Geastrum minimum Schwein., syn.: Geastrum marginatum Vittad.

Family: Geastraceae Corda

EN: Tiny earthstar, DE: Kleiner Erdstern, Zwergerdstern

Slo.: drobcena zvezdica

 

Dat.: Aug. 31. 2021

Lat.: 46.37725 Long.: 13.743703

Code: Bot_1398/2021_DSC08259

 

Habitat: At the base of a mountain ravine, slightly inclined mountain slope; east aspect; about 30 m distance to river Soča and about 10 m higher; close to a Pinus sp. forest; fine, older colluvial, calcareous deposits partly overgrown (Erica carnea dominant); dry, open, sunny, relatively warm place; exposed to direct rain; average precipitations ~ 3.000 mm/year, average temperature 7-9 deg C, elevation 595 m (1.950 feet), alpine phytogeographical region.

 

Substratum: soil, fine sandy colluvial deposits among larger rocks.

 

Place: Lower Trenta valley, south of village Trenta, right bank of river Soča, East Julian Alps, Posočje, Slovenia EC

 

Comments: This find is the smallest earthstar I've ever found. Based on the number of non-hygroscopic exoperidial rays, persistent mycelial layer, encrusted with debris, very small endoperidial body, which is stalked and provided with an apophysis, whitish crystals on endoperidial body surface and a clearly delimited, rather flat, fibrillose, not furrowed peristome I am pretty sure this is Geastrum minimum. The fungus is a rather rare find in Slovenia.

 

Growing in a group of three sporocarps. Endoperidium diameter; 6, 7 and 9 mm; SP on mass black. Spores verrucose, dark. Dimensions: (3.8) 4.4 - 5.5 (5.6) × (3.6) 4.2 - 5.2 (5.7) µm; Q = (0.9) 1 - 1.1 (1.2); N = 52; Me = 4.9 × 4.7 µm; Qe = 1. Olympus CH20, NEA 100x/1.25, magnification 1.000 x, oil (spores); NEA 40x/0.65, magnification 400x (capillitium), in water, fresh material. AmScope MA500 digital camera.

 

Ref.:

(1) www.funghiitaliani.it/topic/88099-geastrum-minimum/ (last accessed Oct.17. 2021)

(2) T. Lӕssøe, J.H. Petersen, Fungi of temperate Europe, Vol. 2., Princeton University Press (2019), p 1249.

(3) G.J. Krieglsteiner (Hrsg.), Die Grosspilze Baden-Württembergs, Band 2., Ulmer (2000), p 112.

(4) www.123pilze.de/DreamHC/Download/KleinerErdstern.htm (last accessed Oct.17. 2021),

(5) J.C. Zamora1, F.D. Calonge, M.P. Martín, Integrative taxonomy reveals an unexpected diversity in Geastrum section Geastrum (Geastrales, Basidiomycota), Persoonia 34, 2015: 130–165; geastrum_sect_geastrum_1422974412556.pdf,

(6) M. Jeppson, R. H. Nilsson, E. Larsson, 2013, European earthstars in Geastraceae (Geastrales, Phallomycetidae) – a systematic approach using morphology and molecular sequence data; Systematics and Biodiversity, 11:4, pp 437-465, Natural History Museum, Taylor & Francis Online.

(5) S. Buczacki, Collins Fungi Guide, Collins (2012), p 440.

(6) D. Arora, Mushrooms Demystified, Ten Speed Press, Berkeley (1986), p 702.

(7) W. Rothmaler, Exkursionsflora von Deutschland, Vol.1, Niedere Pflanzen, Elsevier, 3. Auflage, (1994), p 519.

  

Phallus impudicus.

 

synonyms: Gemeine Stinkmorchel, Oeuf du diable, Phallus Impudique, Satyre puant, Stinkhorn

 

location: North America, Europe

edibility: Inedible

fungus colour: White to cream, Black or blackish

normal size: 5-15cm

cap type: Other

stem type: Volva on stem, Stem much longer than cap diameter

flesh: Mushroom has distinct or odd smell (non mushroomy), Mushroom slimy or sticky

spore colour: Light to dark brown

habitat: Grows in woods, Grows on the ground

 

Phallus impudicus Pers. syn. Ithyphallus impudicus (L.) Fr. Gemeine Stinkmorchel Phallus Impudique, Satyre puant, Oeuf du diable, Stinkhorn. Fruit body initially semi-submerged and covered by leaf-litter, egg-like, 3–6cm across, attached to substrate by a cord-like mycelial strand. The outer wall of the egg is white to pinkish but there is a thick gelatinous middle layer held between the membranous inner and outer layers. The egg is soon ruptured, as the white hollow stalk-like receptacle extends to 10–25cm high, the pendulous, bell-shaped head is covered by a meshwork of raised ribs covered in dark olive slime which contains the spores. This slime has a strong sickly offensive smell which attracts flies from large distances, the slime sticks to the legs of the flies and thus acts as a means of spore dispersal which takes place very rapidly, exposing the underlying mesh of the cap. Spores pale yellow, oblong, 3.5–4 x 1.5–2µ. Habitat associated with rotting wood which may be buried in the soil, in gardens and woodland. Season summer to late autumn. Very common. The egg stage, which lacks the disgusting smell, is edible though not tasty; it is said to be an aphrodisiac presumably through association with its phallic shape. Distribution, America and Europe.

 

info by Roger Phillips:

 

www.rogersmushrooms.com

 

location: North America, Europe

edibility: Inedible

fungus colour: White to cream, Black or blackish

normal size: 5-15cm

cap type: Other

stem type: Volva on stem, Stem much longer than cap diameter

flesh: Mushroom has distinct or odd smell (non mushroomy), Mushroom slimy or sticky

spore colour: Light to dark brown

habitat: Grows in woods, Grows on the ground

 

Phallus impudicus Pers. syn. Ithyphallus impudicus (L.) Fr. Gemeine Stinkmorchel Phallus Impudique, Satyre puant, Oeuf du diable, Stinkhorn. Fruit body initially semi-submerged and covered by leaf-litter, egg-like, 3–6cm across, attached to substrate by a cord-like mycelial strand. The outer wall of the egg is white to pinkish but there is a thick gelatinous middle layer held between the membranous inner and outer layers. The egg is soon ruptured, as the white hollow stalk-like receptacle extends to 10–25cm high, the pendulous, bell-shaped head is covered by a meshwork of raised ribs covered in dark olive slime which contains the spores. This slime has a strong sickly offensive smell which attracts flies from large distances, the slime sticks to the legs of the flies and thus acts as a means of spore dispersal which takes place very rapidly, exposing the underlying mesh of the cap. Spores pale yellow, oblong, 3.5–4 x 1.5–2µ. Habitat associated with rotting wood which may be buried in the soil, in gardens and woodland. Season summer to late autumn. Very common. The egg stage, which lacks the disgusting smell, is edible though not tasty; it is said to be an aphrodisiac presumably through association with its phallic shape. Distribution, America and Europe.

 

info by Roger Phillips:

 

www.rogersmushrooms.com

A fungus (pl.: fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as one of the traditional eukaryotic kingdoms, along with Animalia, Plantae and either Protista or Protozoa and Chromista.

 

A characteristic that places fungi in a different kingdom from plants, bacteria, and some protists is chitin in their cell walls. Fungi, like animals, are heterotrophs; they acquire their food by absorbing dissolved molecules, typically by secreting digestive enzymes into their environment. Fungi do not photosynthesize. Growth is their means of mobility, except for spores (a few of which are flagellated), which may travel through the air or water. Fungi are the principal decomposers in ecological systems. These and other differences place fungi in a single group of related organisms, named the Eumycota (true fungi or Eumycetes), that share a common ancestor (i.e. they form a monophyletic group), an interpretation that is also strongly supported by molecular phylogenetics. This fungal group is distinct from the structurally similar myxomycetes (slime molds) and oomycetes (water molds). The discipline of biology devoted to the study of fungi is known as mycology (from the Greek μύκης mykes, mushroom). In the past mycology was regarded as a branch of botany, although it is now known that fungi are genetically more closely related to animals than to plants.

 

Abundant worldwide, most fungi are inconspicuous because of the small size of their structures, and their cryptic lifestyles in soil or on dead matter. Fungi include symbionts of plants, animals, or other fungi and also parasites. They may become noticeable when fruiting, either as mushrooms or as molds. Fungi perform an essential role in the decomposition of organic matter and have fundamental roles in nutrient cycling and exchange in the environment. They have long been used as a direct source of human food, in the form of mushrooms and truffles; as a leavening agent for bread; and in the fermentation of various food products, such as wine, beer, and soy sauce. Since the 1940s, fungi have been used for the production of antibiotics, and, more recently, various enzymes produced by fungi are used industrially and in detergents. Fungi are also used as biological pesticides to control weeds, plant diseases, and insect pests. Many species produce bioactive compounds called mycotoxins, such as alkaloids and polyketides, that are toxic to animals, including humans. The fruiting structures of a few species contain psychotropic compounds and are consumed recreationally or in traditional spiritual ceremonies. Fungi can break down manufactured materials and buildings, and become significant pathogens of humans and other animals. Losses of crops due to fungal diseases (e.g., rice blast disease) or food spoilage can have a large impact on human food supplies and local economies.

 

The fungus kingdom encompasses an enormous diversity of taxa with varied ecologies, life cycle strategies, and morphologies ranging from unicellular aquatic chytrids to large mushrooms. However, little is known of the true biodiversity of the fungus kingdom, which has been estimated at 2.2 million to 3.8 million species. Of these, only about 148,000 have been described, with over 8,000 species known to be detrimental to plants and at least 300 that can be pathogenic to humans. Ever since the pioneering 18th and 19th century taxonomical works of Carl Linnaeus, Christiaan Hendrik Persoon, and Elias Magnus Fries, fungi have been classified according to their morphology (e.g., characteristics such as spore color or microscopic features) or physiology. Advances in molecular genetics have opened the way for DNA analysis to be incorporated into taxonomy, which has sometimes challenged the historical groupings based on morphology and other traits. Phylogenetic studies published in the first decade of the 21st century have helped reshape the classification within the fungi kingdom, which is divided into one subkingdom, seven phyla, and ten subphyla.

 

Etymology

The English word fungus is directly adopted from the Latin fungus (mushroom), used in the writings of Horace and Pliny. This in turn is derived from the Greek word sphongos (σφόγγος 'sponge'), which refers to the macroscopic structures and morphology of mushrooms and molds; the root is also used in other languages, such as the German Schwamm ('sponge') and Schimmel ('mold').

 

The word mycology is derived from the Greek mykes (μύκης 'mushroom') and logos (λόγος 'discourse'). It denotes the scientific study of fungi. The Latin adjectival form of "mycology" (mycologicæ) appeared as early as 1796 in a book on the subject by Christiaan Hendrik Persoon. The word appeared in English as early as 1824 in a book by Robert Kaye Greville. In 1836 the English naturalist Miles Joseph Berkeley's publication The English Flora of Sir James Edward Smith, Vol. 5. also refers to mycology as the study of fungi.

 

A group of all the fungi present in a particular region is known as mycobiota (plural noun, no singular). The term mycota is often used for this purpose, but many authors use it as a synonym of Fungi. The word funga has been proposed as a less ambiguous term morphologically similar to fauna and flora. The Species Survival Commission (SSC) of the International Union for Conservation of Nature (IUCN) in August 2021 asked that the phrase fauna and flora be replaced by fauna, flora, and funga.

 

Characteristics

 

Fungal hyphae cells

Hyphal wall

Septum

Mitochondrion

Vacuole

Ergosterol crystal

Ribosome

Nucleus

Endoplasmic reticulum

Lipid body

Plasma membrane

Spitzenkörper

Golgi apparatus

 

Fungal cell cycle showing Dikaryons typical of Higher Fungi

Before the introduction of molecular methods for phylogenetic analysis, taxonomists considered fungi to be members of the plant kingdom because of similarities in lifestyle: both fungi and plants are mainly immobile, and have similarities in general morphology and growth habitat. Although inaccurate, the common misconception that fungi are plants persists among the general public due to their historical classification, as well as several similarities. Like plants, fungi often grow in soil and, in the case of mushrooms, form conspicuous fruit bodies, which sometimes resemble plants such as mosses. The fungi are now considered a separate kingdom, distinct from both plants and animals, from which they appear to have diverged around one billion years ago (around the start of the Neoproterozoic Era). Some morphological, biochemical, and genetic features are shared with other organisms, while others are unique to the fungi, clearly separating them from the other kingdoms:

 

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. 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).

With animals: Fungi lack chloroplasts and are heterotrophic organisms and so require preformed organic compounds as energy sources.

With plants: Fungi have a cell wall and vacuoles. They reproduce by both sexual and asexual means, and like basal plant groups (such as ferns and mosses) produce spores. Similar to mosses and algae, fungi typically have haploid nuclei.

With euglenoids and bacteria: Higher fungi, euglenoids, and some bacteria produce the amino acid L-lysine in specific biosynthesis steps, called the α-aminoadipate pathway.

The cells of most fungi grow as tubular, elongated, and thread-like (filamentous) structures called hyphae, which may contain multiple nuclei and extend by growing at their tips. Each tip contains a set of aggregated vesicles—cellular structures consisting of proteins, lipids, and other organic molecules—called the Spitzenkörper. Both fungi and oomycetes grow as filamentous hyphal cells. In contrast, similar-looking organisms, such as filamentous green algae, grow by repeated cell division within a chain of cells. There are also single-celled fungi (yeasts) that do not form hyphae, and some fungi have both hyphal and yeast forms.

In common with some plant and animal species, more than one hundred fungal species display bioluminescence.

Unique features:

 

Some species grow as unicellular yeasts that reproduce by budding or fission. Dimorphic fungi can switch between a yeast phase and a hyphal phase in response to environmental conditions.

The fungal cell wall is made of a chitin-glucan complex; while glucans are also found in plants and chitin in the exoskeleton of arthropods, fungi are the only organisms that combine these two structural molecules in their cell wall. Unlike those of plants and oomycetes, fungal cell walls do not contain cellulose.

A whitish fan or funnel-shaped mushroom growing at the base of a tree.

Omphalotus nidiformis, a bioluminescent mushroom

Most fungi lack an efficient system for the long-distance transport of water and nutrients, such as the xylem and phloem in many plants. To overcome this limitation, some fungi, such as Armillaria, form rhizomorphs, which resemble and perform functions similar to the roots of plants. As eukaryotes, fungi possess a biosynthetic pathway for producing terpenes that uses mevalonic acid and pyrophosphate as chemical building blocks. Plants and some other organisms have an additional terpene biosynthesis pathway in their chloroplasts, a structure that fungi and animals do not have. Fungi produce several secondary metabolites that are similar or identical in structure to those made by plants. Many of 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.

 

Diversity

Fungi have a worldwide distribution, and grow in a wide range of habitats, including extreme environments such as deserts or areas with high salt concentrations or ionizing radiation, as well as in deep sea sediments. Some can survive the intense UV and cosmic radiation encountered during space travel. Most grow in terrestrial environments, though several species live partly or solely in aquatic habitats, such as the chytrid fungi Batrachochytrium dendrobatidis and B. salamandrivorans, parasites that have been responsible for a worldwide decline in amphibian populations. These organisms spend part of their life cycle as a motile zoospore, enabling them to propel itself through water and enter their amphibian host. Other examples of aquatic fungi include those living in hydrothermal areas of the ocean.

 

As of 2020, around 148,000 species of fungi have been described by taxonomists, but the global biodiversity of the fungus kingdom is not fully understood. A 2017 estimate suggests there may be between 2.2 and 3.8 million species The number of new fungi species discovered yearly has increased from 1,000 to 1,500 per year about 10 years ago, to about 2000 with a peak of more than 2,500 species in 2016. In the year 2019, 1882 new species of fungi were described, and it was estimated that more than 90% of fungi remain unknown The following year, 2905 new species were described—the highest annual record of new fungus names. In mycology, species have historically been distinguished by a variety of methods and concepts. Classification based on morphological characteristics, such as the size and shape of spores or fruiting structures, has traditionally dominated fungal taxonomy. 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.

 

Mycology

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.

 

The use of fungi by humans dates back to prehistory; Ötzi the Iceman, a well-preserved mummy of a 5,300-year-old Neolithic man found frozen in the Austrian Alps, carried two species of polypore mushrooms that may have been used as tinder (Fomes fomentarius), or for medicinal purposes (Piptoporus betulinus). Ancient peoples have used fungi as food sources—often unknowingly—for millennia, in the preparation of leavened bread and fermented juices. Some of the oldest written records contain references to the destruction of crops that were probably caused by pathogenic fungi.

 

History

Mycology became a systematic science after the development of the microscope 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. Micheli not only observed spores but also showed that, under the proper conditions, they could be induced into growing into the same species of fungi from which they originated. Extending the use of the binomial system of nomenclature introduced by Carl Linnaeus in his Species plantarum (1753), the Dutch Christiaan Hendrik Persoon (1761–1836) established the first classification of mushrooms with such skill as to be considered a founder of modern mycology. Later, Elias Magnus Fries (1794–1878) further elaborated the classification of fungi, using spore color and microscopic characteristics, methods still used by taxonomists today. Other notable early contributors to mycology in the 17th–19th and early 20th centuries include Miles Joseph Berkeley, August Carl Joseph Corda, Anton de Bary, the brothers Louis René and Charles Tulasne, Arthur H. R. Buller, Curtis G. Lloyd, and Pier Andrea Saccardo. In the 20th and 21st centuries, advances in biochemistry, genetics, molecular biology, biotechnology, DNA sequencing and phylogenetic analysis has provided new insights into fungal relationships and biodiversity, and has challenged traditional morphology-based groupings in fungal taxonomy.

 

Morphology

Microscopic structures

Monochrome micrograph showing Penicillium hyphae as long, transparent, tube-like structures a few micrometres across. Conidiophores branch out laterally from the hyphae, terminating in bundles of phialides on which spherical condidiophores are arranged like beads on a string. Septa are faintly visible as dark lines crossing the hyphae.

An environmental isolate of Penicillium

Hypha

Conidiophore

Phialide

Conidia

Septa

Most fungi grow as hyphae, which are cylindrical, thread-like structures 2–10 µm in diameter and up to several centimeters in length. Hyphae grow at their tips (apices); new hyphae are typically formed by emergence of new tips along existing hyphae by a process called branching, or occasionally growing hyphal tips fork, giving rise to two parallel-growing hyphae. Hyphae also sometimes fuse when they come into contact, a process called hyphal fusion (or anastomosis). These growth processes lead to the development of a mycelium, an interconnected network of hyphae. Hyphae can be either septate or coenocytic. Septate hyphae are divided into compartments separated by cross walls (internal cell walls, called septa, that are formed at right angles to the cell wall giving the hypha its shape), with each compartment containing one or more nuclei; coenocytic hyphae are not compartmentalized. Septa have pores that allow cytoplasm, organelles, and sometimes nuclei to pass through; an example is the dolipore septum in fungi of the phylum Basidiomycota. Coenocytic hyphae are in essence multinucleate supercells.

 

Many species have developed specialized hyphal structures for nutrient uptake from living hosts; examples include haustoria in plant-parasitic species of most fungal phyla,[63] and arbuscules of several mycorrhizal fungi, which penetrate into the host cells to consume nutrients.

 

Although fungi are opisthokonts—a grouping of evolutionarily related organisms broadly characterized by a single posterior flagellum—all phyla except for the chytrids have lost their posterior flagella. Fungi are unusual among the eukaryotes in having a cell wall that, in addition to glucans (e.g., β-1,3-glucan) and other typical components, also contains the biopolymer chitin.

 

Macroscopic structures

Fungal mycelia can become visible to the naked eye, for example, on various surfaces and substrates, such as damp walls and spoiled food, where they are commonly called 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. Some individual fungal colonies can reach extraordinary dimensions and ages as in the case of a clonal colony of Armillaria solidipes, which extends over an area of more than 900 ha (3.5 square miles), with an estimated age of nearly 9,000 years.

 

The apothecium—a specialized structure important in sexual reproduction in the ascomycetes—is a cup-shaped fruit body that is often macroscopic and holds the hymenium, a layer of tissue containing the spore-bearing cells. The fruit bodies of the basidiomycetes (basidiocarps) and some ascomycetes can sometimes grow very large, and many are well known as mushrooms.

 

Growth and physiology

Time-lapse photography sequence of a peach becoming progressively discolored and disfigured

Mold growth covering a decaying peach. The frames were taken approximately 12 hours apart over a period of six days.

The growth of fungi as hyphae on or in solid substrates or as single cells in aquatic environments is adapted for the efficient extraction of nutrients, because these growth forms have high surface area to volume ratios. Hyphae are specifically adapted for growth on solid surfaces, and to invade substrates and tissues. They can exert large penetrative mechanical forces; for example, many plant pathogens, including Magnaporthe grisea, form a structure called an appressorium that evolved to puncture plant tissues.[71] The pressure generated by the appressorium, directed against the plant epidermis, can exceed 8 megapascals (1,200 psi).[71] The filamentous fungus Paecilomyces lilacinus uses a similar structure to penetrate the eggs of nematodes.

 

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

 

Fungi are traditionally considered heterotrophs, organisms that rely solely on carbon fixed by other organisms for metabolism. Fungi have evolved a high degree of metabolic versatility that allows them to use a diverse range of organic substrates for growth, including simple compounds such as nitrate, ammonia, acetate, or ethanol. In some species the pigment melanin may play a role in extracting energy from ionizing radiation, such as gamma radiation. This form of "radiotrophic" growth has been described for only a few species, the effects on growth rates are small, and the underlying biophysical and biochemical processes are not well known. This process might bear similarity to CO2 fixation via visible light, but instead uses ionizing radiation as a source of energy.

 

Reproduction

Two thickly stemmed brownish mushrooms with scales on the upper surface, growing out of a tree trunk

Polyporus squamosus

Fungal reproduction is complex, reflecting the differences in lifestyles and genetic makeup within this diverse kingdom of organisms. It is estimated that a third of all fungi reproduce using more than one method of propagation; for example, reproduction may occur in two well-differentiated stages within the life cycle of a species, the teleomorph (sexual reproduction) and the anamorph (asexual reproduction). Environmental conditions trigger genetically determined developmental states that lead to the creation of specialized structures for sexual or asexual reproduction. These structures aid reproduction by efficiently dispersing spores or spore-containing propagules.

 

Asexual reproduction

Asexual reproduction occurs via vegetative spores (conidia) or through mycelial fragmentation. Mycelial fragmentation occurs when a fungal mycelium separates into pieces, and each component grows into a separate mycelium. Mycelial fragmentation and vegetative spores maintain clonal populations adapted to a specific niche, and allow more rapid dispersal than sexual reproduction. The "Fungi imperfecti" (fungi lacking the perfect or sexual stage) or Deuteromycota comprise all the species that lack an observable sexual cycle. Deuteromycota (alternatively known as Deuteromycetes, conidial fungi, or mitosporic fungi) is not an accepted taxonomic clade and is now taken to mean simply fungi that lack a known sexual stage.

 

Sexual reproduction

See also: Mating in fungi and Sexual selection in fungi

Sexual reproduction with meiosis has been directly observed in all fungal phyla except Glomeromycota (genetic analysis suggests meiosis in Glomeromycota as well). It differs in many aspects from sexual reproduction in animals or plants. Differences also exist between fungal groups and can be used to discriminate species by morphological differences in sexual structures and reproductive strategies. Mating experiments between fungal isolates may identify species on the basis of biological species concepts. The major fungal groupings have initially been delineated based on the morphology of their sexual structures and spores; for example, the spore-containing structures, asci and basidia, can be used in the identification of ascomycetes and basidiomycetes, respectively. Fungi employ two mating systems: heterothallic species allow mating only between individuals of the opposite mating type, whereas homothallic species can mate, and sexually reproduce, with any other individual or itself.

 

Most fungi have both a haploid and a diploid stage in their life cycles. In sexually reproducing fungi, compatible individuals may combine by fusing their hyphae together into an interconnected network; this process, anastomosis, is required for the initiation of the sexual cycle. Many ascomycetes and basidiomycetes go through a dikaryotic stage, in which the nuclei inherited from the two parents do not combine immediately after cell fusion, but remain separate in the hyphal cells (see heterokaryosis).

 

In ascomycetes, dikaryotic hyphae of the hymenium (the spore-bearing tissue layer) form a characteristic hook (crozier) at the hyphal septum. During cell division, the formation of the hook ensures proper distribution of the newly divided nuclei into the apical and basal hyphal compartments. An ascus (plural asci) is then formed, in which karyogamy (nuclear fusion) occurs. Asci are embedded in an ascocarp, or fruiting body. Karyogamy in the asci is followed immediately by meiosis and the production of ascospores. After dispersal, the ascospores may germinate and form a new haploid mycelium.

 

Sexual reproduction in basidiomycetes is similar to that of the ascomycetes. Compatible haploid hyphae fuse to produce a dikaryotic mycelium. However, the dikaryotic phase is more extensive in the basidiomycetes, often also present in the vegetatively growing mycelium. A specialized anatomical structure, called a clamp connection, is formed at each hyphal septum. As with the structurally similar hook in the ascomycetes, the clamp connection in the basidiomycetes is required for controlled transfer of nuclei during cell division, to maintain the dikaryotic stage with two genetically different nuclei in each hyphal compartment. A basidiocarp is formed in which club-like structures known as basidia generate haploid basidiospores after karyogamy and meiosis. The most commonly known basidiocarps are mushrooms, but they may also take other forms (see Morphology section).

 

In fungi formerly classified as Zygomycota, haploid hyphae of two individuals fuse, forming 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.

 

Spore dispersal

The spores of most of the researched species of fungi are transported by wind. Such species often produce dry or hydrophobic spores that do not absorb water and are readily scattered by raindrops, for example. In other species, both asexual and sexual spores or sporangiospores are often actively dispersed by forcible ejection from their reproductive structures. This ejection ensures exit of the spores from the reproductive structures as well as traveling through the air over long distances.

 

Specialized mechanical and physiological mechanisms, as well as spore surface structures (such as hydrophobins), enable efficient spore ejection. For example, the structure of the spore-bearing cells in some ascomycete species is such that the buildup of substances affecting cell volume and fluid balance enables the explosive discharge of spores into the air. The forcible discharge of single spores termed ballistospores involves formation of a small drop of water (Buller's drop), which upon contact with the spore leads to its projectile release with an initial acceleration of more than 10,000 g; the net result is that the spore is ejected 0.01–0.02 cm, sufficient distance for it to fall through the gills or pores into the air below. Other fungi, like the puffballs, rely on alternative mechanisms for spore release, such as external mechanical forces. The hydnoid fungi (tooth fungi) produce spores on pendant, tooth-like or spine-like projections. The bird's nest fungi use the force of falling water drops to liberate the spores from cup-shaped fruiting bodies. Another strategy is seen in the stinkhorns, a group of fungi with lively colors and putrid odor that attract insects to disperse their spores.

 

Homothallism

In homothallic sexual reproduction, two haploid nuclei derived from the same individual fuse to form a zygote that can then undergo meiosis. Homothallic fungi include species with an Aspergillus-like asexual stage (anamorphs) occurring in numerous different genera, several species of the ascomycete genus Cochliobolus, and the ascomycete Pneumocystis jirovecii. The earliest mode of sexual reproduction among eukaryotes was likely homothallism, that is, self-fertile unisexual reproduction.

 

Other sexual processes

Besides regular sexual reproduction with meiosis, certain fungi, such as those in the genera Penicillium and Aspergillus, may exchange genetic material via parasexual processes, initiated by anastomosis between hyphae and plasmogamy of fungal cells. The frequency and relative importance of parasexual events is unclear and may be lower than other sexual processes. It is known to play a role in intraspecific hybridization and is likely required for hybridization between species, which has been associated with major events in fungal evolution.

 

Evolution

In contrast to plants 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. Often recovered from a permineralized plant or animal host, these samples are typically studied by making thin-section preparations that can be examined with light microscopy or transmission electron microscopy. Researchers study compression fossils by dissolving the surrounding matrix with acid and then using light or scanning electron microscopy to examine surface details.

 

The earliest fossils possessing features typical of fungi date to the Paleoproterozoic era, some 2,400 million years ago (Ma); these multicellular benthic organisms had filamentous structures capable of anastomosis. Other studies (2009) estimate the arrival of fungal organisms at about 760–1060 Ma on the basis of comparisons of the rate of evolution in closely related groups. The oldest fossilizied mycelium to be identified from its molecular composition is between 715 and 810 million years old. For much of the Paleozoic Era (542–251 Ma), the fungi appear to have been aquatic and consisted of organisms similar to the extant chytrids in having flagellum-bearing spores. The evolutionary adaptation from an aquatic to a terrestrial lifestyle necessitated a diversification of ecological strategies for obtaining nutrients, including parasitism, saprobism, and the development of mutualistic relationships such as mycorrhiza and lichenization. Studies suggest that the ancestral ecological state of the Ascomycota was saprobism, and that independent lichenization events have occurred multiple times.

