FUNGUS

A fungus (pronounced /ˈfʌŋgəs/) is a eukaryotic organism that is a member of

the kingdom Fungi (pronounced /ˈfʌndʒaɪ/).[2] The fungi are heterotrophic
organisms possessing a chitinous cell wall. The majority of species grow as
multicellular filaments called hyphae forming a mycelium; some fungal species
also grow as single cells. Sexual and asexual reproduction of the fungi is
commonly via spores, often produced on specialized structures or in fruiting
bodies. Some species have lost the ability to form reproductive structures, and
propagate solely by vegetative growth. Yeasts, molds, and mushrooms are
examples of fungi. The fungi are a monophyletic group that is phylogenetically
clearly distinct from the morphologically similar slime molds (myxomycetes) and
water molds (oomycetes). The fungi are more closely related to animals than
plants, yet the discipline of biology devoted to the study of fungi, known as
mycology, often falls under a branch of botany.

Occurring worldwide, most fungi are largely invisible to the naked eye, living for
the most part in soil, dead matter, and as symbionts of plants, animals, or other
fungi. They perform an essential role in all ecosystems in decomposing organic
matter and are indispensable in nutrient cycling and exchange. Some fungi
become noticeable when fruiting, either as mushrooms or molds. Many fungal
species have long been used as a direct source of food, such as mushrooms and
truffles and in fermentation of various food products, such as wine, beer, and soy
sauce. More recently, fungi are being used as sources for antibiotics used in
medicine and various enzymes, such as cellulases, pectinases, and proteases,
important for industrial use or as active ingredients of detergents. Many fungi
produce bioactive compounds called mycotoxins, such as alkaloids and
polyketides that are toxic to animals including humans. Some fungi are used
recreationally or in traditional ceremonies as a source of psychotropic
compounds. Several species of the fungi are significant pathogens of humans
and other animals, and losses due to diseases of crops (e.g., rice blast disease)
or food spoilage caused by fungi can have a large impact on human food supply
and local economies.

Etymology and definition

The English word fungus is directly adopted from the Latin fungus, meaning
"mushroom", used in Horace and Pliny.[3] This in turn is derived from the Greek
word sphongos/σφογγος ("sponge"), referring to the macroscopic structures and
morphology of some mushrooms and molds and also used in other languages
(e.g., the German Schwamm ("sponge") or Schwammerl for some types of
mushroom).

Characteristics

The fungi have a range of features defining the fungal kingdom, some of which
are shared with other organisms while others are unique to the fungi.[4]

Shared features:

• With eukaryotes: All fungi are eukaryotic, containing membrane-bound

nuclei with chromosomes. Fungal cells contain membrane-bound
cytoplasmic organelles, DNA with noncoding regions called introns, sterol-
containing membranes, and ribosomes of the 80S type. Fungi have a
characteristic range of soluble carbohydrates and storage compounds,
including mannitol and other sugar alcohols, trehalose and glycogen the
latter of which is also found in animals.

• With animals: Fungi lack chloroplasts and are heterotrophic organisms,

requiring preformed organic compounds as energy sources and also as
carbon skeletons for organic synthesis.

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

• With prokaryotes: As in some bacteria, biosynthesis of the amino acid, L-

lysine, is via the α-aminoadipate pathway.

• Some fungi grow as single-celled yeasts which reproduce by budding, and

some dimorphic fungi can switch between a yeast phase and a hyphal
phase in response to environmental conditions.
 • The fungal cell wall contains glucans also found in plants, but also chitin
not found in the Plant kingdom, but in some animals. In contrast to plants,
fungal cell walls do not contain cellulose.

• Fungal hyphae may have several nuclei within each hyphal compartment,
and many budding yeasts are diploid.

Diversity

Fungi have a worldwide distribution, and grow in a wide range of habitats,

including deserts, hypersaline environments[5], the deep sea[6], on rocks[7], in
extremely low and high temperatures. They have been shown to be able to
survive the intense UV and cosmic radiation encountered during space travel.[8]

Importance for human use

Sacharomyces cerevisiae cells in DIC microscopy.

Human use of fungi for food preparation or preservation and other purposes is
extensive and has a long history: yeasts are required for fermentation of beer,
wine [13] and bread, some other fungal species are used in the production of soy
sauce and tempeh. Mushroom farming and mushroom gathering are large
industries in many countries. Many fungi are producers of antibiotics, including β-
lactam antibiotics such as penicillin and cephalosporin.[14] Widespread use of
these antibiotics for the treatment of bacterial diseases, such as tuberculosis,
syphilis, leprosy, and many others began in the early 20th century and continues
to play a major part in anti-bacterial chemotherapy. The study of the historical
uses and sociological impact of fungi is known as ethnomycology.

Cultured foods

Baker's yeast or Saccharomyces cerevisiae, a single-cell fungus, is used in the

baking of bread and other wheat-based products, such as pizza and dumplings.
[15]
Several yeast species of the genus Saccharomyces are also used in the
production of alcoholic beverages through fermentation.[16] Mycelial fungi, such as
the shoyu koji mold (Aspergillus oryzae), are used in the brewing of Shoyu (soy
sauce) and preparation of tempeh.[17] Quorn is a high-protein product made from
the mold, Fusarium venenatum, and is used in vegetarian cooking.
Other human uses

Fungi are also used extensively to produce industrial chemicals like citric,
gluconic, lactic, and malic acids, antibiotics, and even to make stonewashed
jeans.[18] Fungi are also sources of industrial enzymes, such as lipases used in
biological detergents[19], amylases[20], cellulases[21], invertases, proteases and
xylanases[22]. Several fungal species are ingested for their psychedelic properties,
both recreationally and religiously (see main article, Psilocybin mushrooms).

