runic attacks.

text: Similar to previous games, there is a "Rage" ability called Spartan Rage. Like the previous versions, the Rage ability has a meter that gradually fills during combat. With this ability, Kratos uses powerful bare-handed attacks, as opposed to weapons, to greatly damage enemies. The game also features elements similar to role-playing video games (RPGs).[7][16] There are crafting resources for the player to find that allow them to create new or upgrade existing armor with better perks.[17] Players also accumulate a currency called Hacksilver, a key component in crafting and purchasing new items. Experience points (XP) are used for learning new combat skills.[16] Throughout the game world, players find chests containing random items, such as Hacksilver and enchantments for improving armor and weapons, as well as two special items, Iðunn's Apples and Horns of Blood Mead, which increase the maximum length of the health and rage meters, respectively. Quick time events have changed from previous games.[6] Enemies display two meters above their heads, one for health and the other for stun. Filling up the stun meter helps to defeat more difficult enemies. When the stun meter is full, a grab-prompt will appear. Depending on the enemy, Kratos may rip it in half or grab them and throw them into other enemies, among other possible outcomes.[18]

text: Although the game is played entirely as Kratos,[19] there are times when the player may choose to passively control Kratos' son, Atreus. One button is dedicated to Atreus and its use depends on the context. For example, if the player needs assistance, they can look at an enemy, press the button, and Atreus will use his Talon Bow to shoot arrows at the enemy.[20] The arrows have little effect on an enemy's health, but do increase the stun meter.[18] Over the course of the game, Atreus helps in combat, traversal, exploration, and puzzle-solving. When facing a large number of enemies, he distracts the weaker ones as Kratos fights the stronger ones.[17] If too many enemies gang up on Atreus, he is knocked out for the remainder of that combat. Just like Kratos, Atreus acquires new skills, armor, special arrows, such as lightning arrows, as well as runic attacks for his Talon Bow, but it only has one slot instead of two. Atreus' runic attacks summon different spectral animals with different abilities. For example, one summons a wolf that attacks enemies, while another summons the squirrel Ratatoskr that will dig up orbs to replenish Kratos' health or rage meters.[21]

text: Synopsis[edit]

text: Setting[edit]

text: While the first seven games were loosely based on Greek mythology, this installment takes the series to Norse mythology,[22] but predates the Vikings.[20][23] Six of the nine realms of Norse mythology can be explored. The majority of the game takes place in the realm of Midgard, inhabited by humans and other creatures and is the same realm that the Greek world had existed in. As more creatures began appearing, many humans fled. Other realms visited as part of the story include Alfheim, the mystical home of the light and dark elves, Helheim, the icy land of the dead, and Jötunheim, the mountainous land of the giants. Other explorable realms include Niflheim, a realm of poisonous fog with a maze-like structure of rewards, and the fire realm Muspelheim, featuring the six Trials of Muspelheim; completing each trial grants rewards and advances Kratos and Atreus closer to the top of a large volcano. Access to the other three realms—Asgard, home of the Æsir gods, Vanaheim, home of the Vanir gods, and Svartalfheim, home of the dwarves—have been blocked by the ruler of Asgard and the Æsir gods, Odin.[24] At the center of the realms is the mythical tree Yggdrasil, which connects each realm together. Although each realm is a different world, they simultaneously exist in the same space. Travel to and from realms can be done by the use of the Bifröst from a root of Yggdrasil contained within a temple located at the center of the Lake of the Nine. The temple was created by the now dead Týr, a peaceful God of War who had traveled to other lands and learned about their mythologies; Odin had him killed as he believed Týr was secretly aiding the giants and would try to overthrow him.[22]

text: Characters[edit]

