Fusarium Sporotrichioides
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''Fusarium sporotrichioides'' is a
fungal A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from th ...
plant pathogen Plant pathology (also phytopathology) is the scientific study of diseases in plants caused by pathogens (infectious organisms) and environmental conditions (physiological factors). Organisms that cause infectious disease include fungi, oomyc ...
, one of various ''
Fusarium ''Fusarium'' is a large genus of filamentous fungi, part of a group often referred to as hyphomycetes, widely distributed in soil and associated with plants. Most species are harmless saprobes, and are relatively abundant members of the soil mi ...
'' species responsible for damaging crops, in particular causing a condition known as
Fusarium head blight Fusarium ear blight (FEB) (also called Fusarium head blight, FHB, or scab), is a fungal disease of cereals, including wheat, barley, oats, rye and triticale. FEB is caused by a range of ''Fusarium'' fungi, which infects the heads of the crop, reduc ...
in wheat, consequently being of notable agricultural and economic importance. The species is ecologically widespread, being found across
tropical The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...
and
temperate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout t ...
regions, and is a significant producer of
mycotoxins A mycotoxin (from the Greek μύκης , "fungus" and τοξίνη , "toxin") is a toxic secondary metabolite produced by organisms of kingdom Fungi and is capable of causing disease and death in both humans and other animals. The term 'mycotoxin' ...
, particularly
trichothecenes The trichothecenes are a large family of chemically related mycotoxins. They are produced by various species of ''Fusarium'', ''Myrothecium'', ''Trichoderma''/''Podostroma'', '' Trichothecium'', ''Cephalosporium'', '' Verticimonosporium'', and '' ...
. Although mainly infecting crops, ''F. sporotrichioides''-derived mycotoxins can have repercussions for human health in the case of the ingestion of infected
cereals A cereal is any grass cultivated for the edible components of its grain (botanically, a type of fruit called a caryopsis), composed of the endosperm, germ, and bran. Cereal grain crops are grown in greater quantities and provide more food en ...
. One such example includes the outbreak of alimentary toxic aleukia (ATA) in Russia, of which ''F. sporotrichioides''-infected crop was suspected to be the cause. Although current studies on ''F. sporotrichioides'' are somewhat limited in comparison to other species in the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
, ''Fusarium sporotrichioides'' has found several applications as a model system for experimentation in
molecular biology Molecular biology is the branch of biology that seeks to understand the molecular basis of biological activity in and between cells, including biomolecular synthesis, modification, mechanisms, and interactions. The study of chemical and physi ...
.


History and taxonomy

The genus ''Fusarium'' was first characterised by the German botanist Johann Link in 1809, prior to the recognition of fungal involvement in plant disease. Over a thousand different species of ''Fusarium'' were identified by the 1930s, however, upon further analysis, these were narrowed down to 65 different species. Despite this reduction in the recognised number of species, differentiating one from the other remained difficult and unclear. Lacking a satisfactory system of classification and identification with which to organise these many, seemingly similar ''Fusarium'' species, the
mycologists Mycology is the branch of biology concerned with the study of fungi, including their genetic and biochemical properties, their taxonomy and their use to humans, including as a source for tinder, traditional medicine, food, and entheogens, as w ...
Snyder and Hansen collected samples of various fusaria from laboratories worldwide, isolated single spores and cultured them for subsequent analysis of their genetic variation. Their work indicated the existence of 9 distinct species of ''Fusarium'', but there have since been several different classification systems that reach different conclusions, and consensus around this has been difficult to establish, perhaps in part because research focuses predominantly on the agriculturally or botanically more significant fusaria. ''F. sporotrichioides'' exemplifies these classification difficulties, as it is usually designated as belonging to the section Sporotrichiella, along with other similar species such as the more well-studied ''F. poae'' and ''F. tricinctum'', although other classification systems have placed ''F. sporotrichioides'' in the ''Arthrosporiella'' section, based on the similarity of its
conidia A conidium ( ; ), sometimes termed an asexual chlamydospore or chlamydoconidium (), is an asexual, non-motile spore of a fungus. The word ''conidium'' comes from the Ancient Greek word for dust, ('). They are also called mitospores due to the ...
l morphology to other species in the section. More currently, the
taxonomy Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
of the genus ''Fusarium'' is studied using high-performance liquid chromatography, with each of the peaks on the resulting chromatograph being detected by a
photodiode array A photodiode is a light-sensitive semiconductor diode. It produces current when it absorbs photons. The package of a photodiode allows light (or infrared or ultraviolet radiation, or X-rays) to reach the sensitive part of the device. The packa ...
and grouped into
chromophore A chromophore is the part of a molecule responsible for its color. The color that is seen by our eyes is the one not absorbed by the reflecting object within a certain wavelength spectrum of visible light. The chromophore is a region in the molec ...
families. As each species produces a different pattern of peaks, this technique allows accurate species identification, and has successfully been applied to several fusaria, including ''F. sporotrichioides''. For larger samples, thin-layer chromatography is usually used as a less expensive alternative.