 

In May 2019, scientists reported the discovery of a fossilized fungus, named Ourasphaira giraldae, in the Canadian Arctic, that may have grown on land a billion years ago, well before plants were living on land. Pyritized fungus-like microfossils preserved in the basal Ediacaran Doushantuo Formation (~635 Ma) have been reported in South China. Earlier, it had been presumed that the fungi colonized the land during the Cambrian (542–488.3 Ma), also long before land plants. Fossilized hyphae and spores recovered from the Ordovician of Wisconsin (460 Ma) resemble modern-day Glomerales, and existed at a time when the land flora likely consisted of only non-vascular bryophyte-like plants. Prototaxites, which was probably a fungus or lichen, would have been the tallest organism of the late Silurian and early Devonian. Fungal fossils do not become common and uncontroversial until the early Devonian (416–359.2 Ma), when they occur abundantly in the Rhynie chert, mostly as Zygomycota and Chytridiomycota. At about this same time, approximately 400 Ma, the Ascomycota and Basidiomycota diverged, and all modern classes of fungi were present by the Late Carboniferous (Pennsylvanian, 318.1–299 Ma).

 

Lichens formed a component of the early terrestrial ecosystems, and the estimated age of the oldest terrestrial lichen fossil is 415 Ma; this date roughly corresponds to the age of the oldest known sporocarp fossil, a Paleopyrenomycites species found in the Rhynie Chert. The oldest fossil with microscopic features resembling modern-day basidiomycetes is Palaeoancistrus, found permineralized with a fern from the Pennsylvanian. Rare in the fossil record are the Homobasidiomycetes (a taxon roughly equivalent to the mushroom-producing species of the Agaricomycetes). Two amber-preserved specimens provide evidence that the earliest known mushroom-forming fungi (the extinct species Archaeomarasmius leggetti) appeared during the late Cretaceous, 90 Ma.

 

Some time after the Permian–Triassic extinction event (251.4 Ma), a fungal spike (originally thought to be an extraordinary abundance of fungal spores in sediments) formed, suggesting that fungi were the dominant life form at this time, representing nearly 100% of the available fossil record for this period. However, the relative proportion of fungal spores relative to spores formed by algal species is difficult to assess, the spike did not appear worldwide, and in many places it did not fall on the Permian–Triassic boundary.

 

Sixty-five million years ago, immediately after the Cretaceous–Paleogene extinction event that famously killed off most dinosaurs, there was a dramatic increase in evidence of fungi; apparently the death of most plant and animal species led to a huge fungal bloom like "a massive compost heap".

 

Taxonomy

Although commonly included in botany curricula and textbooks, fungi are more closely related to animals than to plants and are placed with the animals in the monophyletic group of opisthokonts. Analyses using molecular phylogenetics support a monophyletic origin of fungi. The taxonomy of fungi is in a state of constant flux, especially due to research based on DNA comparisons. These current phylogenetic analyses often overturn classifications based on older and sometimes less discriminative methods based on morphological features and biological species concepts obtained from experimental matings.

 

There is no unique generally accepted system at the higher taxonomic levels and there are frequent name changes at every level, from species upwards. Efforts among researchers are now underway to establish and encourage usage of a unified and more consistent nomenclature. Until relatively recent (2012) changes to the International Code of Nomenclature for algae, fungi and plants, fungal species could also have multiple scientific names depending on their life cycle and mode (sexual or asexual) of reproduction. Web sites such as Index Fungorum and MycoBank are officially recognized nomenclatural repositories and list current names of fungal species (with cross-references to older synonyms).

 

The 2007 classification of Kingdom Fungi is the result of a large-scale collaborative research effort involving dozens of mycologists and other scientists working on fungal taxonomy. It recognizes seven phyla, two of which—the Ascomycota and the Basidiomycota—are contained within a branch representing subkingdom Dikarya, the most species rich and familiar group, including all the mushrooms, most food-spoilage molds, most plant pathogenic fungi, and the beer, wine, and bread yeasts. The accompanying cladogram depicts the major fungal taxa and their relationship to opisthokont and unikont organisms, based on the work of Philippe Silar, "The Mycota: A Comprehensive Treatise on Fungi as Experimental Systems for Basic and Applied Research" and Tedersoo et al. 2018. The lengths of the branches are not proportional to evolutionary distances.

 

The major phyla (sometimes called divisions) of fungi have been classified mainly on the basis of characteristics of their sexual reproductive structures. As of 2019, nine major lineages have been identified: Opisthosporidia, Chytridiomycota, Neocallimastigomycota, Blastocladiomycota, Zoopagomycotina, Mucoromycota, Glomeromycota, Ascomycota and Basidiomycota.

 

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). Previously considered to be "primitive" protozoa, they are now thought to be either a basal branch of the Fungi, or a sister group–each other's closest evolutionary relative.

 

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.

 

The Blastocladiomycota were previously considered a taxonomic clade within the Chytridiomycota. 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.

 

The Neocallimastigomycota were earlier placed in the phylum Chytridiomycota. 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). They lack mitochondria but contain hydrogenosomes of mitochondrial origin. As in the related chrytrids, neocallimastigomycetes form zoospores that are posteriorly uniflagellate or polyflagellate.

 

Microscopic view of a layer of translucent grayish cells, some containing small dark-color spheres

Arbuscular mycorrhiza seen under microscope. Flax root cortical cells containing paired arbuscules.

Cross-section of a cup-shaped structure showing locations of developing meiotic asci (upper edge of cup, left side, arrows pointing to two gray cells containing four and two small circles), sterile hyphae (upper edge of cup, right side, arrows pointing to white cells with a single small circle in them), and mature asci (upper edge of cup, pointing to two gray cells with eight small circles in them)

Diagram of an apothecium (the typical cup-like reproductive structure of Ascomycetes) showing sterile tissues as well as developing and mature asci.

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. The symbiotic association between the Glomeromycota and plants is ancient, with evidence dating to 400 million years ago. 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. 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. 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. Medically relevant genera include Mucor, Rhizomucor, and Rhizopus.

 

The Ascomycota, commonly known as sac fungi or ascomycetes, constitute the largest taxonomic group within the Eumycota. 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. 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).

 

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, human commensal species of the genus Malassezia, and the opportunistic human pathogen, Cryptococcus neoformans.

 

Fungus-like organisms

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 an ambiregnal, duplicated taxonomy.

 

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 take in 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.

 

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.

 

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. Alternatively, Rozella can be classified as a basal fungal group.

 

The nucleariids may be the next sister group to the eumycete clade, and as such could be included in an expanded fungal kingdom. Many Actinomycetales (Actinomycetota), a group with many filamentous bacteria, were also long believed to be fungi.

 

Ecology

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 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.

 

Symbiosis

Many fungi have important symbiotic relationships with organisms from most if not all kingdoms. 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.

 

With plants

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.

 

A microscopic view of blue-stained cells, some with dark wavy lines in them

The dark filaments are hyphae of the endophytic fungus Epichloë coenophiala in the intercellular spaces of tall fescue leaf sheath tissue

The mycorrhizal symbiosis is ancient, dating back to at least 400 million years. 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. The fungal partners may also mediate plant-to-plant transfer of carbohydrates and other nutrients. Such mycorrhizal communities are called "common mycorrhizal networks". A special case of mycorrhiza is myco-heterotrophy, whereby the plant parasitizes the fungus, obtaining all of its nutrients from its fungal symbiont. Some fungal species inhabit the tissues inside roots, stems, and leaves, in which case they are called endophytes. 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.

 

With algae and cyanobacteria

A green, leaf-like structure attached to a tree, with a pattern of ridges and depression on the bottom surface

The lichen Lobaria pulmonaria, a symbiosis of fungal, algal, and cyanobacterial species

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. Lichens occur in every ecosystem on all continents, play a key role in soil formation and the initiation of biological succession, and are prominent in some extreme environments, including polar, alpine, and semiarid desert regions. They are able to grow on inhospitable surfaces, including bare soil, rocks, tree bark, wood, shells, barnacles and leaves. 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 27% of known fungi—more than 19,400 species—are lichenized. 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.

 

With insects

Many insects also engage in mutualistic relationships with fungi. Several groups of ants cultivate fungi in the order Chaetothyriales for several purposes: as a food source, as a structural component of their nests, and as a part of an ant/plant symbiosis in the domatia (tiny chambers in plants that house arthropods). Ambrosia beetles cultivate various species of fungi in the bark of trees that they infest. 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. 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. Termites on the African savannah are also known to cultivate fungi, 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. Fungi growing in dead wood are essential for xylophagous insects (e.g. woodboring beetles). They deliver nutrients needed by xylophages to nutritionally scarce dead wood. Thanks to this nutritional enrichment the larvae of the woodboring insect is able to grow and develop to adulthood. 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.

 

A thin brown stick positioned horizontally with roughly two dozen clustered orange-red leaves originating from a single point in the middle of the stick. These orange leaves are three to four times larger than the few other green leaves growing out of the stick, and are covered on the lower leaf surface with hundreds of tiny bumps. The background shows the green leaves and branches of neighboring shrubs.

The plant pathogen Puccinia magellanicum (calafate rust) causes the defect known as witch's broom, seen here on a barberry shrub in Chile.

 

Gram stain of Candida albicans from a vaginal swab from a woman with candidiasis, showing hyphae, and chlamydospores, which are 2–4 µm in diameter.

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, tree pathogens such as Ophiostoma ulmi and Ophiostoma novo-ulmi causing Dutch elm disease, Cryphonectria parasitica responsible for chestnut blight, and Phymatotrichopsis omnivora causing Texas Root Rot, and plant pathogens in the genera Fusarium, Ustilago, Alternaria, and Cochliobolus. 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. Many fungi that are plant pathogens, such as Magnaporthe oryzae, can switch from being biotrophic (parasitic on living plants) to being necrotrophic (feeding on the dead tissues of plants they have killed). This same principle is applied to fungi-feeding parasites, including Asterotremella albida, which feeds on the fruit bodies of other fungi both while they are living and after they are dead.

 

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, Histoplasma, and Pneumocystis. 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. Fungal spores are also a cause of allergies, and fungi from different taxonomic groups can evoke allergic reactions.

 

As targets of mycoparasites

Organisms that parasitize fungi are known as mycoparasitic organisms. About 300 species of fungi and fungus-like organisms, belonging to 13 classes and 113 genera, are used as biocontrol agents against plant fungal diseases. Fungi can also act as mycoparasites or antagonists of other fungi, such as Hypomyces chrysospermus, which grows on bolete mushrooms. Fungi can also become the target of infection by mycoviruses.

 

Communication

Main article: Mycorrhizal networks

There appears to be electrical communication between fungi in word-like components according to spiking characteristics.

 

Possible impact on climate

According to a study published in the academic journal Current Biology, fungi can soak from the atmosphere around 36% of global fossil fuel greenhouse gas emissions.

 

Mycotoxins

(6aR,9R)-N-((2R,5S,10aS,10bS)-5-benzyl-10b-hydroxy-2-methyl-3,6-dioxooctahydro-2H-oxazolo[3,2-a] pyrrolo[2,1-c]pyrazin-2-yl)-7-methyl-4,6,6a,7,8,9-hexahydroindolo[4,3-fg] quinoline-9-carboxamide

Ergotamine, a major mycotoxin produced by Claviceps species, which if ingested can cause gangrene, convulsions, and hallucinations

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. 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.

 

Mycotoxins are secondary metabolites (or natural products), and research has established the existence of biochemical pathways solely for the purpose of producing mycotoxins and other natural products in fungi. Mycotoxins may provide fitness benefits in terms of physiological adaptation, competition with other microbes and fungi, and protection from consumption (fungivory). Many fungal secondary metabolites (or derivatives) are used medically, as described under Human use below.

 

Pathogenic mechanisms

Ustilago maydis is a pathogenic plant fungus that causes smut disease in maize and teosinte. Plants have evolved efficient defense systems against pathogenic microbes such as U. maydis. A rapid defense reaction after pathogen attack is the oxidative burst where the plant produces reactive oxygen species 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. Furthermore, U. maydis has a well-established recombinational DNA repair system which acts during mitosis and meiosis. The system may assist the pathogen in surviving DNA damage arising from the host plant's oxidative defensive response to infection.

 

Cryptococcus neoformans is an encapsulated yeast that can live in both plants and animals. C. neoformans usually infects the lungs, where it is phagocytosed by alveolar macrophages. 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. 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. 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.

 

Human use

See also: Human interactions with fungi

Microscopic view of five spherical structures; one of the spheres is considerably smaller than the rest and attached to one of the larger spheres

Saccharomyces cerevisiae cells shown with DIC microscopy

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. Methods have been developed for genetic engineering of fungi, enabling metabolic engineering of fungal species. For example, genetic modification of yeast species—which are easy to grow at fast rates in large fermentation vessels—has opened up ways of pharmaceutical production that are potentially more efficient than production by the original source organisms. Fungi-based industries are sometimes considered to be a major part of a growing bioeconomy, with applications under research and development including use for textiles, meat substitution and general fungal biotechnology.

 

Therapeutic uses

Modern chemotherapeutics

Many species produce metabolites that are major sources of pharmacologically active drugs.

 

Antibiotics

Particularly important are the antibiotics, including the 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. 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. 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.

 

Other

Other drugs produced by fungi include griseofulvin isolated from Penicillium griseofulvum, used to treat fungal infections, 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. Psilocybin from fungi is investigated for therapeutic use and appears to cause global increases in brain network integration. Fungi produce compounds that inhibit viruses and cancer cells. 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. In Europe and Japan, polysaccharide-K (brand name Krestin), a chemical derived from Trametes versicolor, is an approved adjuvant for cancer therapy.

 

Traditional medicine

Upper surface view of a kidney-shaped fungus, brownish-red with a lighter yellow-brown margin, and a somewhat varnished or shiny appearance

Two dried yellow-orange caterpillars, one with a curly grayish fungus growing out of one of its ends. The grayish fungus is roughly equal to or slightly greater in length than the caterpillar, and tapers in thickness to a narrow end.

The fungi Ganoderma lucidum (left) and Ophiocordyceps sinensis (right) are used in traditional medicine practices

Certain mushrooms are used as supposed therapeutics in folk medicine practices, such as traditional Chinese medicine. Mushrooms with a history of such use include Agaricus subrufescens, Ganoderma lucidum, and Ophiocordyceps sinensis.

 

Cultured foods

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. Yeast species of the genus Saccharomyces are also used to produce alcoholic beverages through fermentation. Shoyu koji mold (Aspergillus oryzae) is an essential ingredient in brewing Shoyu (soy sauce) and sake, and the preparation of miso while Rhizopus species are used for making tempeh. 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. Quorn, a meat substitute, is made from Fusarium venenatum.

Underneath the bark on a log...

Dutch nature -

  

The fruit bodies of the gloomy honey fungus are bundled together and are brown in color. Scales are on the stem and the hat. The honey fungus is well distinguished by the cuff (collar) underneath the hat. The fruit bodies are visible around the stamina, on the trunk, but also on the trunk from autumn. Characteristic of the honey mushroom are the black rhizomorphs (shoelaces) that are visible on the tree or in the ground.

 

The gloomy honey fungus infects the tree through wounds to roots. The fungus then touches the cambium, after which this outer layer of the tree dies. Often the cambium's attack occurs in wood root of the carrots and rootstocks. Over time, the condition of the tree will deteriorate and this may ultimately result in the death of the tree. Among the dead bark, sometimes the black rhizomorphs are visible and often white mycelial strings (the roots of a fungus).

Done in Ai, Finalized in Photoshop.

 

Fungal entities that root into sorrowful minds.

 

Griefbinders are sentient, parasitic mycelial intelligences native to the emotional decay zones of Thornbleed Vale. Rather than consuming flesh, they root into the grief-nodes of conscious beings — memories of betrayal, regret, abandonment — and feed on their resonance.

 

Psychological Infiltration

 

Sympathetic Invasion: They do not attack physically. They empathize, echo, and expand sorrow until it consumes.

 

Spore of Reflection: Victims begin to hear their own regrets narrated back to them, reframed cruelly.

 

Cognition Mycelia: Neural filaments that bloom behind the eyes of the host, weaving memory and hallucination together.

 

🌿 Manifestation

 

Physical Description: Griefbinders resemble squat, low-slung creatures with bark-like skin crusted in fungal nodules and ringed caps. Their enormous, glassy black eyes dominate a skull-like face etched with natural ridges that suggest sorrowful expressions. A fringe of rootlike tendrils hangs beneath their head like a fungal beard. Their limbs are thick and clawed, not for attack, but for anchoring into the earth — or into you.

 

Roughly the size of a labrador — compact, sturdy, and unsettlingly present when still — large enough to loom, small enough to nestle beneath trees.

 

Moves with an unsettling stillness punctuated by twitchy, purposeful steps — as if every motion is filtered through the memory of loss. When it locks eyes with a target, it tilts its fungal head slightly, as if hearing a sorrow not yet spoken aloud.

 

Appear as lichenous growths, slowly shaping into silhouettes of people the host has lost or wronged.

 

Their form shifts with the emotional state of those nearby.

 

They emit a scent like rain on scorched stone — a memory you wish you forgot.

 

Their caps glisten with memory-sweat; veins run through their stalks like pulsing nerve-roots.

 

Tendrils move slowly, as if listening.

 

⚠️ Threats & Effects

 

Extended exposure causes emotional paralysis, recursive memory loops, and identity drift.

 

Some willingly bond with a Griefbinder to keep the memory of someone alive — at the cost of themselves.

 

Cannot be removed by force. Only release through truth, closure, or total emotional severance.

 

“They don’t feed on pain — they bloom through it.”

location: North America, Europe

edibility: Inedible

fungus colour: Yellow

normal size: 5-15cm

cap type: Other

stem type: Lateral, rudimentary or absent

spore colour: Purplish to black

habitat: Grows in woods, Grows on the ground

 

Scleroderma citrinum Pers. syn. S. aurantium (Vaill.) Pers. syn. S. vulgare Horn. Kartoffelbovist Scléroderme vulgaire, Scléroderme orangé Common Earthball. Fruit body 2–10cm across, subglobose, attached to the substrate by cord-like mycelial threads, wall dirty yellow to ochre-brown, thick and tough, coarsely scaly, breaking open irregularly to liberate the spores. Gleba purplish-black at first patterned by whitish veins, powdery when mature. Spores brown, globose, with a net-like ornamentation, 9–13m in diameter. Habitat on mossy or peaty ground on heaths or in rich woodland, especially on sandy soil. Season late summer to early winter. Common. Not edible. Distribution, America and Europe.

This species is sometimes parasitized by Boletus parasiticus.

 

info by Roger Phillips:

 

www.rogersmushrooms.com

 

Mycilium, OOAK Inamorata doll with Shani head sculpt in Ice resin. Mycilium is being displayed in the Hitogata Ten art doll exhibition in Tokyo.

 

She is a nude OOAK with a white mulberry silk wig with branching out braids and amazing 3D printed wings by @lbxcouture (Thank you for the wings LBX!).

 

I was playing with the idea that fairies were related to mushrooms and that she is symbiotically reaching out and connecting to the environment around her using mycelial networks .

 

There is a waiting list for the exhibition dolls. Email me with "waiting list" as topic to get a chance to buy her if she doesn't sell at the exhibition.

Mycilium, OOAK Inamorata doll with Shani head sculpt in Ice resin. Mycilium is being displayed in the Hitogata Ten art doll exhibition in Tokyo.

 

She is a nude OOAK with a white mulberry silk wig with branching out braids and amazing 3D printed wings by @lbxcouture (Thank you for the wings LBX!).

 

I was playing with the idea that fairies were related to mushrooms and that she is symbiotically reaching out and connecting to the environment around her using mycelial networks .

 

There is a waiting list for the exhibition dolls. Email me with "waiting list" as topic to get a chance to buy her if she doesn't sell at the exhibition.

Mycilium, OOAK Inamorata doll with Shani head sculpt in Ice resin. Mycilium is being displayed in the Hitogata Ten art doll exhibition in Tokyo.

 

She is a nude OOAK with a white mulberry silk wig with branching out braids and amazing 3D printed wings by @lbxcouture (Thank you for the wings LBX!).

 

I was playing with the idea that fairies were related to mushrooms and that she is symbiotically reaching out and connecting to the environment around her using mycelial networks .

 

There is a waiting list for the exhibition dolls. Email me with "waiting list" as topic to get a chance to buy her if she doesn't sell at the exhibition.

Mycelium, OOAK Inamorata art doll featuring Shani head sculpt in Ice resin.

 

I originally made Mycilium for the Hitogata Ten art doll exhibition in Tokyo. Since tattoos are a bit of a taboo in Japan, and I didn't want to risk upsetting the gallery, I didn't fulfil my original vision for he doll at the time. However, when this doll sold after the exhibition, I pitched the idea of painting her entire body with a mycelial network that has little bugs scuttling throughout it. I was so happy that the new owner loved the idea and I got to finish the concept. Her white mulberry silk hard cap wig with branching out braids repeats the branching patterns of a mycelial network.

 

The next preorder for Inamorata dolls is on April 22nd and will include this translucent Ice resin tone.

Mycilium, OOAK Inamorata doll with Shani head sculpt in Ice resin. Mycilium is being displayed in the Hitogata Ten art doll exhibition in Tokyo.

 

She is a nude OOAK with a white mulberry silk wig with branching out braids and amazing 3D printed wings by @lbxcouture (Thank you for the wings LBX!).

 

I was playing with the idea that fairies were related to mushrooms and that she is symbiotically reaching out and connecting to the environment around her using mycelial networks .

 

There is a waiting list for the exhibition dolls. Email me with "waiting list" as topic to get a chance to buy her if she doesn't sell at the exhibition.

Ok, back to October and the Fungi foray i went on.

 

This is Bovista Nigrescens or Brown puffball.

The fruiting body is 3 to 6 cm across and is attached to the substrate by a single mycelial cord which often breaks, leaving the fruit body free to roll about in the wind. The outer wall is white at first, but soon flakes off in large scales at maturity to expose the dark purple-brown to blackish inner wall that encloses the spore mass.

 

Mycilium, OOAK Inamorata doll with Shani head sculpt in Ice resin. Mycilium is being displayed in the Hitogata Ten art doll exhibition in Tokyo.

 

She is a nude OOAK with a white mulberry silk wig with branching out braids and amazing 3D printed wings by @lbxcouture (Thank you for the wings LBX!).

 

I was playing with the idea that fairies were related to mushrooms and that she is symbiotically reaching out and connecting to the environment around her using mycelial networks .

 

There is a waiting list for the exhibition dolls. Email me with "waiting list" as topic to get a chance to buy her if she doesn't sell at the exhibition.

   

Common Name: Lantana.

  

Genre: Lantana.

  

Family: Verbenaceae .

  

Etymology: Linnaeus was to pass this name ( which was previously attributed to Viburnum ) to the genre in question. Note the similarity of the leaves with those of Viburnum lantana .

  

Origin: Central and South America .

  

Description gender : includes about 150 species of plants in shrub , perennial , deciduous or persistent presenting simple, serrated margin , opposite or whorled . The flowers are tubular , formed by five irregular petals , and appear in corymbs axillary or ear . They have the particularity of being able to take a color and bloom as the days pass other shades . It is not uncommon , in fact, be able to see , on the same plant , flowers of different shades. Plants are suitable for growing outdoors in areas with a mild climate , otherwise it is better to raise them in a greenhouse or in an apartment .

  

Species and Varieties

Lantana camara : native to the tropical regions of America, this species has elliptical leaves , dark green in color , with an unpleasant odor. From May to August it produces tubular flowers , in corymbs globose, about 5 cm wide . , Which form the axils of the leaves. The color of the flowers darken as the days go from white to yellow to red brick. The fruit is represented by a dark berry toxic . It can reach heights ranging from 40 cm to 2 m . In the market there are different varieties among which are : " Crocea " , with sulfur yellow flowers that turn yellow saffron ; "Cloth of Gold" , from the bright yellow flowers ; " mutabilis " , which produces flowers that go from white to yellow to pink and lilac ; "Rose Queen" , with yellow flowers that turn pink ; " Spreading Sunset ' , characterized by a prostrate habit and flowers that , at first yellow , then become reddish ; " Snow White " with white flowers.

Lantana involucrata : a native of the West Indies , this shrub has opposite leaves , lanceolate . The flowers , which bloom in corymbs globular, are lilac with yellow throat . It can grow up to 1.5 m .

Lantana selloviana or L. montevidensis : native of Uruguay, this species has a habit semiprostrato long and thin branches which bear small leaves, ovate, hairy and toothed margins . From June to October-November produces inflorescences globose , 2-3 cm . , Consisting of small flowers , tubular and slightly rounded pink - lilac with yellow throat . It grows up to 20-30 cm . height.

Lantana trifolia : native to tropical America , this shrub reaches a meter in height and produces purple flowers .

  

Environmental requirements , substrate , fertilizer and special precautions

Temperature : The ideal temperature during the winter period is 6-10 ° C. With high temperatures the plant withers and becomes easy prey for parasites.

Light: very good, even in bright sunlight , however, be avoided in the months and during the hottest hours , especially in the presence of poor ventilation.

  

Watering and humidity : Water abundantly in summer and moderately in winter. The ambient humidity should be increased by any means , even wetting the floor of the greenhouse.

Soil : rich soil and humifero .

Fertilization and special precautions : from June to September fertilize the plants every two weeks . From May until early autumn can be transferred to the pots outdoors. With the end of the summer will be good to bring in a greenhouse.

Multiplication and pruning

Multiplication is sown in February, at a temperature of 16 ° C. The multiplication is done by taking cuttings in August servings of young branches , length of 8 cm. , Which will be put to root in bed for multiplication at a temperature of 15-18 ° C. The cuttings thus obtained allow themselves to toughen up in pots of 8 cm. in diameter ( filled with a compote made ​​from garden soil, peat, sand and soil ) up to February-March, when they will be subjected to shearing to encourage the formation of branches. Are transplanted in pots final spring.

Pruning: in reality the plants of this kind do not require a real pruning. You can use a shortening of the branches longer ( up to 10-15 cm . Length) developed in specimens too , to be made in February.

  

Diseases, pests and adversity

The white fly in greenhouses runs throughout their life cycle on the undersides of leaves : if the attack is massive , the leaves turn yellow , dry up and fall . The strong production of honeydew favors the development of sooty mold .

Damage can be caused by fungal diseases such as powdery mildew ( on the younger leaves forms a white mold , first as a stain and then as a veil covering ) , the leaf spot (leaf spots appear round, blackish -brown color , which tend to merged to form necrotic areas : laminae dry up ) and collar rot and root ( manifested by browning and softening, followed by formation of mycelial color creamy-white , the plant assumes a prostrate position ) .

 

   

In ancient times, only wild Reishi was available. Reishi was classified by color into 6 types: Red, Green, White, Black, Yellow, Purple.

In 1972, researchers at Kyoto University in Japan successfully cultivated Reishi in the laboratory. From a single species, Ganoderma Lucidum (Red Reishi), all six colors could be grown by varying the temperature, humidity, carbon dioxide content, and the available nutrients.

 

The six types of Reishi are thus shown to be one species.

Wild Reishi is extremely rare. Only one or two mushrooms can be found on a hill. Due to damage by insects and weather, the quality of wild Reishi is unpredictable. Only the fruit body can be harvested, when the active ingredients have already decreased. The dried mushrooms may not have the potency of the fresh mushroom. When buying wild Reishi, expertise is required in order not to confuse Reishi with the hundreds of other mushrooms (some of which are poisonous). After it matures, the fruit body is hardened by fibers which makes it more difficult to extract and digest the active ingredients. The spores are of microscopic dimensions, similar to the size of bacteria. They are protected by two layers of hardened cell walls. These cell walls trap the active ingredients inside and are indigestible.

 

Although wild Reishi fruit body and the spores are all effective products, our ancestors had to use a large quantity of Reishi to get a little benefit. As it was impossible to cultivate, this rare mushroom was available only to emperors.

Modern bio-engineering technology has made Reishi available to the general public in large quantities.

 

The quality can be carefully controlled by providing the best conditions and sufficient nutrients. Further investigations have discovered that the largest amount of active ingredients exist in the mycelium, and that the mycelium is more digestible. The extraction process can be timed at the precise stage when the mycelium contains the largest amount of active ingredients. Fresh mycelium is available, and there is no chance for mistaken identity. Without the obstacles of the fibers, the extraction is more complete and the extract is more digestible. Therefore, the latest research on the medicinal properties of Reishi are done on the mycelial extracts.