text: The protagonists of the game are Kratos (voiced by Christopher Judge) and his young son Atreus (Sunny Suljic). Kratos is a warrior originally from Sparta who became the Greek God of War and is the son of Zeus. After ending up in Midgard, he met his second and now deceased wife, Laufey (addressed as Faye), and they bore their son, Atreus, who does not know about Kratos' past or his divine nature, but can hear other beings' thoughts. The main antagonist is the Æsir god Baldur (Jeremy Davies), the brother of Thor, whose sons Modi and Magni (Nolan North and Troy Baker, respectively) assist Baldur. His parents are Odin and the Vanir god Freya (Danielle Bisutti). Freya tried leaving Odin, as she did not truly love him, and Odin had her banished to Midgard, after which she became known as the Witch of the Woods. To protect her son from a prophecy that foretold his death, Freya cast a spell of immortality on Baldur, which also caused him to not feel pain or any feeling of pleasure, which he resented her for. The only thing that she could not prevent from breaking the spell was mistletoe, which she kept secret. Other characters include Mímir (Alastair Duncan), who claims to be the smartest man alive, and the Huldra Brothers Brok (Roger Craighead) and Sindri (Adam J. Harrington), a pair of dwarves who appear at various points in the world and assist Kratos and Atreus with forging new gear. Their weapons, including Thor's hammer Mjölnir, were used by the Æsir gods and they also had forged Kratos' Leviathan Axe, which originally belonged to Faye, who also gifted Kratos his Guardian Shield.[13] The spirit of the Greek goddess Athena (Carole Ruggier) makes a cameo appearance, and Zeus (Corey Burton) appears as an illusion to Kratos in Helheim.

text: Plot[edit]

text: Many years have passed since Kratos took his vengeance against the Olympian gods,[b] and he now lives with his young son Atreus in Midgard. The game opens following the death of Kratos' second wife and Atreus' mother, Faye, whose last wish was for her ashes to be spread at the highest peak of the nine realms. Before beginning their journey, Kratos is confronted by a mysterious man with godlike powers. After seemingly killing him, Kratos and Atreus set out on their journey.

text: Reaching the Lake of the Nine, Kratos and Atreus encounter the friendly World Serpent, Jörmungandr. After running into impenetrable black mist which can only be extinguished with the Light of Alfheim, they receive aid from the Witch of the Woods to use the Bifröst in order to travel to Alfheim and secure the Light. Upon vanquishing the mist and reaching Midgard's peak, they overhear a conversation between the mysterious man, revealed to be Baldur, his nephews Modi and Magni, and the imprisoned Mímir. After they leave, Kratos and Atreus confront Mímir, who reveals that their goal is actually in Jötunheim, but travel there has been blocked to keep out Odin and Thor. Mímir, however, knows another passage. He instructs Kratos to cut off his head and have it revived by the Witch of the Woods, revealed to be Freya. Kratos immediately distrusts her, but both Freya and Mímir warn him that he must tell Atreus about his true nature.

text: Kratos, Atreus, and Mímir's head journey to collect needed components to open Jötunheim's portal when they are attacked by Modi and Magni. After Kratos kills Magni, Modi flees, but later returns and ambushes them. Atreus collapses ill, which Mímir and Freya explain is a mental contradiction of a god believing himself to be mortal. She tells Kratos that he must retrieve the heart of the Keeper of the Bridge of the Damned in Helheim, but his Leviathan Axe is useless there. Kratos then returns home to unearth his old weapons, the Blades of Chaos, and is haunted by Athena's spirit, who goads him about his past. After retrieving the heart, he has a haunting vision of Zeus. Atreus is cured and Kratos tells him that he is a god. Atreus then becomes increasingly arrogant on their journey, and he murders a weakened Modi, despite Kratos ordering not to. At Midgard's peak, they are ambushed by Baldur, resulting in Jötunheim's portal being destroyed and the group falling into Helheim.

text: Atreus makes amends with Kratos and they find out about Freya and Baldur's familial relationship. Returning to Midgard, Mímir realizes there is another way to reach Jötunheim, but it requires recovering his missing eye. After obtaining it from Jörmungandr's belly, who had inadvertently swallowed it when he ate a statue of Thor, they are attacked by Baldur again, but Freya intervenes in an attempt to protect her son. During the fight, Baldur is pierced by Atreus' mistletoe arrow, breaking Freya's spell on him. Baldur is defeated, and although Kratos gives him an opportunity to retreat, he instead attempts to strangle Freya, forcing Kratos to kill him. A grieving Freya swears revenge on Kratos and taunts him about hiding his true nature from Atreus. Kratos finally tells Atreus about his past and how he had killed his fellow Greek gods, including his father, Zeus. Atreus laments if patricide is all that gods are good for. Kratos tells Atreus that they should both learn from their experiences and not repeat the mistakes of their predecessors. A silent Freya leaves with Baldur's corpse and Mímir hopefully suggests that she will eventually move on from the tragedy and that Kratos did the right thing.