Ecology

''Fusarium sporotrichioides'' are found in many
tropical The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...
and
temperate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout t ...
regions, usually in the soil or on wheat. Several members of the genus ''Fusarium'' are often found colonising the same area, with the proportion of the total ''Fusarium'' population each species constitutes fluctuating depending on extant weather conditions, as for example, colder weather hinders the growth of dominant species such as ''F. avenaceum'', allowing others such as ''F. culmorum'' to dominate. In contrast to other taxonomically-related fusaria that are mainly found on cereal crops, ''F. sporotrichioides'' is often soil-dwelling. Furthermore, other non-pathogenic or opportunistic ''Fusarium'' species are frequently found alongside ''F. sporotrichioides'' as part of a
phylogenetically In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
similar species complex.


Morphology

''Fusarium sporotrichioides'' is usually white in early growth, but syellow, brownish, red, pink, or purple later on. The
hyphae A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one or ...
are usually trinucleated, but can have up to eight nuclei. ''F. sportotrichioides'' usually has many aerial
mycelia Mycelium (plural mycelia) is a root-like structure of a fungus consisting of a mass of branching, thread-like hyphae. Fungal colonies composed of mycelium are found in and on soil and many other substrates. A typical single spore germinates in ...
, and may form reddish- or yellow-brown clusters of hyphae, called sporodochia. Yellow sporodochia turn purple upon addition of
alkaline In chemistry, an alkali (; from ar, القلوي, al-qaly, lit=ashes of the saltwort) is a base (chemistry), basic, ionic compound, ionic salt (chemistry), salt of an alkali metal or an alkaline earth metal. An alkali can also be defined as ...
substances such as
ammonia Ammonia is an inorganic compound of nitrogen and hydrogen with the formula . A stable binary hydride, and the simplest pnictogen hydride, ammonia is a colourless gas with a distinct pungent smell. Biologically, it is a common nitrogenous was ...
, whilst reddish-brown clusters turn yellow under acidic conditions. Members of this species have irregularly shaped, almost globular
microconidia A conidium ( ; ), sometimes termed an asexual chlamydospore or chlamydoconidium (), is an asexual, non-motile spore of a fungus. The word ''conidium'' comes from the Ancient Greek word for dust, ('). They are also called mitospores due to th ...
(referred to as subglobose), that are usually 5-7 μm in diameter, whilst their macroconidia are slightly curved and usually have three to five
septa The Southeastern Pennsylvania Transportation Authority (SEPTA) is a regional public transportation authority that operates bus, rapid transit, commuter rail, light rail, and electric trolleybus services for nearly 4 million people in five coun ...
. Many have numerous brown, globose
chlamydospores A chlamydospore is the thick-walled large resting spore of several kinds of fungi, including Ascomycota such as '' Candida'', Basidiomycota such as ''Panus'', and various Mortierellales species. It is the life-stage which survives in unfavourable c ...
that are 7 to 15 μm in diameter, and serve as an important feature for their distinction from other fusaria.