 

Reishi is now available in capsule or tablet form, which makes it possible to avoid the bitter taste and standardize the dosage. However, not all Reishi capsules are the same. Some capsules are made from the fruit body, which contain a large quantity of dietary fibers. Some capsules contain other herbs, which may lead to side-effects. Some capsules made from the mycelium contain also the grain from which the mycelium is grown (so only a small fraction of the capsule is actually Reishi mycelium). The differences can be readily identified by tasting the powder. Pure Reishi has an intense, pure bitter taste. The best Reishi capsule is the extract of pure Reishi mycelium without the grain.

Several manufacturers have printed misleading brochures to promote the use of their own products. Such conflicting information can be confusing. The intelligent consumer will be careful in checking the source of information. Third-party documentation, especially those written by scientists, are the only reliable source.

An excellent definitive work on Reishi is "Reishi Mushroom, herb of spiritual potency and medical wonder", written by Dr. Terry Willard, Ph.D., member of the Canadian Government's Expert Advisory Committee on Herbs and Botanical Preparations. Also very informative is the Chinese books "Lingzhi and Health Vol. I-III" edited by Dr. Shiuh-Sheng Lee, Professor of Biochemistry at National Yang-Ming University School of Medicine, Taipei, Taiwan.

 

Reishi polysaccharides have high molecular weights of up to 1,050,000, which makes absorptiion by the intestines difficult. Dr. Fukumi Morishige, M.D. found that patients given large doses of Reishi (2-9g/day) had diarrhea, but when a large dose of vitamin C was also given at the same time, there was no diarrhea. Vitamin C breaks down the high molecular weight polysaccharides to a molecular weight of around 30,000, so that they are much more easily absorbed by the intestines.

For general health maintenance and prevention, take 2 Reishi mycelium capsules twice per day on an empty stomach. Also take 500 mg of Vitamin C per day after breakfast.

When a discomfort occurs from minor illness, increase the dosage to 2-3 capsules 3 times per day. Also take 1,000 mg of Vitamin C per day after meals.

For insomnia, take 4-5 capsules before bedtime. Take 2 in the morning. Also take 1,000 mg of Vitamin C per day after meals.

When a serious illness occurs, increase the dosage to 4 capsules 4 times per day. Also take 2,000-3,000 mg of Vitamin C per day after meals.

Some people may experience Vertigo Reaction when they first take Reishi. During the first week, they may experience slight swelling of the head or dizziness. Arthritis patient may experienced a temporary increase in the pain level. This is a cleansing reaction, often called a "healing crisis". Continue to take Reishi and the symptoms will be alleviated within a week.

 

Mycelium, OOAK Inamorata art doll featuring Shani head sculpt in Ice resin.

 

I originally made Mycilium for the Hitogata Ten art doll exhibition in Tokyo. Since tattoos are a bit of a taboo in Japan, and I didn't want to risk upsetting the gallery, I didn't fulfil my original vision for he doll at the time. However, when this doll sold after the exhibition, I pitched the idea of painting her entire body with a mycelial network that has little bugs scuttling throughout it. I was so happy that the new owner loved the idea and I got to finish the concept. Her white mulberry silk hard cap wig with branching out braids repeats the branching patterns of a mycelial network.

 

The next preorder for Inamorata dolls is on April 22nd and will include this translucent Ice resin tone.

Fairy-Ring on the Downs

 

The settlement spreads outwards:

a mycelial sprawl, marked

by a rampart of darker green.

It is like a loving thought:

always seeking, with white

fingers. It creeps, because

it fears it isn't quite

decent - worms furtively

toward ends beyond itself.

It's the grim persistence

that is so self-sacrificial,

so primed for survival.

 

Hills are scarred with ramparts:

fossil ripplings-outwards.

 

Poem by Giles Watson, 2013. The picture shows a "fairy ring", probably caused by the fungus Marasmius oreades, in the turf of a downland "gallop" on the Ridgeway between Ogbourne St. George and the ancient Iron Age hill-fort, Barbury Castle. Another hill-fort, Liddington Castle, can be seen on the edge of the escarpment on the horizon.

 

Geastrum minimum Schwein., syn.: Geastrum marginatum Vittad.

Family: Geastraceae Corda

EN: Tiny earthstar, DE: Kleiner Erdstern, Zwergerdstern

Slo.: drobcena zvezdica

 

Dat.: Aug. 31. 2021

Lat.: 46.37725 Long.: 13.743703

Code: Bot_1398/2021_DSC08259

 

Habitat: At the base of a mountain ravine, slightly inclined mountain slope; east aspect; about 30 m distance to river Soča and about 10 m higher; close to a Pinus sp. forest; fine, older colluvial, calcareous deposits partly overgrown (Erica carnea dominant); dry, open, sunny, relatively warm place; exposed to direct rain; average precipitations ~ 3.000 mm/year, average temperature 7-9 deg C, elevation 595 m (1.950 feet), alpine phytogeographical region.

 

Substratum: soil, fine sandy colluvial deposits among larger rocks.

 

Place: Lower Trenta valley, south of village Trenta, right bank of river Soča, East Julian Alps, Posočje, Slovenia EC

 

Comments: This find is the smallest earthstar I've ever found. Based on the number of non-hygroscopic exoperidial rays, persistent mycelial layer, encrusted with debris, very small endoperidial body, which is stalked and provided with an apophysis, whitish crystals on endoperidial body surface and a clearly delimited, rather flat, fibrillose, not furrowed peristome I am pretty sure this is Geastrum minimum. The fungus is a rather rare find in Slovenia.

 

Growing in a group of three sporocarps. Endoperidium diameter; 6, 7 and 9 mm; SP on mass black. Spores verrucose, dark. Dimensions: (3.8) 4.4 - 5.5 (5.6) × (3.6) 4.2 - 5.2 (5.7) µm; Q = (0.9) 1 - 1.1 (1.2); N = 52; Me = 4.9 × 4.7 µm; Qe = 1. Olympus CH20, NEA 100x/1.25, magnification 1.000 x, oil (spores); NEA 40x/0.65, magnification 400x (capillitium), in water, fresh material. AmScope MA500 digital camera.

 

Ref.:

(1) www.funghiitaliani.it/topic/88099-geastrum-minimum/ (last accessed Oct.17. 2021)

(2) T. Lӕssøe, J.H. Petersen, Fungi of temperate Europe, Vol. 2., Princeton University Press (2019), p 1249.

(3) G.J. Krieglsteiner (Hrsg.), Die Grosspilze Baden-Württembergs, Band 2., Ulmer (2000), p 112.

(4) www.123pilze.de/DreamHC/Download/KleinerErdstern.htm (last accessed Oct.17. 2021),

(5) J.C. Zamora1, F.D. Calonge, M.P. Martín, Integrative taxonomy reveals an unexpected diversity in Geastrum section Geastrum (Geastrales, Basidiomycota), Persoonia 34, 2015: 130–165; geastrum_sect_geastrum_1422974412556.pdf,

(6) M. Jeppson, R. H. Nilsson, E. Larsson, 2013, European earthstars in Geastraceae (Geastrales, Phallomycetidae) – a systematic approach using morphology and molecular sequence data; Systematics and Biodiversity, 11:4, pp 437-465, Natural History Museum, Taylor & Francis Online.

(5) S. Buczacki, Collins Fungi Guide, Collins (2012), p 440.

(6) D. Arora, Mushrooms Demystified, Ten Speed Press, Berkeley (1986), p 702.

(7) W. Rothmaler, Exkursionsflora von Deutschland, Vol.1, Niedere Pflanzen, Elsevier, 3. Auflage, (1994), p 519.

  

Mycelium, OOAK Inamorata art doll featuring Shani head sculpt in Ice resin.

 

I originally made Mycilium for the Hitogata Ten art doll exhibition in Tokyo. Since tattoos are a bit of a taboo in Japan, and I didn't want to risk upsetting the gallery, I didn't fulfil my original vision for he doll at the time. However, when this doll sold after the exhibition, I pitched the idea of painting her entire body with a mycelial network that has little bugs scuttling throughout it. I was so happy that the new owner loved the idea and I got to finish the concept. Her white mulberry silk hard cap wig with branching out braids repeats the branching patterns of a mycelial network.

 

The next preorder for Inamorata dolls is on April 22nd and will include this translucent Ice resin tone.

Mycelium, OOAK Inamorata art doll featuring Shani head sculpt in Ice resin.

 

I originally made Mycilium for the Hitogata Ten art doll exhibition in Tokyo. Since tattoos are a bit of a taboo in Japan, and I didn't want to risk upsetting the gallery, I didn't fulfil my original vision for he doll at the time. However, when this doll sold after the exhibition, I pitched the idea of painting her entire body with a mycelial network that has little bugs scuttling throughout it. I was so happy that the new owner loved the idea and I got to finish the concept. Her white mulberry silk hard cap wig with branching out braids repeats the branching patterns of a mycelial network.

 

The next preorder for Inamorata dolls is on April 22nd and will include this translucent Ice resin tone.

A fungus (pl.: fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as one of the traditional eukaryotic kingdoms, along with Animalia, Plantae and either Protista or Protozoa and Chromista.

 

A characteristic that places fungi in a different kingdom from plants, bacteria, and some protists is chitin in their cell walls. Fungi, like animals, are heterotrophs; they acquire their food by absorbing dissolved molecules, typically by secreting digestive enzymes into their environment. Fungi do not photosynthesize. Growth is their means of mobility, except for spores (a few of which are flagellated), which may travel through the air or water. Fungi are the principal decomposers in ecological systems. These and other differences place fungi in a single group of related organisms, named the Eumycota (true fungi or Eumycetes), that share a common ancestor (i.e. they form a monophyletic group), an interpretation that is also strongly supported by molecular phylogenetics. This fungal group is distinct from the structurally similar myxomycetes (slime molds) and oomycetes (water molds). The discipline of biology devoted to the study of fungi is known as mycology (from the Greek μύκης mykes, mushroom). In the past mycology was regarded as a branch of botany, although it is now known that fungi are genetically more closely related to animals than to plants.

 

Abundant worldwide, most fungi are inconspicuous because of the small size of their structures, and their cryptic lifestyles in soil or on dead matter. Fungi include symbionts of plants, animals, or other fungi and also parasites. They may become noticeable when fruiting, either as mushrooms or as molds. Fungi perform an essential role in the decomposition of organic matter and have fundamental roles in nutrient cycling and exchange in the environment. They have long been used as a direct source of human food, in the form of mushrooms and truffles; as a leavening agent for bread; and in the fermentation of various food products, such as wine, beer, and soy sauce. Since the 1940s, fungi have been used for the production of antibiotics, and, more recently, various enzymes produced by fungi are used industrially and in detergents. Fungi are also used as biological pesticides to control weeds, plant diseases, and insect pests. Many species produce bioactive compounds called mycotoxins, such as alkaloids and polyketides, that are toxic to animals, including humans. The fruiting structures of a few species contain psychotropic compounds and are consumed recreationally or in traditional spiritual ceremonies. Fungi can break down manufactured materials and buildings, and become significant pathogens of humans and other animals. Losses of crops due to fungal diseases (e.g., rice blast disease) or food spoilage can have a large impact on human food supplies and local economies.

 

The fungus kingdom encompasses an enormous diversity of taxa with varied ecologies, life cycle strategies, and morphologies ranging from unicellular aquatic chytrids to large mushrooms. However, little is known of the true biodiversity of the fungus kingdom, which has been estimated at 2.2 million to 3.8 million species. Of these, only about 148,000 have been described, with over 8,000 species known to be detrimental to plants and at least 300 that can be pathogenic to humans. Ever since the pioneering 18th and 19th century taxonomical works of Carl Linnaeus, Christiaan Hendrik Persoon, and Elias Magnus Fries, fungi have been classified according to their morphology (e.g., characteristics such as spore color or microscopic features) or physiology. Advances in molecular genetics have opened the way for DNA analysis to be incorporated into taxonomy, which has sometimes challenged the historical groupings based on morphology and other traits. Phylogenetic studies published in the first decade of the 21st century have helped reshape the classification within the fungi kingdom, which is divided into one subkingdom, seven phyla, and ten subphyla.

 

Etymology

The English word fungus is directly adopted from the Latin fungus (mushroom), used in the writings of Horace and Pliny. This in turn is derived from the Greek word sphongos (σφόγγος 'sponge'), which refers to the macroscopic structures and morphology of mushrooms and molds; the root is also used in other languages, such as the German Schwamm ('sponge') and Schimmel ('mold').

 

The word mycology is derived from the Greek mykes (μύκης 'mushroom') and logos (λόγος 'discourse'). It denotes the scientific study of fungi. The Latin adjectival form of "mycology" (mycologicæ) appeared as early as 1796 in a book on the subject by Christiaan Hendrik Persoon. The word appeared in English as early as 1824 in a book by Robert Kaye Greville. In 1836 the English naturalist Miles Joseph Berkeley's publication The English Flora of Sir James Edward Smith, Vol. 5. also refers to mycology as the study of fungi.

 

A group of all the fungi present in a particular region is known as mycobiota (plural noun, no singular). The term mycota is often used for this purpose, but many authors use it as a synonym of Fungi. The word funga has been proposed as a less ambiguous term morphologically similar to fauna and flora. The Species Survival Commission (SSC) of the International Union for Conservation of Nature (IUCN) in August 2021 asked that the phrase fauna and flora be replaced by fauna, flora, and funga.

 

Characteristics

 

Fungal hyphae cells

Hyphal wall

Septum

Mitochondrion

Vacuole

Ergosterol crystal

Ribosome

Nucleus

Endoplasmic reticulum

Lipid body

Plasma membrane

Spitzenkörper

Golgi apparatus

 

Fungal cell cycle showing Dikaryons typical of Higher Fungi

Before the introduction of molecular methods for phylogenetic analysis, taxonomists considered fungi to be members of the plant kingdom because of similarities in lifestyle: both fungi and plants are mainly immobile, and have similarities in general morphology and growth habitat. Although inaccurate, the common misconception that fungi are plants persists among the general public due to their historical classification, as well as several similarities. Like plants, fungi often grow in soil and, in the case of mushrooms, form conspicuous fruit bodies, which sometimes resemble plants such as mosses. The fungi are now considered a separate kingdom, distinct from both plants and animals, from which they appear to have diverged around one billion years ago (around the start of the Neoproterozoic Era). Some morphological, biochemical, and genetic features are shared with other organisms, while others are unique to the fungi, clearly separating them from the other kingdoms:

 

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. 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).

With animals: Fungi lack chloroplasts and are heterotrophic organisms and so require preformed organic compounds as energy sources.

With plants: Fungi have a cell wall and vacuoles. They reproduce by both sexual and asexual means, and like basal plant groups (such as ferns and mosses) produce spores. Similar to mosses and algae, fungi typically have haploid nuclei.

With euglenoids and bacteria: Higher fungi, euglenoids, and some bacteria produce the amino acid L-lysine in specific biosynthesis steps, called the α-aminoadipate pathway.

The cells of most fungi grow as tubular, elongated, and thread-like (filamentous) structures called hyphae, which may contain multiple nuclei and extend by growing at their tips. Each tip contains a set of aggregated vesicles—cellular structures consisting of proteins, lipids, and other organic molecules—called the Spitzenkörper. Both fungi and oomycetes grow as filamentous hyphal cells. In contrast, similar-looking organisms, such as filamentous green algae, grow by repeated cell division within a chain of cells. There are also single-celled fungi (yeasts) that do not form hyphae, and some fungi have both hyphal and yeast forms.

In common with some plant and animal species, more than one hundred fungal species display bioluminescence.

Unique features:

 

Some species grow as unicellular yeasts that reproduce by budding or fission. Dimorphic fungi can switch between a yeast phase and a hyphal phase in response to environmental conditions.

The fungal cell wall is made of a chitin-glucan complex; while glucans are also found in plants and chitin in the exoskeleton of arthropods, fungi are the only organisms that combine these two structural molecules in their cell wall. Unlike those of plants and oomycetes, fungal cell walls do not contain cellulose.

A whitish fan or funnel-shaped mushroom growing at the base of a tree.

Omphalotus nidiformis, a bioluminescent mushroom

Most fungi lack an efficient system for the long-distance transport of water and nutrients, such as the xylem and phloem in many plants. To overcome this limitation, some fungi, such as Armillaria, form rhizomorphs, which resemble and perform functions similar to the roots of plants. As eukaryotes, fungi possess a biosynthetic pathway for producing terpenes that uses mevalonic acid and pyrophosphate as chemical building blocks. Plants and some other organisms have an additional terpene biosynthesis pathway in their chloroplasts, a structure that fungi and animals do not have. Fungi produce several secondary metabolites that are similar or identical in structure to those made by plants. Many of 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.

 

Diversity

Fungi have a worldwide distribution, and grow in a wide range of habitats, including extreme environments such as deserts or areas with high salt concentrations or ionizing radiation, as well as in deep sea sediments. Some can survive the intense UV and cosmic radiation encountered during space travel. Most grow in terrestrial environments, though several species live partly or solely in aquatic habitats, such as the chytrid fungi Batrachochytrium dendrobatidis and B. salamandrivorans, parasites that have been responsible for a worldwide decline in amphibian populations. These organisms spend part of their life cycle as a motile zoospore, enabling them to propel itself through water and enter their amphibian host. Other examples of aquatic fungi include those living in hydrothermal areas of the ocean.

 

As of 2020, around 148,000 species of fungi have been described by taxonomists, but the global biodiversity of the fungus kingdom is not fully understood. A 2017 estimate suggests there may be between 2.2 and 3.8 million species The number of new fungi species discovered yearly has increased from 1,000 to 1,500 per year about 10 years ago, to about 2000 with a peak of more than 2,500 species in 2016. In the year 2019, 1882 new species of fungi were described, and it was estimated that more than 90% of fungi remain unknown The following year, 2905 new species were described—the highest annual record of new fungus names. In mycology, species have historically been distinguished by a variety of methods and concepts. Classification based on morphological characteristics, such as the size and shape of spores or fruiting structures, has traditionally dominated fungal taxonomy. 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.

 

Mycology

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.

 

The use of fungi by humans dates back to prehistory; Ötzi the Iceman, a well-preserved mummy of a 5,300-year-old Neolithic man found frozen in the Austrian Alps, carried two species of polypore mushrooms that may have been used as tinder (Fomes fomentarius), or for medicinal purposes (Piptoporus betulinus). Ancient peoples have used fungi as food sources—often unknowingly—for millennia, in the preparation of leavened bread and fermented juices. Some of the oldest written records contain references to the destruction of crops that were probably caused by pathogenic fungi.

 

History

Mycology became a systematic science after the development of the microscope 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. Micheli not only observed spores but also showed that, under the proper conditions, they could be induced into growing into the same species of fungi from which they originated. Extending the use of the binomial system of nomenclature introduced by Carl Linnaeus in his Species plantarum (1753), the Dutch Christiaan Hendrik Persoon (1761–1836) established the first classification of mushrooms with such skill as to be considered a founder of modern mycology. Later, Elias Magnus Fries (1794–1878) further elaborated the classification of fungi, using spore color and microscopic characteristics, methods still used by taxonomists today. Other notable early contributors to mycology in the 17th–19th and early 20th centuries include Miles Joseph Berkeley, August Carl Joseph Corda, Anton de Bary, the brothers Louis René and Charles Tulasne, Arthur H. R. Buller, Curtis G. Lloyd, and Pier Andrea Saccardo. In the 20th and 21st centuries, advances in biochemistry, genetics, molecular biology, biotechnology, DNA sequencing and phylogenetic analysis has provided new insights into fungal relationships and biodiversity, and has challenged traditional morphology-based groupings in fungal taxonomy.

 

Morphology

Microscopic structures

Monochrome micrograph showing Penicillium hyphae as long, transparent, tube-like structures a few micrometres across. Conidiophores branch out laterally from the hyphae, terminating in bundles of phialides on which spherical condidiophores are arranged like beads on a string. Septa are faintly visible as dark lines crossing the hyphae.

An environmental isolate of Penicillium

Hypha

Conidiophore

Phialide

Conidia

Septa

Most fungi grow as hyphae, which are cylindrical, thread-like structures 2–10 µm in diameter and up to several centimeters in length. Hyphae grow at their tips (apices); new hyphae are typically formed by emergence of new tips along existing hyphae by a process called branching, or occasionally growing hyphal tips fork, giving rise to two parallel-growing hyphae. Hyphae also sometimes fuse when they come into contact, a process called hyphal fusion (or anastomosis). These growth processes lead to the development of a mycelium, an interconnected network of hyphae. Hyphae can be either septate or coenocytic. Septate hyphae are divided into compartments separated by cross walls (internal cell walls, called septa, that are formed at right angles to the cell wall giving the hypha its shape), with each compartment containing one or more nuclei; coenocytic hyphae are not compartmentalized. Septa have pores that allow cytoplasm, organelles, and sometimes nuclei to pass through; an example is the dolipore septum in fungi of the phylum Basidiomycota. Coenocytic hyphae are in essence multinucleate supercells.

 

Many species have developed specialized hyphal structures for nutrient uptake from living hosts; examples include haustoria in plant-parasitic species of most fungal phyla,[63] and arbuscules of several mycorrhizal fungi, which penetrate into the host cells to consume nutrients.

 

Although fungi are opisthokonts—a grouping of evolutionarily related organisms broadly characterized by a single posterior flagellum—all phyla except for the chytrids have lost their posterior flagella. Fungi are unusual among the eukaryotes in having a cell wall that, in addition to glucans (e.g., β-1,3-glucan) and other typical components, also contains the biopolymer chitin.

 

Macroscopic structures

Fungal mycelia can become visible to the naked eye, for example, on various surfaces and substrates, such as damp walls and spoiled food, where they are commonly called 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. Some individual fungal colonies can reach extraordinary dimensions and ages as in the case of a clonal colony of Armillaria solidipes, which extends over an area of more than 900 ha (3.5 square miles), with an estimated age of nearly 9,000 years.

 

The apothecium—a specialized structure important in sexual reproduction in the ascomycetes—is a cup-shaped fruit body that is often macroscopic and holds the hymenium, a layer of tissue containing the spore-bearing cells. The fruit bodies of the basidiomycetes (basidiocarps) and some ascomycetes can sometimes grow very large, and many are well known as mushrooms.

 

Growth and physiology

Time-lapse photography sequence of a peach becoming progressively discolored and disfigured

Mold growth covering a decaying peach. The frames were taken approximately 12 hours apart over a period of six days.

The growth of fungi as hyphae on or in solid substrates or as single cells in aquatic environments is adapted for the efficient extraction of nutrients, because these growth forms have high surface area to volume ratios. Hyphae are specifically adapted for growth on solid surfaces, and to invade substrates and tissues. They can exert large penetrative mechanical forces; for example, many plant pathogens, including Magnaporthe grisea, form a structure called an appressorium that evolved to puncture plant tissues.[71] The pressure generated by the appressorium, directed against the plant epidermis, can exceed 8 megapascals (1,200 psi).[71] The filamentous fungus Paecilomyces lilacinus uses a similar structure to penetrate the eggs of nematodes.

 

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

 

Fungi are traditionally considered heterotrophs, organisms that rely solely on carbon fixed by other organisms for metabolism. Fungi have evolved a high degree of metabolic versatility that allows them to use a diverse range of organic substrates for growth, including simple compounds such as nitrate, ammonia, acetate, or ethanol. In some species the pigment melanin may play a role in extracting energy from ionizing radiation, such as gamma radiation. This form of "radiotrophic" growth has been described for only a few species, the effects on growth rates are small, and the underlying biophysical and biochemical processes are not well known. This process might bear similarity to CO2 fixation via visible light, but instead uses ionizing radiation as a source of energy.

 

Reproduction

Two thickly stemmed brownish mushrooms with scales on the upper surface, growing out of a tree trunk

Polyporus squamosus

Fungal reproduction is complex, reflecting the differences in lifestyles and genetic makeup within this diverse kingdom of organisms. It is estimated that a third of all fungi reproduce using more than one method of propagation; for example, reproduction may occur in two well-differentiated stages within the life cycle of a species, the teleomorph (sexual reproduction) and the anamorph (asexual reproduction). Environmental conditions trigger genetically determined developmental states that lead to the creation of specialized structures for sexual or asexual reproduction. These structures aid reproduction by efficiently dispersing spores or spore-containing propagules.

 

Asexual reproduction

Asexual reproduction occurs via vegetative spores (conidia) or through mycelial fragmentation. Mycelial fragmentation occurs when a fungal mycelium separates into pieces, and each component grows into a separate mycelium. Mycelial fragmentation and vegetative spores maintain clonal populations adapted to a specific niche, and allow more rapid dispersal than sexual reproduction. The "Fungi imperfecti" (fungi lacking the perfect or sexual stage) or Deuteromycota comprise all the species that lack an observable sexual cycle. Deuteromycota (alternatively known as Deuteromycetes, conidial fungi, or mitosporic fungi) is not an accepted taxonomic clade and is now taken to mean simply fungi that lack a known sexual stage.

 

Sexual reproduction

See also: Mating in fungi and Sexual selection in fungi

Sexual reproduction with meiosis has been directly observed in all fungal phyla except Glomeromycota (genetic analysis suggests meiosis in Glomeromycota as well). It differs in many aspects from sexual reproduction in animals or plants. Differences also exist between fungal groups and can be used to discriminate species by morphological differences in sexual structures and reproductive strategies. Mating experiments between fungal isolates may identify species on the basis of biological species concepts. The major fungal groupings have initially been delineated based on the morphology of their sexual structures and spores; for example, the spore-containing structures, asci and basidia, can be used in the identification of ascomycetes and basidiomycetes, respectively. Fungi employ two mating systems: heterothallic species allow mating only between individuals of the opposite mating type, whereas homothallic species can mate, and sexually reproduce, with any other individual or itself.

 

Most fungi have both a haploid and a diploid stage in their life cycles. In sexually reproducing fungi, compatible individuals may combine by fusing their hyphae together into an interconnected network; this process, anastomosis, is required for the initiation of the sexual cycle. Many ascomycetes and basidiomycetes go through a dikaryotic stage, in which the nuclei inherited from the two parents do not combine immediately after cell fusion, but remain separate in the hyphal cells (see heterokaryosis).

 

In ascomycetes, dikaryotic hyphae of the hymenium (the spore-bearing tissue layer) form a characteristic hook (crozier) at the hyphal septum. During cell division, the formation of the hook ensures proper distribution of the newly divided nuclei into the apical and basal hyphal compartments. An ascus (plural asci) is then formed, in which karyogamy (nuclear fusion) occurs. Asci are embedded in an ascocarp, or fruiting body. Karyogamy in the asci is followed immediately by meiosis and the production of ascospores. After dispersal, the ascospores may germinate and form a new haploid mycelium.

 

Sexual reproduction in basidiomycetes is similar to that of the ascomycetes. Compatible haploid hyphae fuse to produce a dikaryotic mycelium. However, the dikaryotic phase is more extensive in the basidiomycetes, often also present in the vegetatively growing mycelium. A specialized anatomical structure, called a clamp connection, is formed at each hyphal septum. As with the structurally similar hook in the ascomycetes, the clamp connection in the basidiomycetes is required for controlled transfer of nuclei during cell division, to maintain the dikaryotic stage with two genetically different nuclei in each hyphal compartment. A basidiocarp is formed in which club-like structures known as basidia generate haploid basidiospores after karyogamy and meiosis. The most commonly known basidiocarps are mushrooms, but they may also take other forms (see Morphology section).

 

In fungi formerly classified as Zygomycota, haploid hyphae of two individuals fuse, forming 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.

 

Spore dispersal

The spores of most of the researched species of fungi are transported by wind. Such species often produce dry or hydrophobic spores that do not absorb water and are readily scattered by raindrops, for example. In other species, both asexual and sexual spores or sporangiospores are often actively dispersed by forcible ejection from their reproductive structures. This ejection ensures exit of the spores from the reproductive structures as well as traveling through the air over long distances.

 

Specialized mechanical and physiological mechanisms, as well as spore surface structures (such as hydrophobins), enable efficient spore ejection. For example, the structure of the spore-bearing cells in some ascomycete species is such that the buildup of substances affecting cell volume and fluid balance enables the explosive discharge of spores into the air. The forcible discharge of single spores termed ballistospores involves formation of a small drop of water (Buller's drop), which upon contact with the spore leads to its projectile release with an initial acceleration of more than 10,000 g; the net result is that the spore is ejected 0.01–0.02 cm, sufficient distance for it to fall through the gills or pores into the air below. Other fungi, like the puffballs, rely on alternative mechanisms for spore release, such as external mechanical forces. The hydnoid fungi (tooth fungi) produce spores on pendant, tooth-like or spine-like projections. The bird's nest fungi use the force of falling water drops to liberate the spores from cup-shaped fruiting bodies. Another strategy is seen in the stinkhorns, a group of fungi with lively colors and putrid odor that attract insects to disperse their spores.