text: In Jötunheim, they find a temple with a mural depicting their adventures, showing that the giants had foreseen everything that would happen and vaguely foretold what was to come. In addition, they discover that Faye was a giant who decided to stay behind in Midgard, making Atreus a hybrid of giant, god, and mortal. Their fight with Baldur was shown, revealing he was actually after Faye the whole time. It is also revealed that Atreus was named Loki by his mother. Wondering if Faye planned this in advance, Kratos and Atreus fulfill their promise and spread her ashes at the peak. Afterwards, Kratos reveals to Atreus that his given name was also the name of a compassionate Spartan comrade. When they return to Midgard, Mímir warns them that the three-year long Fimbulwinter has started, meaning Ragnarök is soon to follow, which was not supposed to occur for another hundred years.

text: In the game's secret ending, Kratos and Atreus return home and slumber. Atreus has a vision that at the end of Fimbulwinter, Thor will arrive at their home to confront them.

text: Development[edit]

text: At the first annual PlayStation Experience on December 6, 2014, Santa Monica Studios's creative director Cory Barlog confirmed that a new God of War was in very early development. He said that the game would not be a prequel, but it might be a reboot.[25] In April 2016, Polygon reported that concept art of the next installment had been leaked. The images showed Kratos in the world of Norse mythology; a concept originally considered by series creator David Jaffe after Kratos eliminated the Greek gods.[26] At the 2016 Electronic Entertainment Expo (E3), the next installment was officially announced with a gameplay demo and confirmed the concept art to be true. The demo showed a fully bearded Kratos with a son, and Kratos was teaching the boy how to hunt. The pair also battled a troll. The end of the demo showed the title God of War and confirmed it was in development for the PlayStation 4.[7][1] E3 also confirmed that Barlog had returned to the series as game director for the new installment. Barlog has been a major contributor in the development of the God of War series since the original installment in 2005, with his prior most notable role being game director of God of War II (2007). This new installment was his fifth God of War game.[20]

text: In naming the game, Barlog stated that it was deliberately titled God of War with no numeral or subtitle because although it is a continuation of the series, "we are reimagining everything."[22][19] Head of Santa Monica Studio Shannon Studstill and Barlog said that Sony Interactive Entertainment had to be convinced to do another God of War game as a lot of people at Sony wanted the series to "sleep and rest" due to the lackluster response of the previous game, Ascension.[27] In explaining why Barlog was brought back, Studstill said that he knew the series very well, "and bringing in someone that understands that history is the respect the franchise deserves."[10] Barlog followed up and said that "You gotta know the rules to break the rules."[10] Series creator David Jaffe was also considered, but was unavailable.[28]

text: In explaining the transition from Greek mythology to Norse mythology, Barlog said: "it's kind of this BC–AD change over kind of thing. We're moving and starting from zero and kind of moving forward on that."[22][19] Before settling on Norse mythology, Egyptian mythology was also considered. Barlog said that half of the team was for it, but since "there's a lot more about civilization - it's less isolated, less barren", he had to make the decision and decided on the Norse setting because they wanted the focus to stay on Kratos: "Having too much around distracts from that central theme of a stranger in a strange land."[23] In explaining why Kratos was now in the Norse world, Barlog said that different cultures' belief systems coexisted, but they were "separated by geography", suggesting that Kratos traveled from Greece to Norway (Scandinavia) after the conclusion of God of War III;[29] in clarifying the conclusion of that game, Barlog said that Kratos did not destroy what was believed to be the entire world, but only the portion that was ruled over by the Greeks.[9] Barlog said that the new game predates the Vikings; it is the time in which their gods walked the Earth.[20][23] It was also confirmed that this would not be Kratos' last game.[4] Barlog said that future games could see the series tackling Egyptian or Mayan mythology,[27] and that although this game focuses on Norse mythology, it alludes to the fact that there are other mythologies co-existing in the world.[30] Barlog also said that he liked the idea of having different directors for each game, seen throughout the first seven games, and although he may not direct another God of War, he would still be at Santa Monica to work on future games.[9]