Mycotoxins
Main article: Mycotoxins

Many fungi produce compounds with biological activity. Several of these

compounds are toxic and are therefore called mycotoxins, referring to their fungal
origin and toxic activity. Of particular relevance to humans are those mycotoxins
that are produced by moulds causing food spoilage and poisonous mushrooms
(see below). Particularly infamous are the aflatoxins, which are insidious liver
toxins and highly carcinogenic metabolites produced by Aspergillus species often
growing in or on grains and nuts consumed by humans, and the lethal amatoxins
produced by mushrooms of the genus Amanita. Other notable mycotoxins
include ochratoxins, patulin, ergot alkaloids, and trichothecenes and fumonisins,
all of which have significant impact on human food supplies or animal livestock.
[23]

Edible and poisonous fungi

Asian mushrooms, clockwise from left, enokitake, buna-shimeji, bunapi-shimeji,

king oyster mushroom and shiitake.

Black Périgord Truffle (Tuber melanosporum), cut in half.

Stilton cheese veined with Penicillium roqueforti.

Some of the best known types of fungi are the edible and the poisonous
mushrooms. Many species are commercially raised, but others must be
harvested from the wild. Agaricus bisporus, sold as button mushrooms when
small or Portobello mushrooms when larger, are the most commonly eaten
species, used in salads, soups, and many other dishes. Many Asian fungi are
commercially grown and have gained in popularity in the West. They are often
available fresh in grocery stores and markets, including straw mushrooms
(Volvariella volvacea), oyster mushrooms (Pleurotus ostreatus), shiitakes
(Lentinula edodes), and enokitake (Flammulina spp.).

Fungi in the biological control of pests

In agricultural settings, fungi that actively compete for nutrients and space with,
and eventually prevail over, pathogenic microorganisms, such as bacteria or
other fungi, via the competitive exclusion principle,[33] or are parasites of these
pathogens, may be beneficial agents for human use. For example, some fungi
may be used to suppress growth or eliminate harmful plant pathogens, such as
insects, mites, weeds, nematodes and other fungi that cause diseases of
important crop plants.[34] This has generated strong interest in the use and
practical application of these fungi for the biological control of these agricultural
pests. Entomopathogenic fungi can be used as biopesticides, as they actively kill
insects.[35] Examples of fungi that have been used as biological insecticides are
Beauveria bassiana, Metarhizium anisopliae, Hirsutella spp, Paecilomyces spp,
and Verticillium lecanii.[36] [37] Endophytic fungi of grasses of the genus
Neotyphodium, such as N. coenophialum produce alkaloids that are toxic to a
range of invertebrate and vertebrate herbivores. These alkaloids protect grass
plants from herbivory, but some endophyte alkaloids can cause poisoning of
grazing animals, such as cattle and sheep. [38] Infection of grass cultivars of
pasture or forage grasses with Neotyphodium endophytes selected for producing
only alkaloids that increase resistance to herbivores such as insects, while being
non-toxic to livestock, is being used in grass breeding programs.[39]
 Phylogeny and classification

The parasitic mushroom Oudemansiella nocturnum, infesting the wood of a tree.

For a long time taxonomists considered fungi to be members of the Plant

Kingdom. This early classification was based mainly on similarities in lifestyle:
both fungi and plant are mainly sessile, have similarities in general morphology
and growth habitat (like plants, fungi often grow in soil, in the case of mushrooms
forming conspicuous fruiting bodies, which sometimes bear resemblance to
plants such as mosses). Moreover, both groups possess a cell wall, which is
absent in the Animal Kingdom. However, the fungi are now considered a
separate kingdom, distinct from both plants and animals, from which they appear
to have diverged approximately one billion years ago.[88] Many studies have
identified several distinct morphological, biochemical, and genetic features in the
Fungi, clearly delineating this group from the other kingdoms. For these reasons,
the fungi are placed in their own kingdom.

Physiological and morphological traits

Similar to animals and unlike most plants, fungi lack the capacity to synthesize
organic carbon by chlorophyll-based photosynthesis; whereas plants store the
reduced carbon as starch, fungi, like animals and some bacteria, use glycogen [89]
for storage of carbohydrates. A major component of the cell wall in many fungal
species is the nitrogen-containing carbohydrate, chitin,[90] also present in some
animals, such as the insects and crustaceans, while the plant cell wall consists
chiefly of the carbohydrate cellulose. The defining and unique characteristics of
fungal cells include growth as hyphae, which are microscopic filaments of
between 2-10 microns in diameter and up to several centimetres in length, and
which combined form the fungal mycelium. Some fungi, such as yeasts, grow as
single ovoid cells, similar to unicellular algae and the protists.

Unlike many plants, most fungi lack an efficient vascular system, such as xylem
or phloem for long-distance transport of water and nutrients; as an example for
convergent evolution, some fungi, such as Armillaria, form rhizomorphs or
mycelial cords,[91] resembling and functionally related to, but morphologically
distinct from, plant roots.

Some characteristics shared between plants and fungi include the presence of
vacuoles in the cell,[92] and a similar pathway in the biosynthesis of terpenes
using mevalonic acid and pyrophosphate as biochemical precursors; plants
however use an additional terpene biosynthesis pathway in the chloroplasts that
is apparently absent in fungi.[93] Ancestral traits shared among members of the
fungi include chitinous cell walls and heterotrophy by absorption.[82] A further
characteristic of the fungi that is absent from other eukaryotes, and shared only
with some bacteria, is the biosynthesis of the amino acid, L-lysine, via the α-
aminoadipate pathway. [94]

Similar to plants, fungi produce a plethora of secondary metabolites functioning

as defensive compounds or for niche adaptation; however, biochemical pathways
for the synthesis of similar or even identical compounds often differ markedly
between fungi and plants. [95][96]