 

Homothallism

In homothallic sexual reproduction, two haploid nuclei derived from the same individual fuse to form a zygote that can then undergo meiosis. Homothallic fungi include species with an Aspergillus-like asexual stage (anamorphs) occurring in numerous different genera, several species of the ascomycete genus Cochliobolus, and the ascomycete Pneumocystis jirovecii. The earliest mode of sexual reproduction among eukaryotes was likely homothallism, that is, self-fertile unisexual reproduction.

 

Other sexual processes

Besides regular sexual reproduction with meiosis, certain fungi, such as those in the genera Penicillium and Aspergillus, may exchange genetic material via parasexual processes, initiated by anastomosis between hyphae and plasmogamy of fungal cells. The frequency and relative importance of parasexual events is unclear and may be lower than other sexual processes. It is known to play a role in intraspecific hybridization and is likely required for hybridization between species, which has been associated with major events in fungal evolution.

 

Evolution

In contrast to plants 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. Often recovered from a permineralized plant or animal host, these samples are typically studied by making thin-section preparations that can be examined with light microscopy or transmission electron microscopy. Researchers study compression fossils by dissolving the surrounding matrix with acid and then using light or scanning electron microscopy to examine surface details.

 

The earliest fossils possessing features typical of fungi date to the Paleoproterozoic era, some 2,400 million years ago (Ma); these multicellular benthic organisms had filamentous structures capable of anastomosis. Other studies (2009) estimate the arrival of fungal organisms at about 760–1060 Ma on the basis of comparisons of the rate of evolution in closely related groups. The oldest fossilizied mycelium to be identified from its molecular composition is between 715 and 810 million years old. For much of the Paleozoic Era (542–251 Ma), the fungi appear to have been aquatic and consisted of organisms similar to the extant chytrids in having flagellum-bearing spores. The evolutionary adaptation from an aquatic to a terrestrial lifestyle necessitated a diversification of ecological strategies for obtaining nutrients, including parasitism, saprobism, and the development of mutualistic relationships such as mycorrhiza and lichenization. Studies suggest that the ancestral ecological state of the Ascomycota was saprobism, and that independent lichenization events have occurred multiple times.

 

In May 2019, scientists reported the discovery of a fossilized fungus, named Ourasphaira giraldae, in the Canadian Arctic, that may have grown on land a billion years ago, well before plants were living on land. Pyritized fungus-like microfossils preserved in the basal Ediacaran Doushantuo Formation (~635 Ma) have been reported in South China. Earlier, it had been presumed that the fungi colonized the land during the Cambrian (542–488.3 Ma), also long before land plants. Fossilized hyphae and spores recovered from the Ordovician of Wisconsin (460 Ma) resemble modern-day Glomerales, and existed at a time when the land flora likely consisted of only non-vascular bryophyte-like plants. Prototaxites, which was probably a fungus or lichen, would have been the tallest organism of the late Silurian and early Devonian. Fungal fossils do not become common and uncontroversial until the early Devonian (416–359.2 Ma), when they occur abundantly in the Rhynie chert, mostly as Zygomycota and Chytridiomycota. At about this same time, approximately 400 Ma, the Ascomycota and Basidiomycota diverged, and all modern classes of fungi were present by the Late Carboniferous (Pennsylvanian, 318.1–299 Ma).

 

Lichens formed a component of the early terrestrial ecosystems, and the estimated age of the oldest terrestrial lichen fossil is 415 Ma; this date roughly corresponds to the age of the oldest known sporocarp fossil, a Paleopyrenomycites species found in the Rhynie Chert. The oldest fossil with microscopic features resembling modern-day basidiomycetes is Palaeoancistrus, found permineralized with a fern from the Pennsylvanian. Rare in the fossil record are the Homobasidiomycetes (a taxon roughly equivalent to the mushroom-producing species of the Agaricomycetes). Two amber-preserved specimens provide evidence that the earliest known mushroom-forming fungi (the extinct species Archaeomarasmius leggetti) appeared during the late Cretaceous, 90 Ma.

 

Some time after the Permian–Triassic extinction event (251.4 Ma), a fungal spike (originally thought to be an extraordinary abundance of fungal spores in sediments) formed, suggesting that fungi were the dominant life form at this time, representing nearly 100% of the available fossil record for this period. However, the relative proportion of fungal spores relative to spores formed by algal species is difficult to assess, the spike did not appear worldwide, and in many places it did not fall on the Permian–Triassic boundary.

 

Sixty-five million years ago, immediately after the Cretaceous–Paleogene extinction event that famously killed off most dinosaurs, there was a dramatic increase in evidence of fungi; apparently the death of most plant and animal species led to a huge fungal bloom like "a massive compost heap".

 

Taxonomy

Although commonly included in botany curricula and textbooks, fungi are more closely related to animals than to plants and are placed with the animals in the monophyletic group of opisthokonts. Analyses using molecular phylogenetics support a monophyletic origin of fungi. The taxonomy of fungi is in a state of constant flux, especially due to research based on DNA comparisons. These current phylogenetic analyses often overturn classifications based on older and sometimes less discriminative methods based on morphological features and biological species concepts obtained from experimental matings.

 

There is no unique generally accepted system at the higher taxonomic levels and there are frequent name changes at every level, from species upwards. Efforts among researchers are now underway to establish and encourage usage of a unified and more consistent nomenclature. Until relatively recent (2012) changes to the International Code of Nomenclature for algae, fungi and plants, fungal species could also have multiple scientific names depending on their life cycle and mode (sexual or asexual) of reproduction. Web sites such as Index Fungorum and MycoBank are officially recognized nomenclatural repositories and list current names of fungal species (with cross-references to older synonyms).

 

The 2007 classification of Kingdom Fungi is the result of a large-scale collaborative research effort involving dozens of mycologists and other scientists working on fungal taxonomy. It recognizes seven phyla, two of which—the Ascomycota and the Basidiomycota—are contained within a branch representing subkingdom Dikarya, the most species rich and familiar group, including all the mushrooms, most food-spoilage molds, most plant pathogenic fungi, and the beer, wine, and bread yeasts. The accompanying cladogram depicts the major fungal taxa and their relationship to opisthokont and unikont organisms, based on the work of Philippe Silar, "The Mycota: A Comprehensive Treatise on Fungi as Experimental Systems for Basic and Applied Research" and Tedersoo et al. 2018. The lengths of the branches are not proportional to evolutionary distances.

 

The major phyla (sometimes called divisions) of fungi have been classified mainly on the basis of characteristics of their sexual reproductive structures. As of 2019, nine major lineages have been identified: Opisthosporidia, Chytridiomycota, Neocallimastigomycota, Blastocladiomycota, Zoopagomycotina, Mucoromycota, Glomeromycota, Ascomycota and Basidiomycota.

 

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). Previously considered to be "primitive" protozoa, they are now thought to be either a basal branch of the Fungi, or a sister group–each other's closest evolutionary relative.

 

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.

 

The Blastocladiomycota were previously considered a taxonomic clade within the Chytridiomycota. 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.

 

The Neocallimastigomycota were earlier placed in the phylum Chytridiomycota. 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). They lack mitochondria but contain hydrogenosomes of mitochondrial origin. As in the related chrytrids, neocallimastigomycetes form zoospores that are posteriorly uniflagellate or polyflagellate.

 

Microscopic view of a layer of translucent grayish cells, some containing small dark-color spheres

Arbuscular mycorrhiza seen under microscope. Flax root cortical cells containing paired arbuscules.

Cross-section of a cup-shaped structure showing locations of developing meiotic asci (upper edge of cup, left side, arrows pointing to two gray cells containing four and two small circles), sterile hyphae (upper edge of cup, right side, arrows pointing to white cells with a single small circle in them), and mature asci (upper edge of cup, pointing to two gray cells with eight small circles in them)

Diagram of an apothecium (the typical cup-like reproductive structure of Ascomycetes) showing sterile tissues as well as developing and mature asci.

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. The symbiotic association between the Glomeromycota and plants is ancient, with evidence dating to 400 million years ago. 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. 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. 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. Medically relevant genera include Mucor, Rhizomucor, and Rhizopus.

 

The Ascomycota, commonly known as sac fungi or ascomycetes, constitute the largest taxonomic group within the Eumycota. 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. 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).

 

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, human commensal species of the genus Malassezia, and the opportunistic human pathogen, Cryptococcus neoformans.

 

Fungus-like organisms

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 an ambiregnal, duplicated taxonomy.

 

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 take in 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.

 

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.

 

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. Alternatively, Rozella can be classified as a basal fungal group.

 

The nucleariids may be the next sister group to the eumycete clade, and as such could be included in an expanded fungal kingdom. Many Actinomycetales (Actinomycetota), a group with many filamentous bacteria, were also long believed to be fungi.

 

Ecology

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 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.

 

Symbiosis

Many fungi have important symbiotic relationships with organisms from most if not all kingdoms. 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.

 

With plants

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.

 

A microscopic view of blue-stained cells, some with dark wavy lines in them

The dark filaments are hyphae of the endophytic fungus Epichloë coenophiala in the intercellular spaces of tall fescue leaf sheath tissue

The mycorrhizal symbiosis is ancient, dating back to at least 400 million years. 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. The fungal partners may also mediate plant-to-plant transfer of carbohydrates and other nutrients. Such mycorrhizal communities are called "common mycorrhizal networks". A special case of mycorrhiza is myco-heterotrophy, whereby the plant parasitizes the fungus, obtaining all of its nutrients from its fungal symbiont. Some fungal species inhabit the tissues inside roots, stems, and leaves, in which case they are called endophytes. 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.

 

With algae and cyanobacteria

A green, leaf-like structure attached to a tree, with a pattern of ridges and depression on the bottom surface

The lichen Lobaria pulmonaria, a symbiosis of fungal, algal, and cyanobacterial species

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. Lichens occur in every ecosystem on all continents, play a key role in soil formation and the initiation of biological succession, and are prominent in some extreme environments, including polar, alpine, and semiarid desert regions. They are able to grow on inhospitable surfaces, including bare soil, rocks, tree bark, wood, shells, barnacles and leaves. 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 27% of known fungi—more than 19,400 species—are lichenized. 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.

 

With insects

Many insects also engage in mutualistic relationships with fungi. Several groups of ants cultivate fungi in the order Chaetothyriales for several purposes: as a food source, as a structural component of their nests, and as a part of an ant/plant symbiosis in the domatia (tiny chambers in plants that house arthropods). Ambrosia beetles cultivate various species of fungi in the bark of trees that they infest. 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. 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. Termites on the African savannah are also known to cultivate fungi, 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. Fungi growing in dead wood are essential for xylophagous insects (e.g. woodboring beetles). They deliver nutrients needed by xylophages to nutritionally scarce dead wood. Thanks to this nutritional enrichment the larvae of the woodboring insect is able to grow and develop to adulthood. 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.

 

A thin brown stick positioned horizontally with roughly two dozen clustered orange-red leaves originating from a single point in the middle of the stick. These orange leaves are three to four times larger than the few other green leaves growing out of the stick, and are covered on the lower leaf surface with hundreds of tiny bumps. The background shows the green leaves and branches of neighboring shrubs.

The plant pathogen Puccinia magellanicum (calafate rust) causes the defect known as witch's broom, seen here on a barberry shrub in Chile.

 

Gram stain of Candida albicans from a vaginal swab from a woman with candidiasis, showing hyphae, and chlamydospores, which are 2–4 µm in diameter.

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, tree pathogens such as Ophiostoma ulmi and Ophiostoma novo-ulmi causing Dutch elm disease, Cryphonectria parasitica responsible for chestnut blight, and Phymatotrichopsis omnivora causing Texas Root Rot, and plant pathogens in the genera Fusarium, Ustilago, Alternaria, and Cochliobolus. 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. Many fungi that are plant pathogens, such as Magnaporthe oryzae, can switch from being biotrophic (parasitic on living plants) to being necrotrophic (feeding on the dead tissues of plants they have killed). This same principle is applied to fungi-feeding parasites, including Asterotremella albida, which feeds on the fruit bodies of other fungi both while they are living and after they are dead.

 

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, Histoplasma, and Pneumocystis. 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. Fungal spores are also a cause of allergies, and fungi from different taxonomic groups can evoke allergic reactions.

 

As targets of mycoparasites

Organisms that parasitize fungi are known as mycoparasitic organisms. About 300 species of fungi and fungus-like organisms, belonging to 13 classes and 113 genera, are used as biocontrol agents against plant fungal diseases. Fungi can also act as mycoparasites or antagonists of other fungi, such as Hypomyces chrysospermus, which grows on bolete mushrooms. Fungi can also become the target of infection by mycoviruses.

 

Communication

Main article: Mycorrhizal networks

There appears to be electrical communication between fungi in word-like components according to spiking characteristics.

 

Possible impact on climate

According to a study published in the academic journal Current Biology, fungi can soak from the atmosphere around 36% of global fossil fuel greenhouse gas emissions.

 

Mycotoxins

(6aR,9R)-N-((2R,5S,10aS,10bS)-5-benzyl-10b-hydroxy-2-methyl-3,6-dioxooctahydro-2H-oxazolo[3,2-a] pyrrolo[2,1-c]pyrazin-2-yl)-7-methyl-4,6,6a,7,8,9-hexahydroindolo[4,3-fg] quinoline-9-carboxamide

Ergotamine, a major mycotoxin produced by Claviceps species, which if ingested can cause gangrene, convulsions, and hallucinations

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. 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.

 

Mycotoxins are secondary metabolites (or natural products), and research has established the existence of biochemical pathways solely for the purpose of producing mycotoxins and other natural products in fungi. Mycotoxins may provide fitness benefits in terms of physiological adaptation, competition with other microbes and fungi, and protection from consumption (fungivory). Many fungal secondary metabolites (or derivatives) are used medically, as described under Human use below.

 

Pathogenic mechanisms

Ustilago maydis is a pathogenic plant fungus that causes smut disease in maize and teosinte. Plants have evolved efficient defense systems against pathogenic microbes such as U. maydis. A rapid defense reaction after pathogen attack is the oxidative burst where the plant produces reactive oxygen species 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. Furthermore, U. maydis has a well-established recombinational DNA repair system which acts during mitosis and meiosis. The system may assist the pathogen in surviving DNA damage arising from the host plant's oxidative defensive response to infection.

 

Cryptococcus neoformans is an encapsulated yeast that can live in both plants and animals. C. neoformans usually infects the lungs, where it is phagocytosed by alveolar macrophages. 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. 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. 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.

 

Human use

See also: Human interactions with fungi

Microscopic view of five spherical structures; one of the spheres is considerably smaller than the rest and attached to one of the larger spheres

Saccharomyces cerevisiae cells shown with DIC microscopy

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. Methods have been developed for genetic engineering of fungi, enabling metabolic engineering of fungal species. For example, genetic modification of yeast species—which are easy to grow at fast rates in large fermentation vessels—has opened up ways of pharmaceutical production that are potentially more efficient than production by the original source organisms. Fungi-based industries are sometimes considered to be a major part of a growing bioeconomy, with applications under research and development including use for textiles, meat substitution and general fungal biotechnology.

 

Therapeutic uses

Modern chemotherapeutics

Many species produce metabolites that are major sources of pharmacologically active drugs.

 

Antibiotics

Particularly important are the antibiotics, including the 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. 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. 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.

 

Other

Other drugs produced by fungi include griseofulvin isolated from Penicillium griseofulvum, used to treat fungal infections, 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. Psilocybin from fungi is investigated for therapeutic use and appears to cause global increases in brain network integration. Fungi produce compounds that inhibit viruses and cancer cells. 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. In Europe and Japan, polysaccharide-K (brand name Krestin), a chemical derived from Trametes versicolor, is an approved adjuvant for cancer therapy.

 

Traditional medicine

Upper surface view of a kidney-shaped fungus, brownish-red with a lighter yellow-brown margin, and a somewhat varnished or shiny appearance

Two dried yellow-orange caterpillars, one with a curly grayish fungus growing out of one of its ends. The grayish fungus is roughly equal to or slightly greater in length than the caterpillar, and tapers in thickness to a narrow end.

The fungi Ganoderma lucidum (left) and Ophiocordyceps sinensis (right) are used in traditional medicine practices

Certain mushrooms are used as supposed therapeutics in folk medicine practices, such as traditional Chinese medicine. Mushrooms with a history of such use include Agaricus subrufescens, Ganoderma lucidum, and Ophiocordyceps sinensis.

 

Cultured foods

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. Yeast species of the genus Saccharomyces are also used to produce alcoholic beverages through fermentation. Shoyu koji mold (Aspergillus oryzae) is an essential ingredient in brewing Shoyu (soy sauce) and sake, and the preparation of miso while Rhizopus species are used for making tempeh. 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. Quorn, a meat substitute, is made from Fusarium venenatum.

Mycilium, OOAK Inamorata doll with Shani head sculpt in Ice resin. Mycilium is being displayed in the Hitogata Ten art doll exhibition in Tokyo.

 

She is a nude OOAK with a white mulberry silk wig with branching out braids and amazing 3D printed wings by @lbxcouture (Thank you for the wings LBX!).

 

I was playing with the idea that fairies were related to mushrooms and that she is symbiotically reaching out and connecting to the environment around her using mycelial networks .

 

There is a waiting list for the exhibition dolls. Email me with "waiting list" as topic to get a chance to buy her if she doesn't sell at the exhibition.

A fungus (pl.: fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as one of the traditional eukaryotic kingdoms, along with Animalia, Plantae and either Protista or Protozoa and Chromista.

 

A characteristic that places fungi in a different kingdom from plants, bacteria, and some protists is chitin in their cell walls. Fungi, like animals, are heterotrophs; they acquire their food by absorbing dissolved molecules, typically by secreting digestive enzymes into their environment. Fungi do not photosynthesize. Growth is their means of mobility, except for spores (a few of which are flagellated), which may travel through the air or water. Fungi are the principal decomposers in ecological systems. These and other differences place fungi in a single group of related organisms, named the Eumycota (true fungi or Eumycetes), that share a common ancestor (i.e. they form a monophyletic group), an interpretation that is also strongly supported by molecular phylogenetics. This fungal group is distinct from the structurally similar myxomycetes (slime molds) and oomycetes (water molds). The discipline of biology devoted to the study of fungi is known as mycology (from the Greek μύκης mykes, mushroom). In the past mycology was regarded as a branch of botany, although it is now known that fungi are genetically more closely related to animals than to plants.

 

Abundant worldwide, most fungi are inconspicuous because of the small size of their structures, and their cryptic lifestyles in soil or on dead matter. Fungi include symbionts of plants, animals, or other fungi and also parasites. They may become noticeable when fruiting, either as mushrooms or as molds. Fungi perform an essential role in the decomposition of organic matter and have fundamental roles in nutrient cycling and exchange in the environment. They have long been used as a direct source of human food, in the form of mushrooms and truffles; as a leavening agent for bread; and in the fermentation of various food products, such as wine, beer, and soy sauce. Since the 1940s, fungi have been used for the production of antibiotics, and, more recently, various enzymes produced by fungi are used industrially and in detergents. Fungi are also used as biological pesticides to control weeds, plant diseases, and insect pests. Many species produce bioactive compounds called mycotoxins, such as alkaloids and polyketides, that are toxic to animals, including humans. The fruiting structures of a few species contain psychotropic compounds and are consumed recreationally or in traditional spiritual ceremonies. Fungi can break down manufactured materials and buildings, and become significant pathogens of humans and other animals. Losses of crops due to fungal diseases (e.g., rice blast disease) or food spoilage can have a large impact on human food supplies and local economies.

 

The fungus kingdom encompasses an enormous diversity of taxa with varied ecologies, life cycle strategies, and morphologies ranging from unicellular aquatic chytrids to large mushrooms. However, little is known of the true biodiversity of the fungus kingdom, which has been estimated at 2.2 million to 3.8 million species. Of these, only about 148,000 have been described, with over 8,000 species known to be detrimental to plants and at least 300 that can be pathogenic to humans. Ever since the pioneering 18th and 19th century taxonomical works of Carl Linnaeus, Christiaan Hendrik Persoon, and Elias Magnus Fries, fungi have been classified according to their morphology (e.g., characteristics such as spore color or microscopic features) or physiology. Advances in molecular genetics have opened the way for DNA analysis to be incorporated into taxonomy, which has sometimes challenged the historical groupings based on morphology and other traits. Phylogenetic studies published in the first decade of the 21st century have helped reshape the classification within the fungi kingdom, which is divided into one subkingdom, seven phyla, and ten subphyla.

 

Etymology

The English word fungus is directly adopted from the Latin fungus (mushroom), used in the writings of Horace and Pliny. This in turn is derived from the Greek word sphongos (σφόγγος 'sponge'), which refers to the macroscopic structures and morphology of mushrooms and molds; the root is also used in other languages, such as the German Schwamm ('sponge') and Schimmel ('mold').

 

The word mycology is derived from the Greek mykes (μύκης 'mushroom') and logos (λόγος 'discourse'). It denotes the scientific study of fungi. The Latin adjectival form of "mycology" (mycologicæ) appeared as early as 1796 in a book on the subject by Christiaan Hendrik Persoon. The word appeared in English as early as 1824 in a book by Robert Kaye Greville. In 1836 the English naturalist Miles Joseph Berkeley's publication The English Flora of Sir James Edward Smith, Vol. 5. also refers to mycology as the study of fungi.

 

A group of all the fungi present in a particular region is known as mycobiota (plural noun, no singular). The term mycota is often used for this purpose, but many authors use it as a synonym of Fungi. The word funga has been proposed as a less ambiguous term morphologically similar to fauna and flora. The Species Survival Commission (SSC) of the International Union for Conservation of Nature (IUCN) in August 2021 asked that the phrase fauna and flora be replaced by fauna, flora, and funga.

 

Characteristics

 

Fungal hyphae cells

Hyphal wall

Septum

Mitochondrion

Vacuole

Ergosterol crystal

Ribosome

Nucleus

Endoplasmic reticulum

Lipid body

Plasma membrane

Spitzenkörper

Golgi apparatus

 

Fungal cell cycle showing Dikaryons typical of Higher Fungi

Before the introduction of molecular methods for phylogenetic analysis, taxonomists considered fungi to be members of the plant kingdom because of similarities in lifestyle: both fungi and plants are mainly immobile, and have similarities in general morphology and growth habitat. Although inaccurate, the common misconception that fungi are plants persists among the general public due to their historical classification, as well as several similarities. Like plants, fungi often grow in soil and, in the case of mushrooms, form conspicuous fruit bodies, which sometimes resemble plants such as mosses. The fungi are now considered a separate kingdom, distinct from both plants and animals, from which they appear to have diverged around one billion years ago (around the start of the Neoproterozoic Era). Some morphological, biochemical, and genetic features are shared with other organisms, while others are unique to the fungi, clearly separating them from the other kingdoms:

 

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. 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).

With animals: Fungi lack chloroplasts and are heterotrophic organisms and so require preformed organic compounds as energy sources.

With plants: Fungi have a cell wall and vacuoles. They reproduce by both sexual and asexual means, and like basal plant groups (such as ferns and mosses) produce spores. Similar to mosses and algae, fungi typically have haploid nuclei.

With euglenoids and bacteria: Higher fungi, euglenoids, and some bacteria produce the amino acid L-lysine in specific biosynthesis steps, called the α-aminoadipate pathway.

The cells of most fungi grow as tubular, elongated, and thread-like (filamentous) structures called hyphae, which may contain multiple nuclei and extend by growing at their tips. Each tip contains a set of aggregated vesicles—cellular structures consisting of proteins, lipids, and other organic molecules—called the Spitzenkörper. Both fungi and oomycetes grow as filamentous hyphal cells. In contrast, similar-looking organisms, such as filamentous green algae, grow by repeated cell division within a chain of cells. There are also single-celled fungi (yeasts) that do not form hyphae, and some fungi have both hyphal and yeast forms.

In common with some plant and animal species, more than one hundred fungal species display bioluminescence.

Unique features:

 

Some species grow as unicellular yeasts that reproduce by budding or fission. Dimorphic fungi can switch between a yeast phase and a hyphal phase in response to environmental conditions.

The fungal cell wall is made of a chitin-glucan complex; while glucans are also found in plants and chitin in the exoskeleton of arthropods, fungi are the only organisms that combine these two structural molecules in their cell wall. Unlike those of plants and oomycetes, fungal cell walls do not contain cellulose.

A whitish fan or funnel-shaped mushroom growing at the base of a tree.

Omphalotus nidiformis, a bioluminescent mushroom

Most fungi lack an efficient system for the long-distance transport of water and nutrients, such as the xylem and phloem in many plants. To overcome this limitation, some fungi, such as Armillaria, form rhizomorphs, which resemble and perform functions similar to the roots of plants. As eukaryotes, fungi possess a biosynthetic pathway for producing terpenes that uses mevalonic acid and pyrophosphate as chemical building blocks. Plants and some other organisms have an additional terpene biosynthesis pathway in their chloroplasts, a structure that fungi and animals do not have. Fungi produce several secondary metabolites that are similar or identical in structure to those made by plants. Many of 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.

 

Diversity

Fungi have a worldwide distribution, and grow in a wide range of habitats, including extreme environments such as deserts or areas with high salt concentrations or ionizing radiation, as well as in deep sea sediments. Some can survive the intense UV and cosmic radiation encountered during space travel. Most grow in terrestrial environments, though several species live partly or solely in aquatic habitats, such as the chytrid fungi Batrachochytrium dendrobatidis and B. salamandrivorans, parasites that have been responsible for a worldwide decline in amphibian populations. These organisms spend part of their life cycle as a motile zoospore, enabling them to propel itself through water and enter their amphibian host. Other examples of aquatic fungi include those living in hydrothermal areas of the ocean.

 

As of 2020, around 148,000 species of fungi have been described by taxonomists, but the global biodiversity of the fungus kingdom is not fully understood. A 2017 estimate suggests there may be between 2.2 and 3.8 million species The number of new fungi species discovered yearly has increased from 1,000 to 1,500 per year about 10 years ago, to about 2000 with a peak of more than 2,500 species in 2016. In the year 2019, 1882 new species of fungi were described, and it was estimated that more than 90% of fungi remain unknown The following year, 2905 new species were described—the highest annual record of new fungus names. In mycology, species have historically been distinguished by a variety of methods and concepts. Classification based on morphological characteristics, such as the size and shape of spores or fruiting structures, has traditionally dominated fungal taxonomy. 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.

 

Mycology

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.

 

The use of fungi by humans dates back to prehistory; Ötzi the Iceman, a well-preserved mummy of a 5,300-year-old Neolithic man found frozen in the Austrian Alps, carried two species of polypore mushrooms that may have been used as tinder (Fomes fomentarius), or for medicinal purposes (Piptoporus betulinus). Ancient peoples have used fungi as food sources—often unknowingly—for millennia, in the preparation of leavened bread and fermented juices. Some of the oldest written records contain references to the destruction of crops that were probably caused by pathogenic fungi.

 

History

Mycology became a systematic science after the development of the microscope 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. Micheli not only observed spores but also showed that, under the proper conditions, they could be induced into growing into the same species of fungi from which they originated. Extending the use of the binomial system of nomenclature introduced by Carl Linnaeus in his Species plantarum (1753), the Dutch Christiaan Hendrik Persoon (1761–1836) established the first classification of mushrooms with such skill as to be considered a founder of modern mycology. Later, Elias Magnus Fries (1794–1878) further elaborated the classification of fungi, using spore color and microscopic characteristics, methods still used by taxonomists today. Other notable early contributors to mycology in the 17th–19th and early 20th centuries include Miles Joseph Berkeley, August Carl Joseph Corda, Anton de Bary, the brothers Louis René and Charles Tulasne, Arthur H. R. Buller, Curtis G. Lloyd, and Pier Andrea Saccardo. In the 20th and 21st centuries, advances in biochemistry, genetics, molecular biology, biotechnology, DNA sequencing and phylogenetic analysis has provided new insights into fungal relationships and biodiversity, and has challenged traditional morphology-based groupings in fungal taxonomy.

 

Morphology

Microscopic structures

Monochrome micrograph showing Penicillium hyphae as long, transparent, tube-like structures a few micrometres across. Conidiophores branch out laterally from the hyphae, terminating in bundles of phialides on which spherical condidiophores are arranged like beads on a string. Septa are faintly visible as dark lines crossing the hyphae.

An environmental isolate of Penicillium

Hypha

Conidiophore

Phialide

Conidia

Septa

Most fungi grow as hyphae, which are cylindrical, thread-like structures 2–10 µm in diameter and up to several centimeters in length. Hyphae grow at their tips (apices); new hyphae are typically formed by emergence of new tips along existing hyphae by a process called branching, or occasionally growing hyphal tips fork, giving rise to two parallel-growing hyphae. Hyphae also sometimes fuse when they come into contact, a process called hyphal fusion (or anastomosis). These growth processes lead to the development of a mycelium, an interconnected network of hyphae. Hyphae can be either septate or coenocytic. Septate hyphae are divided into compartments separated by cross walls (internal cell walls, called septa, that are formed at right angles to the cell wall giving the hypha its shape), with each compartment containing one or more nuclei; coenocytic hyphae are not compartmentalized. Septa have pores that allow cytoplasm, organelles, and sometimes nuclei to pass through; an example is the dolipore septum in fungi of the phylum Basidiomycota. Coenocytic hyphae are in essence multinucleate supercells.