Growth and reproduction

''Fusarium sporotrichioides'' carries out both sexual and asexual mechanisms of reproduction, with its teleomorphs including ''Nectria'' and ''Gibberella''. It is a fast-growing fungus, usually able to grow up to 8-8.8 cm in diameter within four days. Its optimal growth temperature ranges from , with the minimum and maximum temperatures required for growth being and , respectively. The minimum humidity level required for vegetative growth is 88%. This species uses mainly
maltose } Maltose ( or ), also known as maltobiose or malt sugar, is a disaccharide formed from two units of glucose joined with an α(1→4) bond. In the isomer isomaltose, the two glucose molecules are joined with an α(1→6) bond. Maltose is the two- ...
,
starch Starch or amylum is a polymeric carbohydrate consisting of numerous glucose units joined by glycosidic bonds. This polysaccharide is produced by most green plants for energy storage. Worldwide, it is the most common carbohydrate in human diets ...
and
raffinose Raffinose is a trisaccharide composed of galactose, glucose, and fructose. It can be found in beans, cabbage, brussels sprouts, broccoli, asparagus, other vegetables, and whole grains. Raffinose can be hydrolyzed to D-galactose and sucrose by ...
as carbon sources for growth. Fusarial growth is also iron-dependent, and is therefore inhibited by
siderophores Siderophores (Greek: "iron carrier") are small, high-affinity iron-chelating compounds that are secreted by microorganisms such as bacteria and fungi. They help the organism accumulate iron. Although a widening range of siderophore functions is no ...
. These are small molecules with a high affinity for iron, that are produced by other soil-dwelling microorganisms and act as their iron-delivery system, thus interfering with the uptake of iron by ''Fusarium'' species and consequently preventing their germination.


Identification and detection methods

In earlier work, the differentiation of ''F. sporotrichioides'' from other fusaria is mainly based on differences in conidial morphology. For example, the basal cells of macroconidia in some ''Fusarium'' species have hooks or notches whilst others do not, but these differences are not always sufficient to distinguish closely related fusaria from one another. A feature unique to ''F. sporotrichioides'' compared to taxonomically related species is the presence of multiporous cells, known as polyphialides, which are now carefully considered in its identification. The fact that these polyphialides produce pyriform as well as fusiform microconidia is a further distinctive feature of ''F. sporotrichioides'', and the
blastospore A blastospore is an asexual fungal spore produced by budding. Produced by fungi within the phylum Glomeromycota and others. It is also known as a blastoconidium (plural = blastoconidia). An example of a fungus that forms blastospores is ''Candi ...
s of ''F. sporotrichioides'' are a crucial characteristic in its distinction from the similar ''F. tricinctum''. Over the past years, advances in molecular biology and the introduction of the
polymerase chain reaction The polymerase chain reaction (PCR) is a method widely used to rapidly make millions to billions of copies (complete or partial) of a specific DNA sample, allowing scientists to take a very small sample of DNA and amplify it (or a part of it) t ...
(PCR) have made the identification of ''Fusarium'' species a far more precise process. Today, species of ''Fusarium'' can be identified through the cloning and sequencing of
RAPD RAPD may refer to: *Relative afferent pupillary defect *Random amplification of polymorphic DNA Random amplification of polymorphic DNA (RAPD), pronounced "rapid", is a type of polymerase chain reaction (PCR), but the segments of DNA that are amp ...
fragments to produce primers for use in PCR that will consequently only amplify the DNA sequence of a specific species. Restriction Fragment Length Polymorphisms (RFLP) are also useful in the differentiation of fusaria, as differences in base-pair sequence cause the sample DNA sequences to be fragmented at different sites by
restriction enzymes A restriction enzyme, restriction endonuclease, REase, ENase or'' restrictase '' is an enzyme that cleaves DNA into fragments at or near specific recognition sites within molecules known as restriction sites. Restriction enzymes are one class o ...
, resulting in DNA fragments of different lengths. This identification method is particularly useful for screening large numbers of samples.


Media for identification

As conidial morphology can vary depending not only on factors such as temperature, but also on the components of the growth medium, precision, and consistency in the production and use of growth media is important.
potato dextrose agar Potato dextrose agar (BAM Media M127
from the U.S.
(PDA) is widely used for growing ''Fusarium'' species, but its use is limited by the fact that sporulation can take up to two months on this medium. Peptone
pentachloronitrobenzene Pentachloronitrobenzene, typically abbreviated PCNB, is a registered fungicide formally derived from nitrobenzene. It is either an off-white or yellow solid, depending on its purity, with a musty odor. Preparation PCNB was originally synthesized ...
(PCNB) medium, historically known as Nash medium, is considered the most effective
selective medium A growth medium or culture medium is a solid, liquid, or semi-solid designed to support the growth of a population of microorganisms or cells via the process of cell proliferation or small plants like the moss ''Physcomitrella patens''. Different ...
for the fusaria. Although its formulation is now considered somewhat outdated, PCNB has historically been useful for the rapid distinction between different ''Fusarium ''species in soil samples. Other media used for the identification of'' Fusarium ''species include oatmeal agar and potato sucrose agar, on which the main distinguishing characteristics of the species surface after roughly 10 to 14 days of growth.