text: Most of the development team that worked on the original God of War worked on the new installment.[22] They claimed that they matched the new gameplay with the same level of accessibility as the previous installments.[6] It was confirmed that the game would not feature any morality system or branching story; all players have the same story experience. The developers also confirmed that some of the more controversial mini-games found in previous entries (such as the sex mini-game) would not return.[22] Some gameplay characteristics found in the previous installments were cut, such as jumping, swimming, and instant-death platforming challenges; these were cut due to the camera being closer to Kratos.[10] Although the previous installment, Ascension, introduced multiplayer to the series, the team decided to drop the mode to focus on the single-player experience.[5] In changing the gameplay, Studstill said "I felt like, in order to reinvent, we really needed to turn a lot of things around."[31] In regards to the camera change, Barlog said "We wanted a much more intimate experience, a much more up close, and a much more player-controlled experience, so the camera really is a mechanic that [we leaned] into heavily for everything in the game."[6] Explaining Kratos' axe, lead gameplay designer Jason McDonald, who has worked on the series since the original game, said the axe was chosen because they wanted a more grounded direction for the game. Initially, they were unsure how to make it unique, like how the double-chained blades were. After they came up with the concept of throwing the axe and having it return to Kratos, "things started to fall into place."[32] McDonald said that combat with the axe is a little slower than what it was with the blades, "but it's just as fluid and just as brutal as it's ever been."[32] Barlog took inspiration from Dark Souls (2011), which influenced the game's combat system, particularly its gameplay loop and strategic decision-making,[33] as well as the game's approach to storytelling.[34] In addition, designers Anthony Dimento and Luis Sanchez revealed how God of War's level design and exploration was influenced by Bloodborne (2015), as they wanted to "just have the world breathe a little bit" and expand upon player discovery by including "micro-loops where you're unlocking paths, unlocking shortcuts..." that gave purpose.[35]

text: The entire game was done in a single shot, as in no camera cuts, meaning there are no loading screens or fade-to-black between gameplay and cutscenes.[17] The frame rate was confirmed as 30 frames per second,[4] and the enemy count was increased to up to 100 enemies on-screen; God of War III and Ascension could do up to 50.[22] Unlike the previous games, Santa Monica did not make a demo specifically for public release. Barlog explained that doing so would have delayed the game by a couple of months.[36] He also confirmed that the game was built for the standard PlayStation 4,[36] but the game would "benefit from the power" of the PlayStation 4 Pro; an updated version of the PlayStation 4 that can render games in 4K and was released a few months after God of War was announced.[37] In late December 2016, Barlog confirmed that the game was playable from start to finish.[38] At the 2017 PlayStation Experience, Barlog said that the game's story would take 25–35 hours to complete, which is significantly more than the previous four main installments, which each took an average of 10 hours to complete.[39]

text: At E3 2017, a new trailer was shown featuring new gameplay, cinematics, and characters. In the trailer, Kratos was shown using a shield that he could use offensively and defensively. At one point in the trailer, Kratos finds a Greek vase with himself on it, wielding his infamous double-chained blades. During the trailer, an unnamed woman warns Kratos about the Norse gods, as they know what he did to the Greek gods, while a pair of wolves were also shown. The trailer ends with Kratos and Atreus encountering the World Serpent. Atreus was able to translate what it said, which was that it wanted to help the pair. It was confirmed that the game would release in early 2018.[15] Since E3 2017 until the game's launch, Santa Monica included a section on the God of War website titled "The Lost Pages", detailing some of the lore of God of War's Nordic world.[13] In January 2018, the game's release date was confirmed for April 20, 2018. A trailer was also released that showed that the character Mímir from the mythology would have a role in the game.[40] On March 22, God of War went gold.[41]

text: Characterization[edit]

text: During early development, there was talk about having a different protagonist for the game, but it was decided to keep Kratos. Referencing the Nintendo character Mario and the Mario games, Barlog said that just like Mario, "Kratos is intrinsically tied" to the God of War series.[20] In regards to the new changes, Barlog said that:

text: I knew I didn’t want to simply reboot the franchise, starting over with a retelling of the origin story. I wanted to reimagine the gameplay, give players a fresh perspective and a new tactile experience while delving deeper into the emotional journey of Kratos to explore the compelling drama that unfolds when an immortal demigod makes a decision to change.[42]

text: Barlog explained that Kratos had to change his cycle of violence and learn how to control his rage. He said that Kratos had made many bad decisions, which led to the destruction of Olympus, and wanted to know what would happen if Kratos made a good decision. The birth of Barlog's own son influenced the idea of Kratos' character change.[42] The canceled live-action Star Wars television series was also an influence.[43] The bond between Kratos and his son is at the heart of the game and Barlog said "This game is about Kratos teaching his son how to be a god, and his son teaching Kratos how to be human again."[17] Referencing the Marvel Comics character Hulk, Barlog said that in regards to Kratos, "We've already told the story of The Hulk. We want to tell the story of [Bruce] Banner now."[17] One of their goals was to make Kratos "a more nuanced and interesting character."[44] In changing the narrative focus, Studstill said "I think we inherently knew the franchise needed to evolve in that emotional beat and be something meatier for the older generation of gamers."[44]

text: Christopher Judge, best known as Teal'c from Stargate SG-1, replaced T.C. Carson as the voice of Kratos; Carson had voiced Kratos since the original God of War. Commenting in response to the change, Carson said, "Sony went in a new direction."[45] Barlog explained that the way the previous games were made, they were able to have someone else do the motion capture instead of the voice actor. Although Carson had done the motion capture for Kratos in Ascension, Barlog said the actor change was made because of the type of camera work they wanted to do. For the new camera work, they needed someone who was closer to Kratos' size to do the motion capture along with a child. Carson was unsuitable for this because he was much shorter than Kratos, who is over 6-feet tall: "Offsetting [Carson's height] for the size of a child, it turned out it was going to be almost impossible to try and actually shoot them and go in and redo the animations."[46] Judge was chosen because he was 6-foot-3 and had the body of a professional football player. He was also chosen because of the chemistry with his then-10-year-old co-star, Sunny Suljic, who plays Kratos' son Atreus; Suljic's opinion was also sought in making the decision, and out of all the auditions, he liked Judge the most. The two bonded well, and Judge described his time with Suljic as time he had missed with his own children. In stepping into the role of Kratos, Judge took it as an opportunity to add something new to the character. He researched the character and Carson's performance, but decided not to imitate it. Since Santa Monica were going in a new direction, he decided to start fresh and go from there. Judge was thrown off when he first read the script, stating it "was a real script", and not just "a way to get into battles."[46] He said "it was really this great story of this relationship and this crazy mythology."[46] While Judge did all of Kratos' motion capture for the cinematic scenes, stuntman Eric Jacobus did all of Kratos' combat motion capture; Jacobus was found by God of War's animators on YouTube. Instead of going directly to Santa Monica to audition, Jacobus recorded an audition tape and they immediately hired him.[47]

text: During E3 2016, GameSpot mistakenly reported that Kratos' son's name was Charlie, which Barlog laughingly denied.[48] In January 2017, after a fan downloaded the God of War overture and saw the track's details that said "An introduction to Kratos and Atreus", Barlog confirmed on Twitter that Atreus was in fact the son's name.[49] Barlog said that Atreus was unaware that Kratos was a demigod, and did not know about his past.[4] They did not reveal details of Atreus' mother prior to release due to her being a critical part of the story.[22] Barlog stated that during gameplay, Atreus would be "like magic, an additional combat resource, and [the player is] training him and teaching him."[20] The developers stated that Atreus would not be a burden during gameplay.[13] The team experimented with several different approaches for Atreus to ensure that he was an empowering presence. Barlog said he did not want the game to be an escort-mission where the A.I. caused a problem for the player. Their goal was for Atreus to enhance Kratos' capabilities without Atreus becoming a liability. This resulted in the developers designating a command button for Atreus as well as for him to act freely.[18] During combat, Atreus was also designed to call out enemy locations, as due to the camera being closer to Kratos, some enemies may be difficult for the player to see. Jason McDonald said it took a lot of iterations with the enemies and Atreus to make it all work together.[32]