 

Many species have developed specialized hyphal structures for nutrient uptake from living hosts; examples include haustoria in plant-parasitic species of most fungal phyla,[63] and arbuscules of several mycorrhizal fungi, which penetrate into the host cells to consume nutrients.

 

Although fungi are opisthokonts—a grouping of evolutionarily related organisms broadly characterized by a single posterior flagellum—all phyla except for the chytrids have lost their posterior flagella. Fungi are unusual among the eukaryotes in having a cell wall that, in addition to glucans (e.g., β-1,3-glucan) and other typical components, also contains the biopolymer chitin.

 

Macroscopic structures

Fungal mycelia can become visible to the naked eye, for example, on various surfaces and substrates, such as damp walls and spoiled food, where they are commonly called 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. Some individual fungal colonies can reach extraordinary dimensions and ages as in the case of a clonal colony of Armillaria solidipes, which extends over an area of more than 900 ha (3.5 square miles), with an estimated age of nearly 9,000 years.

 

The apothecium—a specialized structure important in sexual reproduction in the ascomycetes—is a cup-shaped fruit body that is often macroscopic and holds the hymenium, a layer of tissue containing the spore-bearing cells. The fruit bodies of the basidiomycetes (basidiocarps) and some ascomycetes can sometimes grow very large, and many are well known as mushrooms.

 

Growth and physiology

Time-lapse photography sequence of a peach becoming progressively discolored and disfigured

Mold growth covering a decaying peach. The frames were taken approximately 12 hours apart over a period of six days.

The growth of fungi as hyphae on or in solid substrates or as single cells in aquatic environments is adapted for the efficient extraction of nutrients, because these growth forms have high surface area to volume ratios. Hyphae are specifically adapted for growth on solid surfaces, and to invade substrates and tissues. They can exert large penetrative mechanical forces; for example, many plant pathogens, including Magnaporthe grisea, form a structure called an appressorium that evolved to puncture plant tissues.[71] The pressure generated by the appressorium, directed against the plant epidermis, can exceed 8 megapascals (1,200 psi).[71] The filamentous fungus Paecilomyces lilacinus uses a similar structure to penetrate the eggs of nematodes.

 

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

 

Fungi are traditionally considered heterotrophs, organisms that rely solely on carbon fixed by other organisms for metabolism. Fungi have evolved a high degree of metabolic versatility that allows them to use a diverse range of organic substrates for growth, including simple compounds such as nitrate, ammonia, acetate, or ethanol. In some species the pigment melanin may play a role in extracting energy from ionizing radiation, such as gamma radiation. This form of "radiotrophic" growth has been described for only a few species, the effects on growth rates are small, and the underlying biophysical and biochemical processes are not well known. This process might bear similarity to CO2 fixation via visible light, but instead uses ionizing radiation as a source of energy.

 

Reproduction

Two thickly stemmed brownish mushrooms with scales on the upper surface, growing out of a tree trunk

Polyporus squamosus

Fungal reproduction is complex, reflecting the differences in lifestyles and genetic makeup within this diverse kingdom of organisms. It is estimated that a third of all fungi reproduce using more than one method of propagation; for example, reproduction may occur in two well-differentiated stages within the life cycle of a species, the teleomorph (sexual reproduction) and the anamorph (asexual reproduction). Environmental conditions trigger genetically determined developmental states that lead to the creation of specialized structures for sexual or asexual reproduction. These structures aid reproduction by efficiently dispersing spores or spore-containing propagules.

 

Asexual reproduction

Asexual reproduction occurs via vegetative spores (conidia) or through mycelial fragmentation. Mycelial fragmentation occurs when a fungal mycelium separates into pieces, and each component grows into a separate mycelium. Mycelial fragmentation and vegetative spores maintain clonal populations adapted to a specific niche, and allow more rapid dispersal than sexual reproduction. The "Fungi imperfecti" (fungi lacking the perfect or sexual stage) or Deuteromycota comprise all the species that lack an observable sexual cycle. Deuteromycota (alternatively known as Deuteromycetes, conidial fungi, or mitosporic fungi) is not an accepted taxonomic clade and is now taken to mean simply fungi that lack a known sexual stage.

 

Sexual reproduction

See also: Mating in fungi and Sexual selection in fungi

Sexual reproduction with meiosis has been directly observed in all fungal phyla except Glomeromycota (genetic analysis suggests meiosis in Glomeromycota as well). It differs in many aspects from sexual reproduction in animals or plants. Differences also exist between fungal groups and can be used to discriminate species by morphological differences in sexual structures and reproductive strategies. Mating experiments between fungal isolates may identify species on the basis of biological species concepts. The major fungal groupings have initially been delineated based on the morphology of their sexual structures and spores; for example, the spore-containing structures, asci and basidia, can be used in the identification of ascomycetes and basidiomycetes, respectively. Fungi employ two mating systems: heterothallic species allow mating only between individuals of the opposite mating type, whereas homothallic species can mate, and sexually reproduce, with any other individual or itself.

 

Most fungi have both a haploid and a diploid stage in their life cycles. In sexually reproducing fungi, compatible individuals may combine by fusing their hyphae together into an interconnected network; this process, anastomosis, is required for the initiation of the sexual cycle. Many ascomycetes and basidiomycetes go through a dikaryotic stage, in which the nuclei inherited from the two parents do not combine immediately after cell fusion, but remain separate in the hyphal cells (see heterokaryosis).

 

In ascomycetes, dikaryotic hyphae of the hymenium (the spore-bearing tissue layer) form a characteristic hook (crozier) at the hyphal septum. During cell division, the formation of the hook ensures proper distribution of the newly divided nuclei into the apical and basal hyphal compartments. An ascus (plural asci) is then formed, in which karyogamy (nuclear fusion) occurs. Asci are embedded in an ascocarp, or fruiting body. Karyogamy in the asci is followed immediately by meiosis and the production of ascospores. After dispersal, the ascospores may germinate and form a new haploid mycelium.

 

Sexual reproduction in basidiomycetes is similar to that of the ascomycetes. Compatible haploid hyphae fuse to produce a dikaryotic mycelium. However, the dikaryotic phase is more extensive in the basidiomycetes, often also present in the vegetatively growing mycelium. A specialized anatomical structure, called a clamp connection, is formed at each hyphal septum. As with the structurally similar hook in the ascomycetes, the clamp connection in the basidiomycetes is required for controlled transfer of nuclei during cell division, to maintain the dikaryotic stage with two genetically different nuclei in each hyphal compartment. A basidiocarp is formed in which club-like structures known as basidia generate haploid basidiospores after karyogamy and meiosis. The most commonly known basidiocarps are mushrooms, but they may also take other forms (see Morphology section).

 

In fungi formerly classified as Zygomycota, haploid hyphae of two individuals fuse, forming 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.

 

Spore dispersal

The spores of most of the researched species of fungi are transported by wind. Such species often produce dry or hydrophobic spores that do not absorb water and are readily scattered by raindrops, for example. In other species, both asexual and sexual spores or sporangiospores are often actively dispersed by forcible ejection from their reproductive structures. This ejection ensures exit of the spores from the reproductive structures as well as traveling through the air over long distances.

 

Specialized mechanical and physiological mechanisms, as well as spore surface structures (such as hydrophobins), enable efficient spore ejection. For example, the structure of the spore-bearing cells in some ascomycete species is such that the buildup of substances affecting cell volume and fluid balance enables the explosive discharge of spores into the air. The forcible discharge of single spores termed ballistospores involves formation of a small drop of water (Buller's drop), which upon contact with the spore leads to its projectile release with an initial acceleration of more than 10,000 g; the net result is that the spore is ejected 0.01–0.02 cm, sufficient distance for it to fall through the gills or pores into the air below. Other fungi, like the puffballs, rely on alternative mechanisms for spore release, such as external mechanical forces. The hydnoid fungi (tooth fungi) produce spores on pendant, tooth-like or spine-like projections. The bird's nest fungi use the force of falling water drops to liberate the spores from cup-shaped fruiting bodies. Another strategy is seen in the stinkhorns, a group of fungi with lively colors and putrid odor that attract insects to disperse their spores.

 

Homothallism

In homothallic sexual reproduction, two haploid nuclei derived from the same individual fuse to form a zygote that can then undergo meiosis. Homothallic fungi include species with an Aspergillus-like asexual stage (anamorphs) occurring in numerous different genera, several species of the ascomycete genus Cochliobolus, and the ascomycete Pneumocystis jirovecii. The earliest mode of sexual reproduction among eukaryotes was likely homothallism, that is, self-fertile unisexual reproduction.

 

Other sexual processes

Besides regular sexual reproduction with meiosis, certain fungi, such as those in the genera Penicillium and Aspergillus, may exchange genetic material via parasexual processes, initiated by anastomosis between hyphae and plasmogamy of fungal cells. The frequency and relative importance of parasexual events is unclear and may be lower than other sexual processes. It is known to play a role in intraspecific hybridization and is likely required for hybridization between species, which has been associated with major events in fungal evolution.

 

Evolution

In contrast to plants 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. Often recovered from a permineralized plant or animal host, these samples are typically studied by making thin-section preparations that can be examined with light microscopy or transmission electron microscopy. Researchers study compression fossils by dissolving the surrounding matrix with acid and then using light or scanning electron microscopy to examine surface details.

 

The earliest fossils possessing features typical of fungi date to the Paleoproterozoic era, some 2,400 million years ago (Ma); these multicellular benthic organisms had filamentous structures capable of anastomosis. Other studies (2009) estimate the arrival of fungal organisms at about 760–1060 Ma on the basis of comparisons of the rate of evolution in closely related groups. The oldest fossilizied mycelium to be identified from its molecular composition is between 715 and 810 million years old. For much of the Paleozoic Era (542–251 Ma), the fungi appear to have been aquatic and consisted of organisms similar to the extant chytrids in having flagellum-bearing spores. The evolutionary adaptation from an aquatic to a terrestrial lifestyle necessitated a diversification of ecological strategies for obtaining nutrients, including parasitism, saprobism, and the development of mutualistic relationships such as mycorrhiza and lichenization. Studies suggest that the ancestral ecological state of the Ascomycota was saprobism, and that independent lichenization events have occurred multiple times.

 

In May 2019, scientists reported the discovery of a fossilized fungus, named Ourasphaira giraldae, in the Canadian Arctic, that may have grown on land a billion years ago, well before plants were living on land. Pyritized fungus-like microfossils preserved in the basal Ediacaran Doushantuo Formation (~635 Ma) have been reported in South China. Earlier, it had been presumed that the fungi colonized the land during the Cambrian (542–488.3 Ma), also long before land plants. Fossilized hyphae and spores recovered from the Ordovician of Wisconsin (460 Ma) resemble modern-day Glomerales, and existed at a time when the land flora likely consisted of only non-vascular bryophyte-like plants. Prototaxites, which was probably a fungus or lichen, would have been the tallest organism of the late Silurian and early Devonian. Fungal fossils do not become common and uncontroversial until the early Devonian (416–359.2 Ma), when they occur abundantly in the Rhynie chert, mostly as Zygomycota and Chytridiomycota. At about this same time, approximately 400 Ma, the Ascomycota and Basidiomycota diverged, and all modern classes of fungi were present by the Late Carboniferous (Pennsylvanian, 318.1–299 Ma).

 

Lichens formed a component of the early terrestrial ecosystems, and the estimated age of the oldest terrestrial lichen fossil is 415 Ma; this date roughly corresponds to the age of the oldest known sporocarp fossil, a Paleopyrenomycites species found in the Rhynie Chert. The oldest fossil with microscopic features resembling modern-day basidiomycetes is Palaeoancistrus, found permineralized with a fern from the Pennsylvanian. Rare in the fossil record are the Homobasidiomycetes (a taxon roughly equivalent to the mushroom-producing species of the Agaricomycetes). Two amber-preserved specimens provide evidence that the earliest known mushroom-forming fungi (the extinct species Archaeomarasmius leggetti) appeared during the late Cretaceous, 90 Ma.

 

Some time after the Permian–Triassic extinction event (251.4 Ma), a fungal spike (originally thought to be an extraordinary abundance of fungal spores in sediments) formed, suggesting that fungi were the dominant life form at this time, representing nearly 100% of the available fossil record for this period. However, the relative proportion of fungal spores relative to spores formed by algal species is difficult to assess, the spike did not appear worldwide, and in many places it did not fall on the Permian–Triassic boundary.

 

Sixty-five million years ago, immediately after the Cretaceous–Paleogene extinction event that famously killed off most dinosaurs, there was a dramatic increase in evidence of fungi; apparently the death of most plant and animal species led to a huge fungal bloom like "a massive compost heap".

 

Taxonomy

Although commonly included in botany curricula and textbooks, fungi are more closely related to animals than to plants and are placed with the animals in the monophyletic group of opisthokonts. Analyses using molecular phylogenetics support a monophyletic origin of fungi. The taxonomy of fungi is in a state of constant flux, especially due to research based on DNA comparisons. These current phylogenetic analyses often overturn classifications based on older and sometimes less discriminative methods based on morphological features and biological species concepts obtained from experimental matings.

 

There is no unique generally accepted system at the higher taxonomic levels and there are frequent name changes at every level, from species upwards. Efforts among researchers are now underway to establish and encourage usage of a unified and more consistent nomenclature. Until relatively recent (2012) changes to the International Code of Nomenclature for algae, fungi and plants, fungal species could also have multiple scientific names depending on their life cycle and mode (sexual or asexual) of reproduction. Web sites such as Index Fungorum and MycoBank are officially recognized nomenclatural repositories and list current names of fungal species (with cross-references to older synonyms).

 

The 2007 classification of Kingdom Fungi is the result of a large-scale collaborative research effort involving dozens of mycologists and other scientists working on fungal taxonomy. It recognizes seven phyla, two of which—the Ascomycota and the Basidiomycota—are contained within a branch representing subkingdom Dikarya, the most species rich and familiar group, including all the mushrooms, most food-spoilage molds, most plant pathogenic fungi, and the beer, wine, and bread yeasts. The accompanying cladogram depicts the major fungal taxa and their relationship to opisthokont and unikont organisms, based on the work of Philippe Silar, "The Mycota: A Comprehensive Treatise on Fungi as Experimental Systems for Basic and Applied Research" and Tedersoo et al. 2018. The lengths of the branches are not proportional to evolutionary distances.

 

The major phyla (sometimes called divisions) of fungi have been classified mainly on the basis of characteristics of their sexual reproductive structures. As of 2019, nine major lineages have been identified: Opisthosporidia, Chytridiomycota, Neocallimastigomycota, Blastocladiomycota, Zoopagomycotina, Mucoromycota, Glomeromycota, Ascomycota and Basidiomycota.

 

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). Previously considered to be "primitive" protozoa, they are now thought to be either a basal branch of the Fungi, or a sister group–each other's closest evolutionary relative.

 

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.

 

The Blastocladiomycota were previously considered a taxonomic clade within the Chytridiomycota. 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.

 

The Neocallimastigomycota were earlier placed in the phylum Chytridiomycota. 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). They lack mitochondria but contain hydrogenosomes of mitochondrial origin. As in the related chrytrids, neocallimastigomycetes form zoospores that are posteriorly uniflagellate or polyflagellate.

 

Microscopic view of a layer of translucent grayish cells, some containing small dark-color spheres

Arbuscular mycorrhiza seen under microscope. Flax root cortical cells containing paired arbuscules.

Cross-section of a cup-shaped structure showing locations of developing meiotic asci (upper edge of cup, left side, arrows pointing to two gray cells containing four and two small circles), sterile hyphae (upper edge of cup, right side, arrows pointing to white cells with a single small circle in them), and mature asci (upper edge of cup, pointing to two gray cells with eight small circles in them)

Diagram of an apothecium (the typical cup-like reproductive structure of Ascomycetes) showing sterile tissues as well as developing and mature asci.

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. The symbiotic association between the Glomeromycota and plants is ancient, with evidence dating to 400 million years ago. 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. 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. 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. Medically relevant genera include Mucor, Rhizomucor, and Rhizopus.

 

The Ascomycota, commonly known as sac fungi or ascomycetes, constitute the largest taxonomic group within the Eumycota. 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. 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).

 

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, human commensal species of the genus Malassezia, and the opportunistic human pathogen, Cryptococcus neoformans.

 

Fungus-like organisms

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 an ambiregnal, duplicated taxonomy.

 

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 take in 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.

 

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.

 

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. Alternatively, Rozella can be classified as a basal fungal group.

 

The nucleariids may be the next sister group to the eumycete clade, and as such could be included in an expanded fungal kingdom. Many Actinomycetales (Actinomycetota), a group with many filamentous bacteria, were also long believed to be fungi.

 

Ecology

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 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.

 

Symbiosis

Many fungi have important symbiotic relationships with organisms from most if not all kingdoms. 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.

 

With plants

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.

 

A microscopic view of blue-stained cells, some with dark wavy lines in them

The dark filaments are hyphae of the endophytic fungus Epichloë coenophiala in the intercellular spaces of tall fescue leaf sheath tissue

The mycorrhizal symbiosis is ancient, dating back to at least 400 million years. 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. The fungal partners may also mediate plant-to-plant transfer of carbohydrates and other nutrients. Such mycorrhizal communities are called "common mycorrhizal networks". A special case of mycorrhiza is myco-heterotrophy, whereby the plant parasitizes the fungus, obtaining all of its nutrients from its fungal symbiont. Some fungal species inhabit the tissues inside roots, stems, and leaves, in which case they are called endophytes. 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.

 

With algae and cyanobacteria

A green, leaf-like structure attached to a tree, with a pattern of ridges and depression on the bottom surface

The lichen Lobaria pulmonaria, a symbiosis of fungal, algal, and cyanobacterial species

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. Lichens occur in every ecosystem on all continents, play a key role in soil formation and the initiation of biological succession, and are prominent in some extreme environments, including polar, alpine, and semiarid desert regions. They are able to grow on inhospitable surfaces, including bare soil, rocks, tree bark, wood, shells, barnacles and leaves. 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 27% of known fungi—more than 19,400 species—are lichenized. 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.

 

With insects

Many insects also engage in mutualistic relationships with fungi. Several groups of ants cultivate fungi in the order Chaetothyriales for several purposes: as a food source, as a structural component of their nests, and as a part of an ant/plant symbiosis in the domatia (tiny chambers in plants that house arthropods). Ambrosia beetles cultivate various species of fungi in the bark of trees that they infest. 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. 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. Termites on the African savannah are also known to cultivate fungi, 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. Fungi growing in dead wood are essential for xylophagous insects (e.g. woodboring beetles). They deliver nutrients needed by xylophages to nutritionally scarce dead wood. Thanks to this nutritional enrichment the larvae of the woodboring insect is able to grow and develop to adulthood. 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.

 

A thin brown stick positioned horizontally with roughly two dozen clustered orange-red leaves originating from a single point in the middle of the stick. These orange leaves are three to four times larger than the few other green leaves growing out of the stick, and are covered on the lower leaf surface with hundreds of tiny bumps. The background shows the green leaves and branches of neighboring shrubs.

The plant pathogen Puccinia magellanicum (calafate rust) causes the defect known as witch's broom, seen here on a barberry shrub in Chile.

 

Gram stain of Candida albicans from a vaginal swab from a woman with candidiasis, showing hyphae, and chlamydospores, which are 2–4 µm in diameter.

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, tree pathogens such as Ophiostoma ulmi and Ophiostoma novo-ulmi causing Dutch elm disease, Cryphonectria parasitica responsible for chestnut blight, and Phymatotrichopsis omnivora causing Texas Root Rot, and plant pathogens in the genera Fusarium, Ustilago, Alternaria, and Cochliobolus. 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. Many fungi that are plant pathogens, such as Magnaporthe oryzae, can switch from being biotrophic (parasitic on living plants) to being necrotrophic (feeding on the dead tissues of plants they have killed). This same principle is applied to fungi-feeding parasites, including Asterotremella albida, which feeds on the fruit bodies of other fungi both while they are living and after they are dead.

 

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, Histoplasma, and Pneumocystis. 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. Fungal spores are also a cause of allergies, and fungi from different taxonomic groups can evoke allergic reactions.

 

As targets of mycoparasites

Organisms that parasitize fungi are known as mycoparasitic organisms. About 300 species of fungi and fungus-like organisms, belonging to 13 classes and 113 genera, are used as biocontrol agents against plant fungal diseases. Fungi can also act as mycoparasites or antagonists of other fungi, such as Hypomyces chrysospermus, which grows on bolete mushrooms. Fungi can also become the target of infection by mycoviruses.

 

Communication

Main article: Mycorrhizal networks

There appears to be electrical communication between fungi in word-like components according to spiking characteristics.

 

Possible impact on climate

According to a study published in the academic journal Current Biology, fungi can soak from the atmosphere around 36% of global fossil fuel greenhouse gas emissions.

 

Mycotoxins

(6aR,9R)-N-((2R,5S,10aS,10bS)-5-benzyl-10b-hydroxy-2-methyl-3,6-dioxooctahydro-2H-oxazolo[3,2-a] pyrrolo[2,1-c]pyrazin-2-yl)-7-methyl-4,6,6a,7,8,9-hexahydroindolo[4,3-fg] quinoline-9-carboxamide

Ergotamine, a major mycotoxin produced by Claviceps species, which if ingested can cause gangrene, convulsions, and hallucinations

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. 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.

 

Mycotoxins are secondary metabolites (or natural products), and research has established the existence of biochemical pathways solely for the purpose of producing mycotoxins and other natural products in fungi. Mycotoxins may provide fitness benefits in terms of physiological adaptation, competition with other microbes and fungi, and protection from consumption (fungivory). Many fungal secondary metabolites (or derivatives) are used medically, as described under Human use below.

 

Pathogenic mechanisms

Ustilago maydis is a pathogenic plant fungus that causes smut disease in maize and teosinte. Plants have evolved efficient defense systems against pathogenic microbes such as U. maydis. A rapid defense reaction after pathogen attack is the oxidative burst where the plant produces reactive oxygen species 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. Furthermore, U. maydis has a well-established recombinational DNA repair system which acts during mitosis and meiosis. The system may assist the pathogen in surviving DNA damage arising from the host plant's oxidative defensive response to infection.

 

Cryptococcus neoformans is an encapsulated yeast that can live in both plants and animals. C. neoformans usually infects the lungs, where it is phagocytosed by alveolar macrophages. 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. 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. 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.

 

Human use

See also: Human interactions with fungi

Microscopic view of five spherical structures; one of the spheres is considerably smaller than the rest and attached to one of the larger spheres

Saccharomyces cerevisiae cells shown with DIC microscopy

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. Methods have been developed for genetic engineering of fungi, enabling metabolic engineering of fungal species. For example, genetic modification of yeast species—which are easy to grow at fast rates in large fermentation vessels—has opened up ways of pharmaceutical production that are potentially more efficient than production by the original source organisms. Fungi-based industries are sometimes considered to be a major part of a growing bioeconomy, with applications under research and development including use for textiles, meat substitution and general fungal biotechnology.

 

Therapeutic uses

Modern chemotherapeutics

Many species produce metabolites that are major sources of pharmacologically active drugs.

 

Antibiotics

Particularly important are the antibiotics, including the 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. 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. 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.

 

Other

Other drugs produced by fungi include griseofulvin isolated from Penicillium griseofulvum, used to treat fungal infections, 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. Psilocybin from fungi is investigated for therapeutic use and appears to cause global increases in brain network integration. Fungi produce compounds that inhibit viruses and cancer cells. 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. In Europe and Japan, polysaccharide-K (brand name Krestin), a chemical derived from Trametes versicolor, is an approved adjuvant for cancer therapy.

 

Traditional medicine

Upper surface view of a kidney-shaped fungus, brownish-red with a lighter yellow-brown margin, and a somewhat varnished or shiny appearance

Two dried yellow-orange caterpillars, one with a curly grayish fungus growing out of one of its ends. The grayish fungus is roughly equal to or slightly greater in length than the caterpillar, and tapers in thickness to a narrow end.

The fungi Ganoderma lucidum (left) and Ophiocordyceps sinensis (right) are used in traditional medicine practices

Certain mushrooms are used as supposed therapeutics in folk medicine practices, such as traditional Chinese medicine. Mushrooms with a history of such use include Agaricus subrufescens, Ganoderma lucidum, and Ophiocordyceps sinensis.

 

Cultured foods

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. Yeast species of the genus Saccharomyces are also used to produce alcoholic beverages through fermentation. Shoyu koji mold (Aspergillus oryzae) is an essential ingredient in brewing Shoyu (soy sauce) and sake, and the preparation of miso while Rhizopus species are used for making tempeh. 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. Quorn, a meat substitute, is made from Fusarium venenatum.

Near the Undara Lava Tubes, Queensland, Australia.

 

Query - Clathrus? See the below, from Wikipedia.

 

Clathrus ruber is a saprobic species of fungus in the family Phallaceae. It is commonly known as the latticed stinkhorn, the basket stinkhorn, or the red cage, alluding to the fruiting bodies that are shaped somewhat like a round or oval ball with interlaced or latticed branches. Covered with a slime layer on the inner surfaces, this inedible species has a fetid odor, described by some as rotting meat, that attracts flies and other insects to help disperse the spores.

 

Description

Prior to the opening of the volva, the fruiting body is egg-shaped with a gelatinous interior, and white to grayish in color. After opening, it is a red or orange receptacle consisting of a spongy network with meshes of unequal size. A considerable variation in height has been noted for this species, ranging from 8–20 cm (3.1–7.9 in) tall. The dark, foul smelling gleba covers the inner surface of the receptacle and the basal portion of the receptacle is surrounded by a white volva with a central mycelial cord. The spores are elongated, smooth, and have dimensions of 5–6 x 1.7–2 µm.

 

This species may be distinguished from the closely related tropical species Clathrus crispin by the absence of corrugated rims which surround each mesh of the C. crispin sporocarp.

 

Habitat

This fungus grows alone or clustered together near woody debris, in lawns, gardens, and cultivated soil.

 

Edibility

Although edibility for C. ruber has not been officially documented, its foul smell would dissuade most individuals from consuming it.

 

Dr. F. Peyre Porcher, of Charleston, South Carolina, United States (1854) provided an early account of poisoning by this species:

 

"A young person having eaten a bit of it, after six hours suffered from a painful tension of the lower stomach, and violent convulsions. He lost the use of his speech, and fell into a state of stupor, which lasted for forty-eight hours. After taking an emetic he threw up a fragment of the mushroom, with two worms, and mucus, tinged with blood. Milk, oil, and emollient fomentations, were then employed with success."

 

Taxonomy

This species has often been referred to by American authors as C. cancellatus Linnaeus, based on the American Code of Botanical Nomenclature, in which the starting point was Linnaeus, Species plantarum, 1753. According to the International Code for Botanical Nomenclature, the starting date for the nomenclature of the Gasteromycetes is 1801, the publication date of Christian Hendrik Persoon's Synopsis methodica fungorum (1801). Thus, the correct specific epithet for this species is Clathrus ruber.

 

Distribution

C. ruber occurs in the United States, Australasia, southern eastern and western Europe, Canary Islands, Caribbean Islands, the Azores, Canada, Mexico, Japan, France, Poland, and the United Kingdom (Isle of Wight, Cornwall, Devon, and Ireland. Within the United States, C. ruber has been reported from California, Florida, Georgia, Virginia, North Carolina and New York.

 

C. ruber is listed in the Red data book of Ukraine.

 

Biochemistry

Like other stinkhorn mushrooms, C. ruber bioaccumulates the element manganese. It has been postulated that this element plays a role in the enzymatic breakdown of the gleba with simultaneous formation of odorous compounds. A multielement analysis of the gelatinous outer layer, the embryonic receptacle and the gleba were undertaken, and the gelatinous layer proved richest in potassium, calcium, manganese and iron. Calcium stabilizes the polysaccharide gel protecting the embryonic carpophore (the stalk of the sporocarp) from drying out during the growth of the egg. The elevated concentrations of the other elements (compared to those in the developing carpophore) suggest that the gelatinous layer has a placenta-like function.

 

Pigments responsible for the orange to red colors of the mature fruiting bodies have been identified as carotenes, predominantly lycopene and beta carotene.

by Michael Christian and Dallas Swindle

Berkeley, CA

 

ePod (Eidolon Panspermia Ostentatia Duodenum) is a 24 foot-tall steel voronoi-inspired open-cell mycelial ovoid structure representing a possible matrix manifold of the panspermia transportation vesicle designed to inspire a profound sense of apophenia. And it has pretty lights.