Experimental applications in molecular biology

Several ''Fusarium'' species provide useful model systems for research in molecular biology. Considering ''F. sporotrichioides'' specifically, sequences of known genes of the species have been used to study potential virulence genes in other fusaria, for example in the characterisation of the
trichodiene synthase The enzyme trichodiene synthase (EC 4.2.3.6) catalyzes the chemical reaction :(2''E'',6''E'')-farnesyl diphosphate \rightleftharpoons trichodiene + diphosphate This enzyme belongs to the family of lyases, specifically those carbon-oxygen lyases ...
gene in ''F. graminearum''. Moreover, the generation of ''F. sporotrichioides''
mutant In biology, and especially in genetics, a mutant is an organism or a new genetic character arising or resulting from an instance of mutation, which is generally an alteration of the DNA sequence of the genome or chromosome of an organism. It ...
libraries has been a particularly useful approach to studying the phytotoxicity of the fusaria.


Applications to biochemistry and biotechnology

Several species of ''Fusarium'' including ''F. sporotrichioides'' have applications for biotechnology through their usefulness as hosts in which to express recombinant proteins, whilst others are used in the synthesis of
nanoparticles A nanoparticle or ultrafine particle is usually defined as a particle of matter that is between 1 and 100 nanometres (nm) in diameter. The term is sometimes used for larger particles, up to 500 nm, or fibers and tubes that are less than 1 ...
. For example,
zirconium dioxide Zirconium dioxide (), sometimes known as zirconia (not to be confused with zircon), is a white crystalline oxide of zirconium. Its most naturally occurring form, with a monoclinic crystalline structure, is the mineral baddeleyite. A dopant stabi ...
production can be induced in ''F. oxysporum'', and as
zirconium Zirconium is a chemical element with the symbol Zr and atomic number 40. The name ''zirconium'' is taken from the name of the mineral zircon, the most important source of zirconium. The word is related to Persian '' zargun'' (zircon; ''zar-gun'', ...
is a hard metal, this has applications for the production of small cutting tools. A further example includes ''F. semitectum'', which has been used for the synthesis of silver, but similar applications of ''F. sporotrichioides'' remain elusive.


Agricultural and economic importance

The genus ''Fusarium'' includes species that cause several crop diseases, including ear rot in maize, and head blight in wheat, thus contributing to significant crop yield reduction. Although ''F. sporotrichioides'' itself causes only wheat head blight, this disease is a significant concern in the agricultural industry, as crop yield losses due to head blight can be extensive. In Russia for example, head blight has in past years been responsible for 25 to 50% reductions in crop yield. ''F. sporotrichioides'', along with ''F. poae'' and ''F. avenaceum'' also cause the discolouration of cereals such as oats, and several Fusaria have been shown to contribute to the rotting of certain fruits and vegetables in suboptimal storage conditions. In the case of ''F. sporotrichioides'', the affected produce includes peas and apples.