text: Early in development, it was suggested for Atreus to be cut or to significantly minimize his role due to the many developmental challenges and financial expense. Barlog stated that the game could have worked without Atreus, but it would have been completely different, likening it to that of the 2013 film All Is Lost. Barlog said that with just Kratos, it would have been "one character who talks to himself occasionally, but generally, it will be very silent and everyone will talk in old Norse, so that you won't understand anything anybody's saying."[50] After hearing Barlog's case, Sony gave him the freedom to incorporate Atreus. Lead level designer Rob Davis also noted that with Atreus, it allowed for "significant gameplay and storytelling opportunities that might not otherwise [been] possible."[50]

text: Soundtrack[edit]

text: Bear McCreary, best known for his work on television shows such as Battlestar Galactica and The Walking Dead, scored God of War and composed themes for the game.[51] McCreary said that he was called into Santa Monica Studio in November 2014 to meet with music producers Peter Scaturro and Keith Leary to discuss "a secret project"; McCreary had previously collaborated with Scaturro and Leary on 2011's SOCOM 4: U.S. Navy SEALs. Ideas of folk music, mythology, Nordic ethnic instruments, vocal writing, and classical thematic development were discussed, to which McCreary correctly guessed that the discussions were about a new God of War. He met with Barlog early on, and they discussed Barlog's narrative vision for the game. After meeting with Barlog, he felt that the franchise was in good hands because God of War II, which Barlog also directed, was his favorite installment.[52]

text: McCreary's initial efforts focused on writing the new main theme, or what he called the Kratos Theme. He spent several months working with Barlog, Scaturro, Leary, Sony music director Chuck Doud, and the rest of the development team in making this new theme. McCreary described the Kratos Theme as "arguably one of my most structurally satisfying and catchy melodies." The main theme features low orchestral instruments, Icelandic choir, deep male vocals, powerful female vocals (in particular Faroese singer Eivør Pálsdóttir), folk percussion, and Nordic stringed instruments, such as the nyckelharpa and hurdy gurdy. When it was decided that God of War would be revealed at E3 2016, Sony wanted McCreary to perform his original score with a live orchestra at the press conference. McCreary opened the show with the new main theme before the unveiling of God of War, and performed the gameplay demo's music live during the presentation.[52] On January 13, 2017, a live recording from E3 2016 of God of War's overture was released for free for a limited time. Barlog released the overture as a thank you to fans for God of War's E3 2016 trailer reaching fifteen million views on YouTube.[53]

text: Release[edit]

text: The game was released worldwide on April 20, 2018, for the PlayStation 4.[40] In addition to the standard base game, there were three special editions: the Stone Mason Edition, the Collector's Edition, and the Digital Deluxe Edition. Only available in the United States and Canada, the Stone Mason Edition came with several physical items, including the base game in a SteelBook case, a 9-inch (230 mm) statue of Kratos and Atreus that was created by Gentle Giant, 2-inch (51 mm) carvings of the Huldra Brothers, a horse, and a troll, an exclusive lithograph, a cloth map, a stone mason's ring, and a keychain of Mímir's head that talks. There was also various downloadable content (DLC), including an exclusive shield skin, in addition to an armor set and another shield skin for Kratos, a PlayStation 4 dynamic theme, a digital artbook, and God of War #0 by Dark Horse Comics.[54] The Collector's Edition came with many of the same items, minus the ring, the keychain, the 2-inch (51 mm) carvings of the horse and troll, and the exclusive shield skin. The Digital Deluxe Edition comes with all of the digital content, minus the exclusive shield skin. U.S. and Canadian customers also received a Kratos and Atreus pin for pre-ordering the Digital Deluxe Edition. Pre-orders at select retailers received three skins for Kratos' shield, while pre-orders from GameStop or EB Games also received a "Luck of Ages XP Talisman", granting increased XP gain, increased Hacksilver gain, and increased ability to trigger perks.[55]