 

connect.clickandpledge.com/Organization/blackrockarts/cam...

From MATCHMAKER

 

LATIN NAME(S) Marasmius plicatulus Peck Bull. Torrey Bot. Club 24: 124. 1897

ENGLISH NAME(S) pleated mushroom, velvet-cap Marasmius

NOTES distinctive features are velvety wine-red to brown cap, distant gills, and long shining reddish-black brittle stem; collections examined from BC, WA, OR, ID, CA, (Redhead)

CAP 1-4(5)cm across, obtusely conic to bell-shaped, often expanding to convex or flat or with uplifted margin when old; bay-brown to reddish brown, brown, wine-red, or maroon; "dry, with a velvety or frosted appearance when fresh", furrowed or wrinkled when old or dried, (Arora), 1.0-3.5(5.5)cm across, conic or bell-shaped when young, broadly conic or convex when old; dark reddish brown when young, reddish brown when old, margin reddish brown when young, becoming brown, light brown or brownish orange when old; somewhat velvety, dull, opaque, disc smooth or subrugulose [somewhat wrinkled], margin even, becoming striate, (Desjardin), may be umbonate

FLESH thin, pliant, (Arora); 0.05-0.1cm thick; buff, (Desjardin)

GILLS "adnate to nearly free, well-spaced, broad"; "white to buff to pinkish or tinged cap color", (Arora), "adnexed, distant, broad", 1-2 series of subgills; buff or yellowish white, (Desjardin)

STEM 5-13cm x 0.15-0.3cm, equal, usually long and thin, tough but brittle (dig up rather than pluck); reddish black to deep chestnut in lower part, often paler (pinkish or sometimes pallid) in upper part; smooth, polished, base often with whitish mycelium, (Arora), 6.0-9.5cm x 0.2-0.3cm, equal, round in cross-section, fibrous but easily broken, not insititious; top grayish red or brown, base dark brown to black; shiny, bald in upper part, "base covered with white, strigose mycelial hairs", "basal mycelium forming an extensive mycelial mat", (Desjardin), may be curved

VEIL absent

ODOR mild (Arora, Desjardin)

TASTE mild (Desjardin)

EDIBILITY too tough and thin (Arora)

HABITAT widely scattered to gregarious "in humus under trees and shrubs" (Arora), gregarious in humus or grassy areas under oak or various conifers, (Desjardin), July to April (Miller), spring, summer, fall, winter

SPORE DEPOSIT white (Arora)

MICROSCOPIC spores 11-15 x 5-6.5 microns, elliptic, smooth, inamyloid (Arora), spores 12.3-16.2(16.6) x 4.8-6.3 microns, almond-shaped or subfusiform (somewhat spindle-shaped); basidia 4-spored, 38-50 x 7-10 microns, clavate or spheropedunculate; pleurocystidia absent, cheilocystidia "abundant in young specimens, often scarce in older specimens", 15-24 x 3.6-6 microns, "cylindric or clavate, diverticulate", diverticula numerous, 1.8-10.8 x 0.6-2.1 microns, apical, rod-like or conic, often branched, basal part of cells colorless, thin-walled, diverticula "solid, yellow or pale ochraceous, dextrinoid", (Desjardin)

NAME ORIGIN means 'folded'

SIMILAR

SOURCES Desjardin(1), Arora(1)*, Lincoff(2)*, Miller(14)*, Sept(1)*, Redhead(6)

FAMILY Marasmiaceae of Order Agaricales

Mycelium, OOAK Inamorata art doll featuring Shani head sculpt in Ice resin.

 

I originally made Mycilium for the Hitogata Ten art doll exhibition in Tokyo. Since tattoos are a bit of a taboo in Japan, and I didn't want to risk upsetting the gallery, I didn't fulfil my original vision for he doll at the time. However, when this doll sold after the exhibition, I pitched the idea of painting her entire body with a mycelial network that has little bugs scuttling throughout it. I was so happy that the new owner loved the idea and I got to finish the concept. Her white mulberry silk hard cap wig with branching out braids repeats the branching patterns of a mycelial network.

 

The next preorder for Inamorata dolls is on April 22nd and will include this translucent Ice resin tone.

Mycilium, OOAK Inamorata doll with Shani head sculpt in Ice resin. Mycilium is being displayed in the Hitogata Ten art doll exhibition in Tokyo.

 

She is a nude OOAK with a white mulberry silk wig with branching out braids and amazing 3D printed wings by @lbxcouture (Thank you for the wings LBX!).

 

I was playing with the idea that fairies were related to mushrooms and that she is symbiotically reaching out and connecting to the environment around her using mycelial networks .

 

There is a waiting list for the exhibition dolls. Email me with "waiting list" as topic to get a chance to buy her if she doesn't sell at the exhibition.

Mycilium, OOAK Inamorata doll with Shani head sculpt in Ice resin. Mycilium is being displayed in the Hitogata Ten art doll exhibition in Tokyo.

 

She is a nude OOAK with a white mulberry silk wig with branching out braids and amazing 3D printed wings by @lbxcouture (Thank you for the wings LBX!).

 

I was playing with the idea that fairies were related to mushrooms and that she is symbiotically reaching out and connecting to the environment around her using mycelial networks .

 

There is a waiting list for the exhibition dolls. Email me with "waiting list" as topic to get a chance to buy her if she doesn't sell at the exhibition.

Mycilium, OOAK Inamorata doll with Shani head sculpt in Ice resin. Mycilium is being displayed in the Hitogata Ten art doll exhibition in Tokyo.

 

She is a nude OOAK with a white mulberry silk wig with branching out braids and amazing 3D printed wings by @lbxcouture (Thank you for the wings LBX!).

 

I was playing with the idea that fairies were related to mushrooms and that she is symbiotically reaching out and connecting to the environment around her using mycelial networks .

 

There is a waiting list for the exhibition dolls. Email me with "waiting list" as topic to get a chance to buy her if she doesn't sell at the exhibition.

Mycilium, OOAK Inamorata doll with Shani head sculpt in Ice resin. Mycilium is being displayed in the Hitogata Ten art doll exhibition in Tokyo.

 

She is a nude OOAK with a white mulberry silk wig with branching out braids and amazing 3D printed wings by @lbxcouture (Thank you for the wings LBX!).

 

I was playing with the idea that fairies were related to mushrooms and that she is symbiotically reaching out and connecting to the environment around her using mycelial networks .

 

There is a waiting list for the exhibition dolls. Email me with "waiting list" as topic to get a chance to buy her if she doesn't sell at the exhibition.

Original: edible mushroom fungicides - "Wormwood."

Author: flat spring / bacteria shelterbelt

 

Hear each mushroom export business in China, Japan and clamored for the Chinese mushrooms in Japan the most stringent set of "positive list system." The provisions of the mushrooms produced more than 600 technical indicators set, forcing the domestic mushroom export business costs, and make mushroom export business in China in 2010, exports to Japan decreased Liu Cheng.

Japanese products, the reason known, which is their quality system, environmental protection and low-carbon economy, responsible for product recovery system associated.

Japanese emphasis on the natural food of the entrance, which is ranked the world's most longevity in Japan for many years the main cause of national forefront.

China is the largest producer of mushrooms, but mushroom production in China is still generally stay in the extensive cultivation of the state environmental management techniques, which is the global advocate of organic, green agriculture food production system is incompatible.

Notes that the mushroom export business, if you want to occupy the high rate of the Japanese market, have to increase the mushroom production process on the strength of the link, eliminating the use of high residue of pesticides and chemical fungicides, taking advantage of the initiative to adjust and improve, mushroom industry, the production system of high-tech standards, it is better than more constructive than complain year after year, blacksmith must own hard thing!

 

1, the wormwood and the chemical composition of the medical value of

Leaves (Argy Wormwood Leaf)

Scientific name Folium Arte-misiae Argyi

The Compositae (Artemisia argyiLevl.etVant)

Is a perennial herb. Mainly distributed in China's Heilongjiang, Jilin, Liaoning, Hebei, Shandong, Anhui, Jiangsu, Zhejiang, Guangdong, Guangxi, Jiangxi, Hunan, Hubei, Sichuan, Guizhou, Yunnan, Shaanxi, Gansu and other regions.

According to the researchers analyzed the main component of wormwood:

Absinthe alcohol (25.87%), eucalyptol (20.37%), 1R-α-pinene (3.39%), Eucalyptus alcohol (2.73%), 3,3,6 - trimethyl--1,5 - heptadiene -4-- alcohol (2.34%), caryophyllene (2.03%), camphene alcohol (1.43%), etc. In addition Isobornyl formic acid (1.34%), key Eucalyptus oil-ol (0.72%), Se Seer ene (0.56%), naphthalene embedded in pentane (0.54%), Tan Purple triene (0.53%), and for the first time detected in the leaves. To wild leaves smoked, for a variety of pathogenic bacteria also have the role. Ai decoction of wild flooding agents in vitro on a variety of pathogenic fungi have a certain extent.

 

1. ) Anti-fungal effect of wormwood

Clinically, found in the wards with Ai smoked, some patients died of cold can be self-healing, Ai smoked for local herpes zoster, purulent skin infections, rashes, etc. have good therapeutic effect.

Then Ai smoke on the antibacterial effects of various bacterial and thermal stimulation is it?

Ai burning smoke, which contains the volatile oil will evaporate with the smoke, tests showed that the leaves volatile oil of common pathogens such as pneumococcus, white, and Staphylococcus aureus, Influenza A and B streptococci, Neisseria, E. coli , typhoid and paratyphoid bacteria, Shigella flexneri, Haemophilus influenzae, Proteus, etc. have antibacterial, minimum inhibitory concentration of 2 × 10-3 ~ 4 × 10-3 ml.

 

2. ) Antiviral activity of wormwood

Experiments show that, alone with the leaves smoked for adenovirus, rhinovirus, influenza virus and parainfluenza virus inhibition. The leaves extract diluted 1:10 with saline has some inhibitory effects of the virus.

3. ) Wormwood in the treatment of asthma on lung mushroom antitussive effect

A large number of pharmacological experiments show that leaves volatile oil spray administration of oral or have better asthma, antitussive effect, particularly among the most significant role in asthma. Clinically useful leaves volatile oil volatile oil spray or wet leaves inhalation treatment of asthma, allergies to the mushroom spores have a good effect. 2, the wormwood on the application in food

Annual Ching Ming Festival, most people used to collect fresh Ai shoots, put glutinous rice dough, mixed with pounded dough, then steamed cake made seasonal cakes Ai consumption.

   

According to testing, hectogram stem leaves with fresh water 85 g, protein 3.1 g, fat 0.6 grams carbohydrate, 7 g, 2.18 mg of calcium, phosphorus 66 mg, iron 7.4 mg, 2.19 mg of carotene, vitamin B20.24 mg, 1.4 mg niacin, vitamin C28 mg also contain volatile oils, alkaloids, etc.

 

Level of sweet wormwood, with rheumatism, dampness cloud, Huatanzhike effect. "Compendium of Materia Medica." Qingming Festival, edible cake Ai, a "transfer of Qi, Chu Tan stopped the leaks, the pressure when the gas to heat cough," "cure cold cough and sputum, in addition to the lungs cold, soared lung." Modern pharmacological studies, the tender shoots of wormwood ingredient used as ingredients, dishes, also has expanded the local swelling blood vessels, lower blood pressure, treatment of peptic ulcer, antitussive, analgesic and other therapeutic effects.

  

3, the wormwood on the application in mushroom cultivation 1. ) Wormwood Powder

Fresh wormwood is divided into two kinds, one is the big leaves, prolific in the provinces north of the Yangtze River, wide and thick leaves, many and dense dorsal hair, leaves a positive Secheng green, the dorsal hair hard, such Wormwood do more for the Sunburn Sunburn points on the human body moxa treatment processing; the other main production in the country, also known as wild wormwood, chrysanthemum leaves showing leaf-like, Ai Hong volatile than large-leaved species, but its chemical composition and big leaves are the same. Moreover, several provinces across the country regions, more likely to collect. Fresh wormwood in the spring and summer each year, while no buds, leafy when collecting, storing dried. Before use and then crushed ingredients. 2. ) Materials and methods selected oyster strains forest technicians do this comparative test 7 strains. Each recipe each doing 100 mushroom treatment. Moisture content of 60% medium, the medium before sterilization PH7.5. Each mushroom weighing about 600 g of dry material, bacteria bar is 22 cm. Culture medium by conventional spices recipe, adjusting the water, bagging, sterilization, inoculation chamber inoculation, incubation temperature with the temperature 24 degrees. (1) Add 1.5% of wormwood powder formulations: cotton seed shell 79.5%, 1% lime powder, gypsum powder, 1.5%, 15% wheat bran, corn meal 1%, 0.5% potassium dihydrogen phosphate (2) add Wormwood powder 2% of the formula: 79% cotton seed shell, lime powder, 1%, gypsum powder, 1.5%, 15% wheat bran, corn meal 1%, 0.5% potassium dihydrogen phosphate (3) Add 3% of wormwood powder formula: 78% cotton seed shell, lime powder, 1%, gypsum powder 1.5%, 15% wheat bran, corn meal 1%, 0.5% potassium dihydrogen phosphate (4) control group, without adding wormwood powder formulations: cotton seed Shell 80%, 2% lime powder, gypsum powder, 1.5%, 15% wheat bran, corn meal 1%, potassium dihydrogen phosphate 0.5% 3. ) Results and analysis of different combinations of mycelial growth of wormwood, mushroom and fresh mushroom production number pollution (unit: mm / d, Kg, a, g) formula

(1)

(2)

(3)

(4)

Hyphal length of the day speed / average 14.89 14.65 7.9 15.05

  

The number of pollution mushroom 5 2.3 1.5 8.7

Average mushroom yield 365 427.5 346.3 400.2

Biotransformation rate of 60.8% 72% 58% 67%

 

Add wormwood combination of different combinations were made with no added bacteria speed comparison:

3) -1 by the above test data analysis: formula (4) the growth of mycelium on the fastest speed, but (1), (2) a combination of daily growth rate was not significant; (3) hair growth rate of bacteria on the most slow, indicating that the proportion of added wormwood if more than 2% to 2.5%, there will be inhibition of the anti-normal growth rate of mycelium.

3) -2 trial mushroom contamination by the number of the above analysis: the control combination (4) appears the highest number of pollution, (3) minimum, (2) times the pollution is low, (1) is the second highest.

3) -3 Summary and discussion

By the average of the pilot mushroom production data analyzed: a combination of (2) the highest, the average conversion rate reached 72%, combined with other control has a clear gap, which shows medium containing 2% wormwood powder later, not only reduces the rate of bacterial contamination of stick, some compounds in wormwood, also has a hidden role in increasing production.

First country, please indicate the bacteria shelterbelt: www.chjunlin.com/

 

Mycelium, OOAK Inamorata art doll featuring Shani head sculpt in Ice resin.

 

I originally made Mycilium for the Hitogata Ten art doll exhibition in Tokyo. Since tattoos are a bit of a taboo in Japan, and I didn't want to risk upsetting the gallery, I didn't fulfil my original vision for he doll at the time. However, when this doll sold after the exhibition, I pitched the idea of painting her entire body with a mycelial network that has little bugs scuttling throughout it. I was so happy that the new owner loved the idea and I got to finish the concept. Her white mulberry silk hard cap wig with branching out braids repeats the branching patterns of a mycelial network.

 

The next preorder for Inamorata dolls is on April 22nd and will include this translucent Ice resin tone.

A biomechanical terraformer, spreading the reach of the Verdant Protocol into the unstable wilderness of the Shattersea.

 

Function & Purpose:

 

Bloomcaller units are autonomous biological terraforming constructs, designed to expand the Verdant Protocol’s influence by seeding, cultivating, and adapting plant life to harsh or unstable conditions. Unlike the more aggressive Harvesters, which break down and repurpose organic material, Bloomcallers focus on growth, regeneration, and controlled mutation, turning barren landscapes into thriving biomes.

 

They are not combat units, but they are far from defenceless. Their ability to rapidly alter the local environment allows them to make terrain inhospitable to threats, deploying toxic spores, root entanglements, and hallucinogenic pollen clouds as a deterrent. Their primary mission, however, is to spread, adapt, and sustain the Verdant ecosystem, ensuring the continued evolution of organic life within the Shattersea.

 

⸻

 

Physical Characteristics & Capabilities:

 

Mycelial Canopy (Floral Crown)

•The cluster of plant life atop the Bloomcaller is more than just decorative—it acts as a real-time environmental scanner and genetic repository.

•This organic canopy samples the atmosphere, temperature, and soil, adjusting its seeding process accordingly.

•Different Bloomcaller units grow unique flora based on their region of deployment, making them highly adaptable.

•Some act as pollinators, while others introduce hardy, fast-spreading fungal networks that help stabilize terrain.

Spore Chambers & Bioluminescent Vats

•The large, translucent sacs on its body contain specialized spores, bio-serum, and genetically engineered seedlings.

•Depending on the situation, the Bloomcaller can release:

•Hardy Flora Spores: Creates drought-resistant plant growth, stabilizing soil in shifting or crumbling environments.

•Fungal Mycelium Webs: Spreads Verdant neural networks underground, linking plants and extending the Protocol’s influence.

•Hallucinogenic or Defensive Spores: Releases clouds of spores that can disorient intruders or deter hostile wildlife.

•Adaptive Hybrid Seeds: Capable of mutating to better withstand the extreme conditions of the Shattersea.

Hydro-Root Injection Systems

•The tentacle-like appendages and clawed manipulators are designed for direct interaction with terrain.

•Some function as root injectors, implanting fast-growing vegetation deep into the ground.

•Others serve as nutrient distributors, helping spread organic material from decayed sources to sustain new life.

Tripedal Stability & Traversal Adaptability

•The Bloomcaller’s three-legged stance allows for stable movement across unstable ground, including shifting sand, deep mud, and debris-laden terrain.

 

Unlike more predatory Verdant units, it has no offensive weaponry, relying on environmental manipulation and biome-defense mechanisms to avoid threats.

•When under extreme duress, some Bloomcallers self-terminate, bursting into a wave of spores that create an immediate, fast-growing fungal bloom, obscuring their retreat and leaving behind a Verdant “seed” that will later develop into a new outpost.

 

⸻

 

Role in the Shattersea:

•Terraforming & Expansion: Deploys in unstable regions, planting flora and fungal networks that anchor and stabilize terrain.

•Reclamation & Healing: Can be sent to dead zones, restoring them to sustainable ecosystems over time.

•Ecosystem Adaptation: Introduces new plant species tailored to the environment, ensuring biodiversity.

•Tactical Denial & Terrain Manipulation: Uses defensive plant growth to create barriers, obscure pathways, or redirect intruders away from Verdant-controlled zones.

 

⸻

 

Personality & Behaviour:

•Unlike Harvesters, which are more mechanical and purpose-driven, Bloomcallers exhibit a form of curious, slow intelligence.

•They move deliberately and methodically, scanning and adjusting their environment before acting.

•Some Drift Runners claim that Bloomcallers watch them—not with hostility, but with a kind of detached interest, as if measuring whether the land should welcome them or resist them.

•They are not aggressive unless directly threatened, preferring to retreat and let the landscape itself become the deterrent.

 

⸻

 

Interactions with Humans & Drift Runners

•Hostile Factions: Many human groups—especially Bastion forces and industrial factions—see Bloomcallers as a threat to expansion efforts, as they can overtake infrastructure and terraform industrial sites into Verdant territory.

•Tromas & The Howling Hoser: Given Tromas’ collaboration with the Verdant Protocol, he has encountered and

even guided a Bloomcaller unit before. He has a designated safe route through Bloomcaller-controlled zones, allowing for limited trade of bio-fuels and organic materials.

Mycelium, OOAK Inamorata art doll featuring Shani head sculpt in Ice resin.

 

I originally made Mycilium for the Hitogata Ten art doll exhibition in Tokyo. Since tattoos are a bit of a taboo in Japan, and I didn't want to risk upsetting the gallery, I didn't fulfil my original vision for he doll at the time. However, when this doll sold after the exhibition, I pitched the idea of painting her entire body with a mycelial network that has little bugs scuttling throughout it. I was so happy that the new owner loved the idea and I got to finish the concept. Her white mulberry silk hard cap wig with branching out braids repeats the branching patterns of a mycelial network.

 

The next preorder for Inamorata dolls is on April 22nd and will include this translucent Ice resin tone.

view this photo large on black

 

see more interesting photo's from me here:

flickeflu.com/photos/77411963@N07/interesting

 

Armillaria solidipes (formerly Armillaria ostoyae), the honey mushroom, is the most common variant in the western U.S. of the group of species that all used to share the name Armillaria mellea. Armillaria solidipes is quite common on both hardwood and conifer wood in forests west of the Cascade crest. The mycelium attacks the sapwood and is able to travel great distances under the bark or between trees in the form of black rhizomorphs ("shoestrings").

 

In most areas of North America, Armillaria solidipes can be separated from other species by its physical features. Its brown colors, fairly prominent scales featured on its cap, and the well-developed ring on its stem sets it apart from any Armillaria.

 

It is known to be one of the largest living organisms, where scientists have estimated a single specimen found in Malheur National Forest in Oregon to have been growing for some 2,400 years, covering 3.4 square miles (8.4 km^2) and colloquially named the "Humongous Fungus." Armillaria solidipes grows and spreads primarily underground and the bulk of the organism lies in the ground, out of sight. Therefore, the organism is not visible to anyone viewing from the surface. It is only in the autumn when this organism will bloom “honey mushrooms”, visible evidence of the organism lying beneath. Low competition for land and nutrients have allowed this organism to grow so huge; it possibly covers more geographical area than any other living organism.

 

This fungus, like most parasitic fungi, reproduces sexually. The fungi begin their life as spores, released into the environment by a mature mushroom. Armillaria solidipes has a white spore print. There are two types of mating types for spores (not male and female but similar in effect). The spores can be dispersed by environment factors such as wind or they can be redeposited by an animal. Once the spores are in a resting state, the single spore must come in contact with a spore of an opposite mating type and of the same species. If the single spore isolates are from different species, the colonies will not fuse together and they will remain separate. When two isolates of the same species but different mating types fuse together, they soon form coalesced colonies which become dark brown and flat. With this particular fungus it will produce mycelial cords also known as rhizomorphs. These rhizomorphs allow the fungus to obtain nutrients from long distances away. These are also the main factors to its pathogenicity. As the fruiting body continues to grow and obtaining nutrients, it forms into a mature mushroom. Armillaria solidipes in particular grows a wide and thin sheet-like plates radiating from the stem which is known as its gills. The gills hold the spores of a mature mushroom. This is stained white when seen as a spore print. Once spore formation is complete, this signifies a mature mushroom and now is able to spread its spores to start a new generation.

 

The disease is of particular interest to forest managers, as the species is highly pathogenic to a number of commercial softwoods, notably Douglas-fir (Pseudotsuga menziesii), true firs (Abies spp.) and Western Hemlock (Tsuga heterophylla). A commonly prescribed treatment is the clear cutting of an infected stand followed by planting with more resistant species such as Western redcedar (Thuja plicata) or deciduous seedlings. Armillaria can remain viable in stumps for 50 years.

 

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Armillaria ostoyae, Sombere honingzwam.

 

Middelbruine tot geelbruine, vleeskleurige hoed en iets lichtere steel. Hoed met donkerder, afwisbare schubjes. De steel heeft een forse ring, met beneden de ring ook schubjes op de steel. De rand van de hoed is in vochtige toestand doorschijnend gestreept. Groeit meestal in bundels. Kleur van de sporen: wit tot crème. Hoogte: 6-15 cm, breedte: 3-10 cm.

 

Kan van september tot november gevonden worden op de stam, de basis en de wortels van bomen en op stronken van bomen. Meestal op min of meer zure zandgronden, komt algemeen voor.

 

Honingzwammen verspreiden zich niet alleen via hun sporen, maar ook via lange zwarte draden, rhizomorfen. Deze op veters lijkende zwarte draden kunnen onder de bast van aangetaste bomen gevonden worden. Voor de honingzwammen in het algemeen geldt, dat het parasieten zijn, die een sterke vorm van witrot veroorzaken, die uiteindelijk leidt tot het afsterven van de gastheer.

 

In april 2003 werd in het Malheur National Forest in de Amerikaanse staat Oregon een sombere honingzwam ontdekt van naar schatting 2400 jaar oud met een ondergrondse mycelium omvang van 8,9 km². Daarmee is deze schimmel het grootste organisme ter wereld. Ook in Zwitserland in het Nationaal Park in de streek Engadin komt deze schimmel met een grote omvang voor. Hier is de schimmel ongeveer duizend jaar oud en ongeveer 800 meter lang en 500 meter breed.

 

Mycelium, OOAK Inamorata art doll featuring Shani head sculpt in Ice resin.

 

I originally made Mycilium for the Hitogata Ten art doll exhibition in Tokyo. Since tattoos are a bit of a taboo in Japan, and I didn't want to risk upsetting the gallery, I didn't fulfil my original vision for he doll at the time. However, when this doll sold after the exhibition, I pitched the idea of painting her entire body with a mycelial network that has little bugs scuttling throughout it. I was so happy that the new owner loved the idea and I got to finish the concept. Her white mulberry silk hard cap wig with branching out braids repeats the branching patterns of a mycelial network.

 

The next preorder for Inamorata dolls is on April 22nd and will include this translucent Ice resin tone.

Dutch nature -

 

Bij sommige van deze gevallen reuzen zie je zelfs de groeidraden van paddenstoelen. Het zijn vaak parasitaire paddenstoelen die jaren het hout hebben opgegeten van de boom. Met hun myceliumdraden ( zwamvlok ) vreten ze zich een weg door het hout. In de loop van tientallen jaren consumeert de zwam met zijn groeidraden een groot gedeelte van het binnenwerk van de boom op. Het gevolg is dat de boom verzwakt op bepaalde plekken. Op een gegeven moment is een herfstige rukwind voldoende om de boom te vellen. De dood ten gevolge..

....................................................................................................................

 

In some of these fallen giants you can even see the growth rates of mushrooms. They are often parasitic mushrooms that have eaten the wood for years. With their mycelial wires (fungal flake) they feed a way through the wood. Over the course of decades, the fungus consumes a large part of the tree's interior with its growth wires. As a result, the tree weakens in certain places. At some point, an autumn blow wind is enough to sheet the tree. Death as a result.

Megacollybia platyphylla (Pers.) Kotl. & Pouzar, syn.: Collybia platyphylla (Pers.) P. Kumm., Clitocybula platyphylla (Pers.) E. Ludw., Oudemansiella platyphylla (Pers.) Moser, Tricholomopsis platyphya and many others

Platterful mushroom, Whitelaced Shank, DE: Gewöhnliches Breitblatt, Breitblätriger Rübling

Slo.: širokolistna velekorenovka

 

Dat.: July 26. 2017

Lat.: 46.35959 Long.: 13.70122

Code: Bot_1077/2017_DSC8488

 

Habitat: mixed wood, Fagus sylvatica and Picea abies dominant trees with some Ostrya carpinifolia and Fraxinus ornus; slightly inclined mountain slope, SE aspect; old colluvial, calcareous ground, in shade, dry and relatively warm place; partly protected from direct rain by tree canopies; average precipitations ~ 3.000 mm/year, average temperature 7-9 deg C, elevation 600 m (1.950), alpine phytogeographical region.

 

Substratum: heavily rotten, moss covered stump of Fagus sylvatica.

 

Place: Lower Trenta valley, between villages Soča and Trenta, right bank of river Soča; near the trail from Trenta 2b to Strgulc farmhouse, Soča 48; East Julian Alps, Posočje, Slovenia EC.

 

Comments: When I noticed this fungus I thought it was a kind of Pluteus (actually Pluteus cervinus (Schaeff.) P. Kumm. looks very similar). However, neither its spore print nor gills showed even a trace of pinkish color. So, it must be almost for sure Megacollybia platyphylla. Its streaked-looking gray-brown cap is typical.

 

Genus Megacollybia was considered monophyletic (that is containing only one specie - Megacollybia platyphylla) not so far ago. Recent DNA sequencing in North America (Ref.3) showed that it can be separated to several distinct species (how many of them scientists do not agree, up to nine worldwide). One of them Megacollybia platyphylla was limited to Europe. Unfortunately newly recognized species in North America cannot practically be separated by macro-morphological traits (particularly not if geographic location is dismissed as a 'character'). They are all variable and all look alike. Even microscopy doesn't help. To determine them in the field one would need 'pocket DNA sequencer' (which doesn't exist). So it is: DNA-defined species do not always show observable differences in physical features.

 

Megacollybia platyphylla is a common mushroom. There are conflicting reports about edibility of it. Some consider it conditionally edible, some weakly poisonous.

 

Description of this find: Growing solitary; heavily eaten by snails; pileus diameter 13 cm, radially streaked, strakes peel off easily; trama thin, gills broad, beige, fragile; stipe 8 cm long, 23 mm in diameter at the base and 19 mm on top, firm, slightly hollow at the center, fibrous, firmly attached to the rotten wood by mycelial strands; smell slightly on sour bread, taste mild, indistinctive; SP faint, beige, oac814.

 

Spores smooth. Dimensions: 7.3 [8 ; 8.3] 9 x 5.9 [6.5 ; 6.8] 7.4 microns; Q = 1.1 [1.2] 1.4; N = 30; C = 95%; Me = 8.1 x 6.7 microns; Qe = 1.2. Olympus CH20, NEA 100x/1.25, magnification 1.000 x, oil; in water, fresh material. AmScope MA500 digital camera.

 

Herbarium: Mycotheca and lichen herbarium (LJU-Li) of Slovenian Forestry Institute, Večna pot 2, Ljubljana, Index Herbariorum LJF

 

Ref.:

(1) R. Phillips, Mushrooms, Macmillan (2006), p 96.

(2) G.J. Krieglsteiner (Hrsg.), Die Grosspilze Baden-Württembergs, Band 3., Ulmer (2001), p 360.

(3) www.mushroomexpert.com/megacollybia.html stanje megacolibija

(4) J. Breitenbach, F. Kraenzlin, Eds., Fungi of Switzerland, Vol.3. Verlag Mykologia (1991), p 246.

(5) L. Hagar, Ottova Encyklopedia Húb, Ottova Nakladatelstvi, Praha (2015) (in Slovakian), p 673.

(6) R. Lueder, Grundkurs Pilzbestimmung, Quelle & Meyer (2008), p 223.

(7) S. Buczacki, Collins Fungi Guide, Collins (2012), p 208.

 

Geastrum quadrifidum (Pers.) Pers., syn.: Geastrum coronatum Scopoli

Rayed Earthstar, DE: Kleiner Nest-Erdstern, Kronen Erdstern

Slo.: četverokraka zvezdica

 

Dat.: Sept. 09. 2014

Lat.: 46.36529 Long.: 13.74988

Code: Bot_835/2014_DSC3951

 

Habitat: mixed wood, Picea abies and Fagus sylvatica dominant, moderately steep, southeast oriented mountain slope, calcareous skeletal ground covered by leaf and needles litter without ground vegetation, under Picea abies, in shade, partly protected from direct rain by tree canopies, average precipitations ~ 3.000 mm/year, average temperature 5-7 deg C, elevation 950 m (3.100 feet), alpine phytogeographical region.

 

Substratum: on moss covering a small Picea abies stump in the last disintegration stage, decomposed to almost soil.

 

Place: Lower Trenta valley, near the trail from Trenta village to Zasavska koča na Prehodavcih mountain cottage, halfway between the village and Planina Lepoč, East Julian Alps, Posočje, Slovenia EC

 

Comments: Genus Geastrum contains very attractive fungi, which almost all are rather uncommon, if not rare. Globose or onion shaped fruit bodies of many start to develop underground. The 'shell' of their fruit bodies consists of four distinct layers (with some exceptions). The outer three form so called exoperidium and the inner one is endoperidium, a 'sack', which contains spores. Exoperidium's outer layer consists of mycelium, the middle layer consists of fibers and the inner one is so called pseudoparenchymal layer. During growth the last one swells and breaks the exoperidium into star like shaped lappets, which curl backward and in this way push the fruit body out of the ground. In some species, like with Geastrum quadrifidum, the outer mycelial layer does not split together with other two layers of the exoperidium but falls off and forms a kind of 'bird's nest' in ground on top of which the fruit body sits. This white mycelial 'nest' can be seen on Fig. 3. The fruit body, when mature, cuts itself almost completely off the mycelium and stands free, like on legs made of exoperidium flaps. Only the far ends of the laps stay in connection with the 'nest'. In this way endoperidium with its 'chimney' (peristom) on top, through which clouds of spores rise like a 'smoke', is positioned as high as possible to facilitate spore spreading by the wind.

 

Geastrum quadrifidum is among the smallest species of about 50 of them worldwide (and ten of them described in Slovenian checklist (Ref.7)). It is a rare find. As its species name suggests it usually has four exoperidium 'legs'. However, sometimes, as in my find, it has five of them.

 

Growing solitary; exoperidium diameter 20 mm, endoperidium diameter 8 mm, its height (without the peristom 'beak') 7 mm; SP and spores on mass dark brown; smell none; taste not tested (too small).

 

Spores coarsely warty. Dimensions excluding warts: 4.4 [5 ; 5.2] 5.7 x 4.1 [4.5 ; 4.7] 5.2 microns; Q = 1 [1.1] 1.2; N = 40; C = 95%; Me = 5.1 x 4.6 microns; Qe = 1.1; number of warts per circumference: AVG = 12.1, SD = 1.2, N = 30. Olympus CH20, NEA 100x/1.25, magnification 1.000 x, oil; AmScope MA500 digital camera.

 

Herbarium: Mycotheca and lichen herbarium (LJU-Li) of Slovenian Forestry Institute, Večna pot 2, Ljubljana, Index Herbariorum LJF

 

Ref.:

(1) G.J. Krieglsteiner (Hrsg.), Die Grosspilze Baden-Württembergs, Band 2., Ulmer (2000), p 116.

(2) S. Buczacki, Collins Fungi Guide, Collins (2012), p 440.

(3) W. Rothmaler, Exkursionsflora von Deutschland, Vol.1, Niedere Pflanzen, Elsevier, 3.Auflage, (1994), p 519.

(4) M. Bon, Parey's Buch der Pilze, Kosmos (2005), p 302.

(5) J. Breitenbach, F. Kraenzlin, Eds., Fungi of Switzerland, Vol.2. Verlag Mykologia(1986), p 382.

(6) R. Phillips, Mushrooms, Macmillan (2006), p 334.

(7) A. Poler, ed., Seznam gliv Slovenije (in Slovene), 2nd Ed., Assoc. of Mycol. Soc. of Slovenia (1998), p 29.

   

Mycelium, OOAK Inamorata art doll featuring Shani head sculpt in Ice resin.

 

I originally made Mycilium for the Hitogata Ten art doll exhibition in Tokyo. Since tattoos are a bit of a taboo in Japan, and I didn't want to risk upsetting the gallery, I didn't fulfil my original vision for he doll at the time. However, when this doll sold after the exhibition, I pitched the idea of painting her entire body with a mycelial network that has little bugs scuttling throughout it. I was so happy that the new owner loved the idea and I got to finish the concept. Her white mulberry silk hard cap wig with branching out braids repeats the branching patterns of a mycelial network.

 

The next preorder for Inamorata dolls is on April 22nd and will include this translucent Ice resin tone.

Geastrum quadrifidum (Pers.) Pers., syn.: Geastrum coronatum Scopoli

Rayed Earthstar, DE: Kleiner Nest-Erdstern, Kronen Erdstern

Slo.: četverokraka zvezdica

 

Dat.: Sept. 09. 2014

Lat.: 46.36529 Long.: 13.74988

Code: Bot_835/2014_DSC3951

 

Habitat: mixed wood, Picea abies and Fagus sylvatica dominant, moderately steep, southeast oriented mountain slope, calcareous skeletal ground covered by leaf and needles litter without ground vegetation, under Picea abies, in shade, partly protected from direct rain by tree canopies, average precipitations ~ 3.000 mm/year, average temperature 5-7 deg C, elevation 950 m (3.100 feet), alpine phytogeographical region.

 

Substratum: on moss covering a small Picea abies stump in the last disintegration stage, decomposed to almost soil.

 

Place: Lower Trenta valley, near the trail from Trenta village to Zasavska koča na Prehodavcih mountain cottage, halfway between the village and Planina Lepoč, East Julian Alps, Posočje, Slovenia EC

 

Comments: Genus Geastrum contains very attractive fungi, which almost all are rather uncommon, if not rare. Globose or onion shaped fruit bodies of many start to develop underground. The 'shell' of their fruit bodies consists of four distinct layers (with some exceptions). The outer three form so called exoperidium and the inner one is endoperidium, a 'sack', which contains spores. Exoperidium's outer layer consists of mycelium, the middle layer consists of fibers and the inner one is so called pseudoparenchymal layer. During growth the last one swells and breaks the exoperidium into star like shaped lappets, which curl backward and in this way push the fruit body out of the ground. In some species, like with Geastrum quadrifidum, the outer mycelial layer does not split together with other two layers of the exoperidium but falls off and forms a kind of 'bird's nest' in ground on top of which the fruit body sits. This white mycelial 'nest' can be seen on Fig. 3. The fruit body, when mature, cuts itself almost completely off the mycelium and stands free, like on legs made of exoperidium flaps. Only the far ends of the laps stay in connection with the 'nest'. In this way endoperidium with its 'chimney' (peristom) on top, through which clouds of spores rise like a 'smoke', is positioned as high as possible to facilitate spore spreading by the wind.

 

Geastrum quadrifidum is among the smallest species of about 50 of them worldwide (and ten of them described in Slovenian checklist (Ref.7)). It is a rare find. As its species name suggests it usually has four exoperidium 'legs'. However, sometimes, as in my find, it has five of them.

 

Growing solitary; exoperidium diameter 20 mm, endoperidium diameter 8 mm, its height (without the peristom 'beak') 7 mm; SP and spores on mass dark brown; smell none; taste not tested (too small).

 

Spores coarsely warty. Dimensions excluding warts: 4.4 [5 ; 5.2] 5.7 x 4.1 [4.5 ; 4.7] 5.2 microns; Q = 1 [1.1] 1.2; N = 40; C = 95%; Me = 5.1 x 4.6 microns; Qe = 1.1; number of warts per circumference: AVG = 12.1, SD = 1.2, N = 30. Olympus CH20, NEA 100x/1.25, magnification 1.000 x, oil; AmScope MA500 digital camera.

 

Herbarium: Mycotheca and lichen herbarium (LJU-Li) of Slovenian Forestry Institute, Večna pot 2, Ljubljana, Index Herbariorum LJF

 

Ref.:

(1) G.J. Krieglsteiner (Hrsg.), Die Grosspilze Baden-Württembergs, Band 2., Ulmer (2000), p 116.

(2) S. Buczacki, Collins Fungi Guide, Collins (2012), p 440.

(3) W. Rothmaler, Exkursionsflora von Deutschland, Vol.1, Niedere Pflanzen, Elsevier, 3.Auflage, (1994), p 519.

(4) M. Bon, Parey's Buch der Pilze, Kosmos (2005), p 302.

(5) J. Breitenbach, F. Kraenzlin, Eds., Fungi of Switzerland, Vol.2. Verlag Mykologia(1986), p 382.

(6) R. Phillips, Mushrooms, Macmillan (2006), p 334.

(7) A. Poler, ed., Seznam gliv Slovenije (in Slovene), 2nd Ed., Assoc. of Mycol. Soc. of Slovenia (1998), p 29.

   

A fungus (pl.: fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as one of the traditional eukaryotic kingdoms, along with Animalia, Plantae and either Protista or Protozoa and Chromista.

 

A characteristic that places fungi in a different kingdom from plants, bacteria, and some protists is chitin in their cell walls. Fungi, like animals, are heterotrophs; they acquire their food by absorbing dissolved molecules, typically by secreting digestive enzymes into their environment. Fungi do not photosynthesize. Growth is their means of mobility, except for spores (a few of which are flagellated), which may travel through the air or water. Fungi are the principal decomposers in ecological systems. These and other differences place fungi in a single group of related organisms, named the Eumycota (true fungi or Eumycetes), that share a common ancestor (i.e. they form a monophyletic group), an interpretation that is also strongly supported by molecular phylogenetics. This fungal group is distinct from the structurally similar myxomycetes (slime molds) and oomycetes (water molds). The discipline of biology devoted to the study of fungi is known as mycology (from the Greek μύκης mykes, mushroom). In the past mycology was regarded as a branch of botany, although it is now known that fungi are genetically more closely related to animals than to plants.

 

Abundant worldwide, most fungi are inconspicuous because of the small size of their structures, and their cryptic lifestyles in soil or on dead matter. Fungi include symbionts of plants, animals, or other fungi and also parasites. They may become noticeable when fruiting, either as mushrooms or as molds. Fungi perform an essential role in the decomposition of organic matter and have fundamental roles in nutrient cycling and exchange in the environment. They have long been used as a direct source of human food, in the form of mushrooms and truffles; as a leavening agent for bread; and in the fermentation of various food products, such as wine, beer, and soy sauce. Since the 1940s, fungi have been used for the production of antibiotics, and, more recently, various enzymes produced by fungi are used industrially and in detergents. Fungi are also used as biological pesticides to control weeds, plant diseases, and insect pests. Many species produce bioactive compounds called mycotoxins, such as alkaloids and polyketides, that are toxic to animals, including humans. The fruiting structures of a few species contain psychotropic compounds and are consumed recreationally or in traditional spiritual ceremonies. Fungi can break down manufactured materials and buildings, and become significant pathogens of humans and other animals. Losses of crops due to fungal diseases (e.g., rice blast disease) or food spoilage can have a large impact on human food supplies and local economies.

 

The fungus kingdom encompasses an enormous diversity of taxa with varied ecologies, life cycle strategies, and morphologies ranging from unicellular aquatic chytrids to large mushrooms. However, little is known of the true biodiversity of the fungus kingdom, which has been estimated at 2.2 million to 3.8 million species. Of these, only about 148,000 have been described, with over 8,000 species known to be detrimental to plants and at least 300 that can be pathogenic to humans. Ever since the pioneering 18th and 19th century taxonomical works of Carl Linnaeus, Christiaan Hendrik Persoon, and Elias Magnus Fries, fungi have been classified according to their morphology (e.g., characteristics such as spore color or microscopic features) or physiology. Advances in molecular genetics have opened the way for DNA analysis to be incorporated into taxonomy, which has sometimes challenged the historical groupings based on morphology and other traits. Phylogenetic studies published in the first decade of the 21st century have helped reshape the classification within the fungi kingdom, which is divided into one subkingdom, seven phyla, and ten subphyla.

 

Etymology

The English word fungus is directly adopted from the Latin fungus (mushroom), used in the writings of Horace and Pliny. This in turn is derived from the Greek word sphongos (σφόγγος 'sponge'), which refers to the macroscopic structures and morphology of mushrooms and molds; the root is also used in other languages, such as the German Schwamm ('sponge') and Schimmel ('mold').

 

The word mycology is derived from the Greek mykes (μύκης 'mushroom') and logos (λόγος 'discourse'). It denotes the scientific study of fungi. The Latin adjectival form of "mycology" (mycologicæ) appeared as early as 1796 in a book on the subject by Christiaan Hendrik Persoon. The word appeared in English as early as 1824 in a book by Robert Kaye Greville. In 1836 the English naturalist Miles Joseph Berkeley's publication The English Flora of Sir James Edward Smith, Vol. 5. also refers to mycology as the study of fungi.

 

A group of all the fungi present in a particular region is known as mycobiota (plural noun, no singular). The term mycota is often used for this purpose, but many authors use it as a synonym of Fungi. The word funga has been proposed as a less ambiguous term morphologically similar to fauna and flora. The Species Survival Commission (SSC) of the International Union for Conservation of Nature (IUCN) in August 2021 asked that the phrase fauna and flora be replaced by fauna, flora, and funga.

 

Characteristics

 

Fungal hyphae cells

Hyphal wall

Septum

Mitochondrion

Vacuole

Ergosterol crystal

Ribosome

Nucleus

Endoplasmic reticulum

Lipid body

Plasma membrane

Spitzenkörper

Golgi apparatus

 

Fungal cell cycle showing Dikaryons typical of Higher Fungi

Before the introduction of molecular methods for phylogenetic analysis, taxonomists considered fungi to be members of the plant kingdom because of similarities in lifestyle: both fungi and plants are mainly immobile, and have similarities in general morphology and growth habitat. Although inaccurate, the common misconception that fungi are plants persists among the general public due to their historical classification, as well as several similarities. Like plants, fungi often grow in soil and, in the case of mushrooms, form conspicuous fruit bodies, which sometimes resemble plants such as mosses. The fungi are now considered a separate kingdom, distinct from both plants and animals, from which they appear to have diverged around one billion years ago (around the start of the Neoproterozoic Era). Some morphological, biochemical, and genetic features are shared with other organisms, while others are unique to the fungi, clearly separating them from the other kingdoms:

 

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. 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).

With animals: Fungi lack chloroplasts and are heterotrophic organisms and so require preformed organic compounds as energy sources.

With plants: Fungi have a cell wall and vacuoles. They reproduce by both sexual and asexual means, and like basal plant groups (such as ferns and mosses) produce spores. Similar to mosses and algae, fungi typically have haploid nuclei.

With euglenoids and bacteria: Higher fungi, euglenoids, and some bacteria produce the amino acid L-lysine in specific biosynthesis steps, called the α-aminoadipate pathway.

The cells of most fungi grow as tubular, elongated, and thread-like (filamentous) structures called hyphae, which may contain multiple nuclei and extend by growing at their tips. Each tip contains a set of aggregated vesicles—cellular structures consisting of proteins, lipids, and other organic molecules—called the Spitzenkörper. Both fungi and oomycetes grow as filamentous hyphal cells. In contrast, similar-looking organisms, such as filamentous green algae, grow by repeated cell division within a chain of cells. There are also single-celled fungi (yeasts) that do not form hyphae, and some fungi have both hyphal and yeast forms.

In common with some plant and animal species, more than one hundred fungal species display bioluminescence.

Unique features:

 

Some species grow as unicellular yeasts that reproduce by budding or fission. Dimorphic fungi can switch between a yeast phase and a hyphal phase in response to environmental conditions.

The fungal cell wall is made of a chitin-glucan complex; while glucans are also found in plants and chitin in the exoskeleton of arthropods, fungi are the only organisms that combine these two structural molecules in their cell wall. Unlike those of plants and oomycetes, fungal cell walls do not contain cellulose.

A whitish fan or funnel-shaped mushroom growing at the base of a tree.

Omphalotus nidiformis, a bioluminescent mushroom

Most fungi lack an efficient system for the long-distance transport of water and nutrients, such as the xylem and phloem in many plants. To overcome this limitation, some fungi, such as Armillaria, form rhizomorphs, which resemble and perform functions similar to the roots of plants. As eukaryotes, fungi possess a biosynthetic pathway for producing terpenes that uses mevalonic acid and pyrophosphate as chemical building blocks. Plants and some other organisms have an additional terpene biosynthesis pathway in their chloroplasts, a structure that fungi and animals do not have. Fungi produce several secondary metabolites that are similar or identical in structure to those made by plants. Many of 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.

 

Diversity

Fungi have a worldwide distribution, and grow in a wide range of habitats, including extreme environments such as deserts or areas with high salt concentrations or ionizing radiation, as well as in deep sea sediments. Some can survive the intense UV and cosmic radiation encountered during space travel. Most grow in terrestrial environments, though several species live partly or solely in aquatic habitats, such as the chytrid fungi Batrachochytrium dendrobatidis and B. salamandrivorans, parasites that have been responsible for a worldwide decline in amphibian populations. These organisms spend part of their life cycle as a motile zoospore, enabling them to propel itself through water and enter their amphibian host. Other examples of aquatic fungi include those living in hydrothermal areas of the ocean.

 

As of 2020, around 148,000 species of fungi have been described by taxonomists, but the global biodiversity of the fungus kingdom is not fully understood. A 2017 estimate suggests there may be between 2.2 and 3.8 million species The number of new fungi species discovered yearly has increased from 1,000 to 1,500 per year about 10 years ago, to about 2000 with a peak of more than 2,500 species in 2016. In the year 2019, 1882 new species of fungi were described, and it was estimated that more than 90% of fungi remain unknown The following year, 2905 new species were described—the highest annual record of new fungus names. In mycology, species have historically been distinguished by a variety of methods and concepts. Classification based on morphological characteristics, such as the size and shape of spores or fruiting structures, has traditionally dominated fungal taxonomy. 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.

 

Mycology

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.

 

The use of fungi by humans dates back to prehistory; Ötzi the Iceman, a well-preserved mummy of a 5,300-year-old Neolithic man found frozen in the Austrian Alps, carried two species of polypore mushrooms that may have been used as tinder (Fomes fomentarius), or for medicinal purposes (Piptoporus betulinus). Ancient peoples have used fungi as food sources—often unknowingly—for millennia, in the preparation of leavened bread and fermented juices. Some of the oldest written records contain references to the destruction of crops that were probably caused by pathogenic fungi.

 

History

Mycology became a systematic science after the development of the microscope 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. Micheli not only observed spores but also showed that, under the proper conditions, they could be induced into growing into the same species of fungi from which they originated. Extending the use of the binomial system of nomenclature introduced by Carl Linnaeus in his Species plantarum (1753), the Dutch Christiaan Hendrik Persoon (1761–1836) established the first classification of mushrooms with such skill as to be considered a founder of modern mycology. Later, Elias Magnus Fries (1794–1878) further elaborated the classification of fungi, using spore color and microscopic characteristics, methods still used by taxonomists today. Other notable early contributors to mycology in the 17th–19th and early 20th centuries include Miles Joseph Berkeley, August Carl Joseph Corda, Anton de Bary, the brothers Louis René and Charles Tulasne, Arthur H. R. Buller, Curtis G. Lloyd, and Pier Andrea Saccardo. In the 20th and 21st centuries, advances in biochemistry, genetics, molecular biology, biotechnology, DNA sequencing and phylogenetic analysis has provided new insights into fungal relationships and biodiversity, and has challenged traditional morphology-based groupings in fungal taxonomy.

 

Morphology

Microscopic structures

Monochrome micrograph showing Penicillium hyphae as long, transparent, tube-like structures a few micrometres across. Conidiophores branch out laterally from the hyphae, terminating in bundles of phialides on which spherical condidiophores are arranged like beads on a string. Septa are faintly visible as dark lines crossing the hyphae.

An environmental isolate of Penicillium

Hypha

Conidiophore

Phialide

Conidia

Septa

Most fungi grow as hyphae, which are cylindrical, thread-like structures 2–10 µm in diameter and up to several centimeters in length. Hyphae grow at their tips (apices); new hyphae are typically formed by emergence of new tips along existing hyphae by a process called branching, or occasionally growing hyphal tips fork, giving rise to two parallel-growing hyphae. Hyphae also sometimes fuse when they come into contact, a process called hyphal fusion (or anastomosis). These growth processes lead to the development of a mycelium, an interconnected network of hyphae. Hyphae can be either septate or coenocytic. Septate hyphae are divided into compartments separated by cross walls (internal cell walls, called septa, that are formed at right angles to the cell wall giving the hypha its shape), with each compartment containing one or more nuclei; coenocytic hyphae are not compartmentalized. Septa have pores that allow cytoplasm, organelles, and sometimes nuclei to pass through; an example is the dolipore septum in fungi of the phylum Basidiomycota. Coenocytic hyphae are in essence multinucleate supercells.

 

Many species have developed specialized hyphal structures for nutrient uptake from living hosts; examples include haustoria in plant-parasitic species of most fungal phyla,[63] and arbuscules of several mycorrhizal fungi, which penetrate into the host cells to consume nutrients.

 

Although fungi are opisthokonts—a grouping of evolutionarily related organisms broadly characterized by a single posterior flagellum—all phyla except for the chytrids have lost their posterior flagella. Fungi are unusual among the eukaryotes in having a cell wall that, in addition to glucans (e.g., β-1,3-glucan) and other typical components, also contains the biopolymer chitin.

 

Macroscopic structures

Fungal mycelia can become visible to the naked eye, for example, on various surfaces and substrates, such as damp walls and spoiled food, where they are commonly called 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. Some individual fungal colonies can reach extraordinary dimensions and ages as in the case of a clonal colony of Armillaria solidipes, which extends over an area of more than 900 ha (3.5 square miles), with an estimated age of nearly 9,000 years.

 

The apothecium—a specialized structure important in sexual reproduction in the ascomycetes—is a cup-shaped fruit body that is often macroscopic and holds the hymenium, a layer of tissue containing the spore-bearing cells. The fruit bodies of the basidiomycetes (basidiocarps) and some ascomycetes can sometimes grow very large, and many are well known as mushrooms.

 

Growth and physiology

Time-lapse photography sequence of a peach becoming progressively discolored and disfigured

Mold growth covering a decaying peach. The frames were taken approximately 12 hours apart over a period of six days.

The growth of fungi as hyphae on or in solid substrates or as single cells in aquatic environments is adapted for the efficient extraction of nutrients, because these growth forms have high surface area to volume ratios. Hyphae are specifically adapted for growth on solid surfaces, and to invade substrates and tissues. They can exert large penetrative mechanical forces; for example, many plant pathogens, including Magnaporthe grisea, form a structure called an appressorium that evolved to puncture plant tissues.[71] The pressure generated by the appressorium, directed against the plant epidermis, can exceed 8 megapascals (1,200 psi).[71] The filamentous fungus Paecilomyces lilacinus uses a similar structure to penetrate the eggs of nematodes.

 

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

 

Fungi are traditionally considered heterotrophs, organisms that rely solely on carbon fixed by other organisms for metabolism. Fungi have evolved a high degree of metabolic versatility that allows them to use a diverse range of organic substrates for growth, including simple compounds such as nitrate, ammonia, acetate, or ethanol. In some species the pigment melanin may play a role in extracting energy from ionizing radiation, such as gamma radiation. This form of "radiotrophic" growth has been described for only a few species, the effects on growth rates are small, and the underlying biophysical and biochemical processes are not well known. This process might bear similarity to CO2 fixation via visible light, but instead uses ionizing radiation as a source of energy.

 

Reproduction

Two thickly stemmed brownish mushrooms with scales on the upper surface, growing out of a tree trunk

Polyporus squamosus

Fungal reproduction is complex, reflecting the differences in lifestyles and genetic makeup within this diverse kingdom of organisms. It is estimated that a third of all fungi reproduce using more than one method of propagation; for example, reproduction may occur in two well-differentiated stages within the life cycle of a species, the teleomorph (sexual reproduction) and the anamorph (asexual reproduction). Environmental conditions trigger genetically determined developmental states that lead to the creation of specialized structures for sexual or asexual reproduction. These structures aid reproduction by efficiently dispersing spores or spore-containing propagules.

 

Asexual reproduction

Asexual reproduction occurs via vegetative spores (conidia) or through mycelial fragmentation. Mycelial fragmentation occurs when a fungal mycelium separates into pieces, and each component grows into a separate mycelium. Mycelial fragmentation and vegetative spores maintain clonal populations adapted to a specific niche, and allow more rapid dispersal than sexual reproduction. The "Fungi imperfecti" (fungi lacking the perfect or sexual stage) or Deuteromycota comprise all the species that lack an observable sexual cycle. Deuteromycota (alternatively known as Deuteromycetes, conidial fungi, or mitosporic fungi) is not an accepted taxonomic clade and is now taken to mean simply fungi that lack a known sexual stage.

 

Sexual reproduction

See also: Mating in fungi and Sexual selection in fungi

Sexual reproduction with meiosis has been directly observed in all fungal phyla except Glomeromycota (genetic analysis suggests meiosis in Glomeromycota as well). It differs in many aspects from sexual reproduction in animals or plants. Differences also exist between fungal groups and can be used to discriminate species by morphological differences in sexual structures and reproductive strategies. Mating experiments between fungal isolates may identify species on the basis of biological species concepts. The major fungal groupings have initially been delineated based on the morphology of their sexual structures and spores; for example, the spore-containing structures, asci and basidia, can be used in the identification of ascomycetes and basidiomycetes, respectively. Fungi employ two mating systems: heterothallic species allow mating only between individuals of the opposite mating type, whereas homothallic species can mate, and sexually reproduce, with any other individual or itself.

 

Most fungi have both a haploid and a diploid stage in their life cycles. In sexually reproducing fungi, compatible individuals may combine by fusing their hyphae together into an interconnected network; this process, anastomosis, is required for the initiation of the sexual cycle. Many ascomycetes and basidiomycetes go through a dikaryotic stage, in which the nuclei inherited from the two parents do not combine immediately after cell fusion, but remain separate in the hyphal cells (see heterokaryosis).

 

In ascomycetes, dikaryotic hyphae of the hymenium (the spore-bearing tissue layer) form a characteristic hook (crozier) at the hyphal septum. During cell division, the formation of the hook ensures proper distribution of the newly divided nuclei into the apical and basal hyphal compartments. An ascus (plural asci) is then formed, in which karyogamy (nuclear fusion) occurs. Asci are embedded in an ascocarp, or fruiting body. Karyogamy in the asci is followed immediately by meiosis and the production of ascospores. After dispersal, the ascospores may germinate and form a new haploid mycelium.