text: In addition to the special editions of the game, a Limited Edition PlayStation 4 Pro bundle was also available the same day as the game's release. The bundle included the standard base game, a PlayStation 4 Pro console decorated with the runes as on Kratos' axe, and a similarly themed DualShock 4 controller with the God of War logo.[56] Game Director Cory Barlog confirmed that God of War would not have microtransactions post-launch, a feature that has become prominent with other recent games and negatively criticized.[57] Since launch, Santa Monica has supported the game via patch updates to address software bugs. Additionally, the developers added a Photo Mode in update patch 1.20, released on May 9, 2018. Photo Mode allows players to take customized in-game screenshots. Players can adjust the field of view, depth of view, filters, borders, the visibility of characters, and the ability to change the facial expressions of Kratos and Atreus.[58] At E3 2018, it was confirmed that a New Game Plus mode would be added in a future update. In addition to players having access to their fully upgraded gear upon the start of a new game, the difficulty will be increased.[59]

text: God of War: A Call from the Wilds[edit]

text: God of War: A Call from the Wilds is a text-based game playable through Facebook Messenger. To help further promote God of War, Sony partnered with Facebook to develop the play-by-web game, which released on February 1, 2018. Completing the game unlocks downloadable concept art. The short story follows Atreus on his first adventure in the Norse wilds. After archery training and learning runes with his mother, Atreus adventures into the wilderness after telepathically hearing the voice of a dying deer; he finds it covered in blood and stays with it during its final moments. A couple of draugrs appear and Atreus attempts to fight them, but is injured. He is saved by his father, Kratos, who was out hunting. The two then battle a revenant before returning home.[60][61][62]

text: Reception[edit]

text: This section needs expansion. You can help by adding to it. (June 2018)

tag: table

text:
Reception

Aggregate score

Aggregator
Score

Metacritic
94/100[63]

Review scores

Publication
Score

Destructoid
10/10[64]

EGM
9.5/10[65]

Game Informer
9.75/10[66]

Game Revolution
[67]

GameSpot
9/10[68]

GamesRadar+
[69]

Giant Bomb
[70]

IGN
10/10[71]

Polygon
10/10[72]

The Guardian
[73]

USgamer
[74]

text: God of War received "universal acclaim" according to review aggregator Metacritic,[63] tying it with the original God of War for the highest score in the franchise. It has the third highest score of all-time for a PlayStation 4 game, and the highest score for an original, non-remastered PlayStation 4 game.[75] It is the highest rated PlayStation 4 game of 2018,[76] and is tied with the Xbox One version of Celeste for the highest score of 2018, regardless of platform.[77] God of War received particular praise for its art direction, graphics, combat system, music, story, use of Norse mythology, characters, and cinematic feeling. Critics called the game a "technical achievement" and one of the most impressive-looking games developed for consoles. Many also felt that it had successfully revitalized the series without losing the core identity of its predecessors.[78]

text: Sales[edit]

text: During its release week in the UK, God of War became the fastest-selling entry in the franchise, selling 35% more physical copies than God of War III.[79] The game remained at the top of the all format sales chart throughout April and May for six consecutive weeks, setting a record for a PlayStation 4 exclusive having the most consecutive weeks at number one.[80] It sold 46,091 copies within its first week on sale in Japan, which placed it at number two on the sales chart.[81]

text: The game sold over 3 million copies in three days after its release, making it the fastest-selling PlayStation 4 exclusive.[82] The game was the fastest selling game of the month of its release and contributed to the PS4 being the best selling console of that month.[83] In total, the game sold over five million copies in its first month, with 2.1 million in digital sales.[84][85]

text: Accolades[edit]

text:
Year
Award
Category
Result
Ref

2016
Game Critics Awards 2016
Special Commendation for Graphics
Won
[86]

IGN's Best of E3 2016 Awards
Game of the Show
Nominated
[87]

Best PlayStation 4 Game
Nominated

Best Action Game
Nominated

The Game Awards 2016
Most Anticipated Game
Nominated
[88]

2017
The Game Awards 2017
Most Anticipated Game
Nominated
[89]

text: Notes[edit]

text:
Jump up ^ Colloquially referred to as God of War 4[1][2][3] and God of War PS4[4][5]
Jump up ^ As depicted in God of War III (2010)