 

Sexual reproduction in basidiomycetes is similar to that of the ascomycetes. Compatible haploid hyphae fuse to produce a dikaryotic mycelium. However, the dikaryotic phase is more extensive in the basidiomycetes, often also present in the vegetatively growing mycelium. A specialized anatomical structure, called a clamp connection, is formed at each hyphal septum. As with the structurally similar hook in the ascomycetes, the clamp connection in the basidiomycetes is required for controlled transfer of nuclei during cell division, to maintain the dikaryotic stage with two genetically different nuclei in each hyphal compartment. A basidiocarp is formed in which club-like structures known as basidia generate haploid basidiospores after karyogamy and meiosis. The most commonly known basidiocarps are mushrooms, but they may also take other forms (see Morphology section).

 

In fungi formerly classified as Zygomycota, haploid hyphae of two individuals fuse, forming 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.

 

Spore dispersal

The spores of most of the researched species of fungi are transported by wind. Such species often produce dry or hydrophobic spores that do not absorb water and are readily scattered by raindrops, for example. In other species, both asexual and sexual spores or sporangiospores are often actively dispersed by forcible ejection from their reproductive structures. This ejection ensures exit of the spores from the reproductive structures as well as traveling through the air over long distances.

 

Specialized mechanical and physiological mechanisms, as well as spore surface structures (such as hydrophobins), enable efficient spore ejection. For example, the structure of the spore-bearing cells in some ascomycete species is such that the buildup of substances affecting cell volume and fluid balance enables the explosive discharge of spores into the air. The forcible discharge of single spores termed ballistospores involves formation of a small drop of water (Buller's drop), which upon contact with the spore leads to its projectile release with an initial acceleration of more than 10,000 g; the net result is that the spore is ejected 0.01–0.02 cm, sufficient distance for it to fall through the gills or pores into the air below. Other fungi, like the puffballs, rely on alternative mechanisms for spore release, such as external mechanical forces. The hydnoid fungi (tooth fungi) produce spores on pendant, tooth-like or spine-like projections. The bird's nest fungi use the force of falling water drops to liberate the spores from cup-shaped fruiting bodies. Another strategy is seen in the stinkhorns, a group of fungi with lively colors and putrid odor that attract insects to disperse their spores.

 

Homothallism

In homothallic sexual reproduction, two haploid nuclei derived from the same individual fuse to form a zygote that can then undergo meiosis. Homothallic fungi include species with an Aspergillus-like asexual stage (anamorphs) occurring in numerous different genera, several species of the ascomycete genus Cochliobolus, and the ascomycete Pneumocystis jirovecii. The earliest mode of sexual reproduction among eukaryotes was likely homothallism, that is, self-fertile unisexual reproduction.

 

Other sexual processes

Besides regular sexual reproduction with meiosis, certain fungi, such as those in the genera Penicillium and Aspergillus, may exchange genetic material via parasexual processes, initiated by anastomosis between hyphae and plasmogamy of fungal cells. The frequency and relative importance of parasexual events is unclear and may be lower than other sexual processes. It is known to play a role in intraspecific hybridization and is likely required for hybridization between species, which has been associated with major events in fungal evolution.

 

Evolution

In contrast to plants 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. Often recovered from a permineralized plant or animal host, these samples are typically studied by making thin-section preparations that can be examined with light microscopy or transmission electron microscopy. Researchers study compression fossils by dissolving the surrounding matrix with acid and then using light or scanning electron microscopy to examine surface details.

 

The earliest fossils possessing features typical of fungi date to the Paleoproterozoic era, some 2,400 million years ago (Ma); these multicellular benthic organisms had filamentous structures capable of anastomosis. Other studies (2009) estimate the arrival of fungal organisms at about 760–1060 Ma on the basis of comparisons of the rate of evolution in closely related groups. The oldest fossilizied mycelium to be identified from its molecular composition is between 715 and 810 million years old. For much of the Paleozoic Era (542–251 Ma), the fungi appear to have been aquatic and consisted of organisms similar to the extant chytrids in having flagellum-bearing spores. The evolutionary adaptation from an aquatic to a terrestrial lifestyle necessitated a diversification of ecological strategies for obtaining nutrients, including parasitism, saprobism, and the development of mutualistic relationships such as mycorrhiza and lichenization. Studies suggest that the ancestral ecological state of the Ascomycota was saprobism, and that independent lichenization events have occurred multiple times.

 

In May 2019, scientists reported the discovery of a fossilized fungus, named Ourasphaira giraldae, in the Canadian Arctic, that may have grown on land a billion years ago, well before plants were living on land. Pyritized fungus-like microfossils preserved in the basal Ediacaran Doushantuo Formation (~635 Ma) have been reported in South China. Earlier, it had been presumed that the fungi colonized the land during the Cambrian (542–488.3 Ma), also long before land plants. Fossilized hyphae and spores recovered from the Ordovician of Wisconsin (460 Ma) resemble modern-day Glomerales, and existed at a time when the land flora likely consisted of only non-vascular bryophyte-like plants. Prototaxites, which was probably a fungus or lichen, would have been the tallest organism of the late Silurian and early Devonian. Fungal fossils do not become common and uncontroversial until the early Devonian (416–359.2 Ma), when they occur abundantly in the Rhynie chert, mostly as Zygomycota and Chytridiomycota. At about this same time, approximately 400 Ma, the Ascomycota and Basidiomycota diverged, and all modern classes of fungi were present by the Late Carboniferous (Pennsylvanian, 318.1–299 Ma).

 

Lichens formed a component of the early terrestrial ecosystems, and the estimated age of the oldest terrestrial lichen fossil is 415 Ma; this date roughly corresponds to the age of the oldest known sporocarp fossil, a Paleopyrenomycites species found in the Rhynie Chert. The oldest fossil with microscopic features resembling modern-day basidiomycetes is Palaeoancistrus, found permineralized with a fern from the Pennsylvanian. Rare in the fossil record are the Homobasidiomycetes (a taxon roughly equivalent to the mushroom-producing species of the Agaricomycetes). Two amber-preserved specimens provide evidence that the earliest known mushroom-forming fungi (the extinct species Archaeomarasmius leggetti) appeared during the late Cretaceous, 90 Ma.

 

Some time after the Permian–Triassic extinction event (251.4 Ma), a fungal spike (originally thought to be an extraordinary abundance of fungal spores in sediments) formed, suggesting that fungi were the dominant life form at this time, representing nearly 100% of the available fossil record for this period. However, the relative proportion of fungal spores relative to spores formed by algal species is difficult to assess, the spike did not appear worldwide, and in many places it did not fall on the Permian–Triassic boundary.

 

Sixty-five million years ago, immediately after the Cretaceous–Paleogene extinction event that famously killed off most dinosaurs, there was a dramatic increase in evidence of fungi; apparently the death of most plant and animal species led to a huge fungal bloom like "a massive compost heap".

 

Taxonomy

Although commonly included in botany curricula and textbooks, fungi are more closely related to animals than to plants and are placed with the animals in the monophyletic group of opisthokonts. Analyses using molecular phylogenetics support a monophyletic origin of fungi. The taxonomy of fungi is in a state of constant flux, especially due to research based on DNA comparisons. These current phylogenetic analyses often overturn classifications based on older and sometimes less discriminative methods based on morphological features and biological species concepts obtained from experimental matings.

 

There is no unique generally accepted system at the higher taxonomic levels and there are frequent name changes at every level, from species upwards. Efforts among researchers are now underway to establish and encourage usage of a unified and more consistent nomenclature. Until relatively recent (2012) changes to the International Code of Nomenclature for algae, fungi and plants, fungal species could also have multiple scientific names depending on their life cycle and mode (sexual or asexual) of reproduction. Web sites such as Index Fungorum and MycoBank are officially recognized nomenclatural repositories and list current names of fungal species (with cross-references to older synonyms).

 

The 2007 classification of Kingdom Fungi is the result of a large-scale collaborative research effort involving dozens of mycologists and other scientists working on fungal taxonomy. It recognizes seven phyla, two of which—the Ascomycota and the Basidiomycota—are contained within a branch representing subkingdom Dikarya, the most species rich and familiar group, including all the mushrooms, most food-spoilage molds, most plant pathogenic fungi, and the beer, wine, and bread yeasts. The accompanying cladogram depicts the major fungal taxa and their relationship to opisthokont and unikont organisms, based on the work of Philippe Silar, "The Mycota: A Comprehensive Treatise on Fungi as Experimental Systems for Basic and Applied Research" and Tedersoo et al. 2018. The lengths of the branches are not proportional to evolutionary distances.

 

The major phyla (sometimes called divisions) of fungi have been classified mainly on the basis of characteristics of their sexual reproductive structures. As of 2019, nine major lineages have been identified: Opisthosporidia, Chytridiomycota, Neocallimastigomycota, Blastocladiomycota, Zoopagomycotina, Mucoromycota, Glomeromycota, Ascomycota and Basidiomycota.

 

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). Previously considered to be "primitive" protozoa, they are now thought to be either a basal branch of the Fungi, or a sister group–each other's closest evolutionary relative.

 

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.

 

The Blastocladiomycota were previously considered a taxonomic clade within the Chytridiomycota. 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.

 

The Neocallimastigomycota were earlier placed in the phylum Chytridiomycota. 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). They lack mitochondria but contain hydrogenosomes of mitochondrial origin. As in the related chrytrids, neocallimastigomycetes form zoospores that are posteriorly uniflagellate or polyflagellate.

 

Microscopic view of a layer of translucent grayish cells, some containing small dark-color spheres

Arbuscular mycorrhiza seen under microscope. Flax root cortical cells containing paired arbuscules.

Cross-section of a cup-shaped structure showing locations of developing meiotic asci (upper edge of cup, left side, arrows pointing to two gray cells containing four and two small circles), sterile hyphae (upper edge of cup, right side, arrows pointing to white cells with a single small circle in them), and mature asci (upper edge of cup, pointing to two gray cells with eight small circles in them)

Diagram of an apothecium (the typical cup-like reproductive structure of Ascomycetes) showing sterile tissues as well as developing and mature asci.

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. The symbiotic association between the Glomeromycota and plants is ancient, with evidence dating to 400 million years ago. 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. 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. 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. Medically relevant genera include Mucor, Rhizomucor, and Rhizopus.

 

The Ascomycota, commonly known as sac fungi or ascomycetes, constitute the largest taxonomic group within the Eumycota. 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. 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).

 

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, human commensal species of the genus Malassezia, and the opportunistic human pathogen, Cryptococcus neoformans.

 

Fungus-like organisms

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 an ambiregnal, duplicated taxonomy.

 

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 take in 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.

 

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.

 

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. Alternatively, Rozella can be classified as a basal fungal group.

 

The nucleariids may be the next sister group to the eumycete clade, and as such could be included in an expanded fungal kingdom. Many Actinomycetales (Actinomycetota), a group with many filamentous bacteria, were also long believed to be fungi.

 

Ecology

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 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.

 

Symbiosis

Many fungi have important symbiotic relationships with organisms from most if not all kingdoms. 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.

 

With plants

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.

 

A microscopic view of blue-stained cells, some with dark wavy lines in them

The dark filaments are hyphae of the endophytic fungus Epichloë coenophiala in the intercellular spaces of tall fescue leaf sheath tissue

The mycorrhizal symbiosis is ancient, dating back to at least 400 million years. 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. The fungal partners may also mediate plant-to-plant transfer of carbohydrates and other nutrients. Such mycorrhizal communities are called "common mycorrhizal networks". A special case of mycorrhiza is myco-heterotrophy, whereby the plant parasitizes the fungus, obtaining all of its nutrients from its fungal symbiont. Some fungal species inhabit the tissues inside roots, stems, and leaves, in which case they are called endophytes. 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.

 

With algae and cyanobacteria

A green, leaf-like structure attached to a tree, with a pattern of ridges and depression on the bottom surface

The lichen Lobaria pulmonaria, a symbiosis of fungal, algal, and cyanobacterial species

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. Lichens occur in every ecosystem on all continents, play a key role in soil formation and the initiation of biological succession, and are prominent in some extreme environments, including polar, alpine, and semiarid desert regions. They are able to grow on inhospitable surfaces, including bare soil, rocks, tree bark, wood, shells, barnacles and leaves. 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 27% of known fungi—more than 19,400 species—are lichenized. 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.

 

With insects

Many insects also engage in mutualistic relationships with fungi. Several groups of ants cultivate fungi in the order Chaetothyriales for several purposes: as a food source, as a structural component of their nests, and as a part of an ant/plant symbiosis in the domatia (tiny chambers in plants that house arthropods). Ambrosia beetles cultivate various species of fungi in the bark of trees that they infest. 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. 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. Termites on the African savannah are also known to cultivate fungi, 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. Fungi growing in dead wood are essential for xylophagous insects (e.g. woodboring beetles). They deliver nutrients needed by xylophages to nutritionally scarce dead wood. Thanks to this nutritional enrichment the larvae of the woodboring insect is able to grow and develop to adulthood. 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.

 

A thin brown stick positioned horizontally with roughly two dozen clustered orange-red leaves originating from a single point in the middle of the stick. These orange leaves are three to four times larger than the few other green leaves growing out of the stick, and are covered on the lower leaf surface with hundreds of tiny bumps. The background shows the green leaves and branches of neighboring shrubs.

The plant pathogen Puccinia magellanicum (calafate rust) causes the defect known as witch's broom, seen here on a barberry shrub in Chile.

 

Gram stain of Candida albicans from a vaginal swab from a woman with candidiasis, showing hyphae, and chlamydospores, which are 2–4 µm in diameter.

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, tree pathogens such as Ophiostoma ulmi and Ophiostoma novo-ulmi causing Dutch elm disease, Cryphonectria parasitica responsible for chestnut blight, and Phymatotrichopsis omnivora causing Texas Root Rot, and plant pathogens in the genera Fusarium, Ustilago, Alternaria, and Cochliobolus. 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. Many fungi that are plant pathogens, such as Magnaporthe oryzae, can switch from being biotrophic (parasitic on living plants) to being necrotrophic (feeding on the dead tissues of plants they have killed). This same principle is applied to fungi-feeding parasites, including Asterotremella albida, which feeds on the fruit bodies of other fungi both while they are living and after they are dead.

 

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, Histoplasma, and Pneumocystis. 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. Fungal spores are also a cause of allergies, and fungi from different taxonomic groups can evoke allergic reactions.

 

As targets of mycoparasites

Organisms that parasitize fungi are known as mycoparasitic organisms. About 300 species of fungi and fungus-like organisms, belonging to 13 classes and 113 genera, are used as biocontrol agents against plant fungal diseases. Fungi can also act as mycoparasites or antagonists of other fungi, such as Hypomyces chrysospermus, which grows on bolete mushrooms. Fungi can also become the target of infection by mycoviruses.

 

Communication

Main article: Mycorrhizal networks

There appears to be electrical communication between fungi in word-like components according to spiking characteristics.

 

Possible impact on climate

According to a study published in the academic journal Current Biology, fungi can soak from the atmosphere around 36% of global fossil fuel greenhouse gas emissions.

 

Mycotoxins

(6aR,9R)-N-((2R,5S,10aS,10bS)-5-benzyl-10b-hydroxy-2-methyl-3,6-dioxooctahydro-2H-oxazolo[3,2-a] pyrrolo[2,1-c]pyrazin-2-yl)-7-methyl-4,6,6a,7,8,9-hexahydroindolo[4,3-fg] quinoline-9-carboxamide

Ergotamine, a major mycotoxin produced by Claviceps species, which if ingested can cause gangrene, convulsions, and hallucinations

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. 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.

 

Mycotoxins are secondary metabolites (or natural products), and research has established the existence of biochemical pathways solely for the purpose of producing mycotoxins and other natural products in fungi. Mycotoxins may provide fitness benefits in terms of physiological adaptation, competition with other microbes and fungi, and protection from consumption (fungivory). Many fungal secondary metabolites (or derivatives) are used medically, as described under Human use below.

 

Pathogenic mechanisms

Ustilago maydis is a pathogenic plant fungus that causes smut disease in maize and teosinte. Plants have evolved efficient defense systems against pathogenic microbes such as U. maydis. A rapid defense reaction after pathogen attack is the oxidative burst where the plant produces reactive oxygen species 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. Furthermore, U. maydis has a well-established recombinational DNA repair system which acts during mitosis and meiosis. The system may assist the pathogen in surviving DNA damage arising from the host plant's oxidative defensive response to infection.

 

Cryptococcus neoformans is an encapsulated yeast that can live in both plants and animals. C. neoformans usually infects the lungs, where it is phagocytosed by alveolar macrophages. 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. 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. 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.

 

Human use

See also: Human interactions with fungi

Microscopic view of five spherical structures; one of the spheres is considerably smaller than the rest and attached to one of the larger spheres

Saccharomyces cerevisiae cells shown with DIC microscopy

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. Methods have been developed for genetic engineering of fungi, enabling metabolic engineering of fungal species. For example, genetic modification of yeast species—which are easy to grow at fast rates in large fermentation vessels—has opened up ways of pharmaceutical production that are potentially more efficient than production by the original source organisms. Fungi-based industries are sometimes considered to be a major part of a growing bioeconomy, with applications under research and development including use for textiles, meat substitution and general fungal biotechnology.

 

Therapeutic uses

Modern chemotherapeutics

Many species produce metabolites that are major sources of pharmacologically active drugs.

 

Antibiotics

Particularly important are the antibiotics, including the 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. 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. 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.

 

Other

Other drugs produced by fungi include griseofulvin isolated from Penicillium griseofulvum, used to treat fungal infections, 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. Psilocybin from fungi is investigated for therapeutic use and appears to cause global increases in brain network integration. Fungi produce compounds that inhibit viruses and cancer cells. 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. In Europe and Japan, polysaccharide-K (brand name Krestin), a chemical derived from Trametes versicolor, is an approved adjuvant for cancer therapy.

 

Traditional medicine

Upper surface view of a kidney-shaped fungus, brownish-red with a lighter yellow-brown margin, and a somewhat varnished or shiny appearance

Two dried yellow-orange caterpillars, one with a curly grayish fungus growing out of one of its ends. The grayish fungus is roughly equal to or slightly greater in length than the caterpillar, and tapers in thickness to a narrow end.

The fungi Ganoderma lucidum (left) and Ophiocordyceps sinensis (right) are used in traditional medicine practices

Certain mushrooms are used as supposed therapeutics in folk medicine practices, such as traditional Chinese medicine. Mushrooms with a history of such use include Agaricus subrufescens, Ganoderma lucidum, and Ophiocordyceps sinensis.

 

Cultured foods

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. Yeast species of the genus Saccharomyces are also used to produce alcoholic beverages through fermentation. Shoyu koji mold (Aspergillus oryzae) is an essential ingredient in brewing Shoyu (soy sauce) and sake, and the preparation of miso while Rhizopus species are used for making tempeh. 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. Quorn, a meat substitute, is made from Fusarium venenatum.

oyster mushroom mycelium growing on petri dishes in the lab. the plate on the left had hydrogen peroxide in the agar.

Identification of MYT1 and its distribution in fungi.

(A) Mycelial growth and perithecium formation of the Z39P105 mutant on potato dextrose agar (PDA) and carrot agar, respectively. Pictures were taken 3 d after inoculation and 7 d after sexual induction from PDA and carrot agar. Arrows indicate protoperithecia. (B) Molecular characterization of the vector insertion event in the Z39P105 genome. (C) Distribution of MYT1 in representative fungal species. The distribution image was constructed by using the BLASTMatrix tool that is available on the Comparative Fungal Genomics Platform (cfgp.riceblast.snu.ac.kr/) [62]. (D) Phylogenetic tree of MYT1 homologs in several fungal species. The alignment was performed with ClustalW, and the MEGA program Version 4.0 was used to perform a 1,000 bootstrap phylogenetic analysis using the neighbor joining method. amp, ampicillin resistance gene; hph, hygromycin B resistance gene. Pi, Phytophthora infestans; Pr, P. ramorum; Ps, P. sojae; Af, Aspergillus fumigatus; An, A. nidulans; Ao, A. oryzae; Bc, Botrytis cinerea; Fo, Fusarium oxysporum; Fv, F. verticillioides; Hc, Histoplasma capsulatum; Mo, Magnaporthe oryzae; Nc, Neurospora crassa; Pa, Podospora anserine; Ca, Candida albicans; Kl, Kluyveromyces lactis; Sc, Saccharomyces cerevisiae; Cc, Coprinus cinereus; Cn, Cryptococcus neoformans; Pc, Phanerochaete chrysosporium; nd, not detected.

view this photo large on black

 

see more interesting photo's from me here:

flickeflu.com/photos/77411963@N07/interesting

 

Armillaria solidipes (formerly Armillaria ostoyae), the honey mushroom, is the most common variant in the western U.S. of the group of species that all used to share the name Armillaria mellea. Armillaria solidipes is quite common on both hardwood and conifer wood in forests west of the Cascade crest. The mycelium attacks the sapwood and is able to travel great distances under the bark or between trees in the form of black rhizomorphs ("shoestrings").

 

In most areas of North America, Armillaria solidipes can be separated from other species by its physical features. Its brown colors, fairly prominent scales featured on its cap, and the well-developed ring on its stem sets it apart from any Armillaria.

 

It is known to be one of the largest living organisms, where scientists have estimated a single specimen found in Malheur National Forest in Oregon to have been growing for some 2,400 years, covering 3.4 square miles (8.4 km^2) and colloquially named the "Humongous Fungus." Armillaria solidipes grows and spreads primarily underground and the bulk of the organism lies in the ground, out of sight. Therefore, the organism is not visible to anyone viewing from the surface. It is only in the autumn when this organism will bloom “honey mushrooms”, visible evidence of the organism lying beneath. Low competition for land and nutrients have allowed this organism to grow so huge; it possibly covers more geographical area than any other living organism.

 

This fungus, like most parasitic fungi, reproduces sexually. The fungi begin their life as spores, released into the environment by a mature mushroom. Armillaria solidipes has a white spore print. There are two types of mating types for spores (not male and female but similar in effect). The spores can be dispersed by environment factors such as wind or they can be redeposited by an animal. Once the spores are in a resting state, the single spore must come in contact with a spore of an opposite mating type and of the same species. If the single spore isolates are from different species, the colonies will not fuse together and they will remain separate. When two isolates of the same species but different mating types fuse together, they soon form coalesced colonies which become dark brown and flat. With this particular fungus it will produce mycelial cords also known as rhizomorphs. These rhizomorphs allow the fungus to obtain nutrients from long distances away. These are also the main factors to its pathogenicity. As the fruiting body continues to grow and obtaining nutrients, it forms into a mature mushroom. Armillaria solidipes in particular grows a wide and thin sheet-like plates radiating from the stem which is known as its gills. The gills hold the spores of a mature mushroom. This is stained white when seen as a spore print. Once spore formation is complete, this signifies a mature mushroom and now is able to spread its spores to start a new generation.

 

The disease is of particular interest to forest managers, as the species is highly pathogenic to a number of commercial softwoods, notably Douglas-fir (Pseudotsuga menziesii), true firs (Abies spp.) and Western Hemlock (Tsuga heterophylla). A commonly prescribed treatment is the clear cutting of an infected stand followed by planting with more resistant species such as Western redcedar (Thuja plicata) or deciduous seedlings. Armillaria can remain viable in stumps for 50 years.

 

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Armillaria ostoyae, Sombere honingzwam.

 

Middelbruine tot geelbruine, vleeskleurige hoed en iets lichtere steel. Hoed met donkerder, afwisbare schubjes. De steel heeft een forse ring, met beneden de ring ook schubjes op de steel. De rand van de hoed is in vochtige toestand doorschijnend gestreept. Groeit meestal in bundels. Kleur van de sporen: wit tot crème. Hoogte: 6-15 cm, breedte: 3-10 cm.

 

Kan van september tot november gevonden worden op de stam, de basis en de wortels van bomen en op stronken van bomen. Meestal op min of meer zure zandgronden, komt algemeen voor.

 

Honingzwammen verspreiden zich niet alleen via hun sporen, maar ook via lange zwarte draden, rhizomorfen. Deze op veters lijkende zwarte draden kunnen onder de bast van aangetaste bomen gevonden worden. Voor de honingzwammen in het algemeen geldt, dat het parasieten zijn, die een sterke vorm van witrot veroorzaken, die uiteindelijk leidt tot het afsterven van de gastheer.

 

In april 2003 werd in het Malheur National Forest in de Amerikaanse staat Oregon een sombere honingzwam ontdekt van naar schatting 2400 jaar oud met een ondergrondse mycelium omvang van 8,9 km². Daarmee is deze schimmel het grootste organisme ter wereld. Ook in Zwitserland in het Nationaal Park in de streek Engadin komt deze schimmel met een grote omvang voor. Hier is de schimmel ongeveer duizend jaar oud en ongeveer 800 meter lang en 500 meter breed.

  

Geastrum quadrifidum (Pers.) Pers., syn.: Geastrum coronatum Scopoli

Rayed Earthstar, DE: Kleiner Nest-Erdstern, Kronen Erdstern

Slo.: četverokraka zvezdica

 

Dat.: Sept. 09. 2014

Lat.: 46.36529 Long.: 13.74988

Code: Bot_835/2014_DSC3951

 

Habitat: mixed wood, Picea abies and Fagus sylvatica dominant, moderately steep, southeast oriented mountain slope, calcareous skeletal ground covered by leaf and needles litter without ground vegetation, under Picea abies, in shade, partly protected from direct rain by tree canopies, average precipitations ~ 3.000 mm/year, average temperature 5-7 deg C, elevation 950 m (3.100 feet), alpine phytogeographical region.

 

Substratum: on moss covering a small Picea abies stump in the last disintegration stage, decomposed to almost soil.

 

Place: Lower Trenta valley, near the trail from Trenta village to Zasavska koča na Prehodavcih mountain cottage, halfway between the village and Planina Lepoč, East Julian Alps, Posočje, Slovenia EC

 

Comments: Genus Geastrum contains very attractive fungi, which almost all are rather uncommon, if not rare. Globose or onion shaped fruit bodies of many start to develop underground. The 'shell' of their fruit bodies consists of four distinct layers (with some exceptions). The outer three form so called exoperidium and the inner one is endoperidium, a 'sack', which contains spores. Exoperidium's outer layer consists of mycelium, the middle layer consists of fibers and the inner one is so called pseudoparenchymal layer. During growth the last one swells and breaks the exoperidium into star like shaped lappets, which curl backward and in this way push the fruit body out of the ground. In some species, like with Geastrum quadrifidum, the outer mycelial layer does not split together with other two layers of the exoperidium but falls off and forms a kind of 'bird's nest' in ground on top of which the fruit body sits. This white mycelial 'nest' can be seen on Fig. 3. The fruit body, when mature, cuts itself almost completely off the mycelium and stands free, like on legs made of exoperidium flaps. Only the far ends of the laps stay in connection with the 'nest'. In this way endoperidium with its 'chimney' (peristom) on top, through which clouds of spores rise like a 'smoke', is positioned as high as possible to facilitate spore spreading by the wind.

 

Geastrum quadrifidum is among the smallest species of about 50 of them worldwide (and ten of them described in Slovenian checklist (Ref.7)). It is a rare find. As its species name suggests it usually has four exoperidium 'legs'. However, sometimes, as in my find, it has five of them.

 

Growing solitary; exoperidium diameter 20 mm, endoperidium diameter 8 mm, its height (without the peristom 'beak') 7 mm; SP and spores on mass dark brown; smell none; taste not tested (too small).

 

Spores coarsely warty. Dimensions excluding warts: 4.4 [5 ; 5.2] 5.7 x 4.1 [4.5 ; 4.7] 5.2 microns; Q = 1 [1.1] 1.2; N = 40; C = 95%; Me = 5.1 x 4.6 microns; Qe = 1.1; number of warts per circumference: AVG = 12.1, SD = 1.2, N = 30. Olympus CH20, NEA 100x/1.25, magnification 1.000 x, oil; AmScope MA500 digital camera.

 

Herbarium: Mycotheca and lichen herbarium (LJU-Li) of Slovenian Forestry Institute, Večna pot 2, Ljubljana, Index Herbariorum LJF

 

Ref.:

(1) G.J. Krieglsteiner (Hrsg.), Die Grosspilze Baden-Württembergs, Band 2., Ulmer (2000), p 116.

(2) S. Buczacki, Collins Fungi Guide, Collins (2012), p 440.

(3) W. Rothmaler, Exkursionsflora von Deutschland, Vol.1, Niedere Pflanzen, Elsevier, 3.Auflage, (1994), p 519.

(4) M. Bon, Parey's Buch der Pilze, Kosmos (2005), p 302.

(5) J. Breitenbach, F. Kraenzlin, Eds., Fungi of Switzerland, Vol.2. Verlag Mykologia(1986), p 382.

(6) R. Phillips, Mushrooms, Macmillan (2006), p 334.

(7) A. Poler, ed., Seznam gliv Slovenije (in Slovene), 2nd Ed., Assoc. of Mycol. Soc. of Slovenia (1998), p 29.