Fusarium head blight

''F. sporotrichioides'' is one of the most common causative agents of head blight in
Scandinavia Scandinavia; Sámi languages: /. ( ) is a subregion#Europe, subregion in Northern Europe, with strong historical, cultural, and linguistic ties between its constituent peoples. In English usage, ''Scandinavia'' most commonly refers to Denmark, ...
, as well as Eastern and Northern Europe, although other species such as ''F. poae'' and ''F. avenaceum'' are usually more prevalent in these areas. Favourable temperature and humidity conditions are associated with an increased likelihood of infection of wheat by ''Fusarium'' species, with higher humidity being more conducive to infection, especially during the flowering period, or
anthesis Anthesis is the period during which a flower is fully open and functional. It may also refer to the onset of that period. The onset of anthesis is spectacular in some species. In ''Banksia'' species, for example, anthesis involves the extension ...
, of wheat. Fusarium head blight is caused by the release of
mycotoxins A mycotoxin (from the Greek μύκης , "fungus" and τοξίνη , "toxin") is a toxic secondary metabolite produced by organisms of kingdom Fungi and is capable of causing disease and death in both humans and other animals. The term 'mycotoxin' ...
from ''Fusarium'' species, which damage wheat kernels or spikelets. The infection of spikelets results in a loss of
chlorophyll Chlorophyll (also chlorophyl) is any of several related green pigments found in cyanobacteria and in the chloroplasts of algae and plants. Its name is derived from the Greek words , ("pale green") and , ("leaf"). Chlorophyll allow plants to a ...
, whilst in infected kernels, ''F. sporotrichioides''
mycelia Mycelium (plural mycelia) is a root-like structure of a fungus consisting of a mass of branching, thread-like hyphae. Fungal colonies composed of mycelium are found in and on soil and many other substrates. A typical single spore germinates in ...
extend from the kernel wall, or pericarp, resulting in a scaliness and discolouration. The infection usually spreads to other areas of the wheat head in favourable environmental conditions.


Fusarium mycotoxins

All pathogenic ''Fusarium'' species produce mycotoxins as secondary metabolites, with the optimal conditions for toxin production being low temperatures, , darkness, and a slightly acidic environment (pH around 5.6). Notably, the specific types of mycotoxins produced depend on the species in question, and significant diversity exists among species in this respect. This diversity of secondary metabolite synthesis and the respective genes involved is thought to have arisen via horizontal gene transfer. ''F. sporotrichioides'' produces the mycotoxins neosolaniol, nivalenol, NT-1 toxin, NT-2 toxin, HT-2 toxin, and T-2 toxin, all of which are
trichothecenes The trichothecenes are a large family of chemically related mycotoxins. They are produced by various species of ''Fusarium'', ''Myrothecium'', ''Trichoderma''/''Podostroma'', '' Trichothecium'', ''Cephalosporium'', '' Verticimonosporium'', and '' ...
. NT-1 and NT-2 toxins are inhibitors of protein synthesis, whilst nivalenol is a skin irritant and emetic, and can cause bone marrow degeneration. T-2 toxin is associated with skin
necrosis Necrosis () is a form of cell injury which results in the premature death of cells in living tissue by autolysis. Necrosis is caused by factors external to the cell or tissue, such as infection, or trauma which result in the unregulated dige ...
in mammals, as well as causing intestinal damage and acting as an emetic in trout and birds, respectively. Other mycotoxins produced by ''F. sporotrichioides'' include butenolide, which causes
mitochondria A mitochondrion (; ) is an organelle found in the Cell (biology), cells of most Eukaryotes, such as animals, plants and Fungus, fungi. Mitochondria have a double lipid bilayer, membrane structure and use aerobic respiration to generate adenosi ...
l damage in mammals and interferes with chlorophyll retention in plants, and moniliformin, which inhibits the
citric acid cycle The citric acid cycle (CAC)—also known as the Krebs cycle or the TCA cycle (tricarboxylic acid cycle)—is a series of chemical reactions to release stored energy through the oxidation of acetyl-CoA derived from carbohydrates, fats, and proteins ...
and consequently the breakdown of carbohydrates. Nontoxic secondary metabolites of ''F. sporotrichioides'' include various sterols, such as
ergosterol Ergosterol (ergosta-5,7,22-trien-3β-ol) is a sterol found in cell membranes of fungi and protozoa, serving many of the same functions that cholesterol serves in animal cells. Because many fungi and protozoa cannot survive without ergosterol, the ...
(an important cell membrane constituent), campesterol, and
sitosterol β-sitosterol (beta-sitosterol) is one of several phytosterols (plant sterols) with chemical structures similar to that of cholesterol. It is a white, waxy powder with a characteristic odor, and is one of the components of the food additive E499. ...
.


Implications for human health

Trichothecene mycotoxins such as HT-2 and T-2 toxin, both produced by ''F. sporotrichioides'', represent a concern for human health due to the possibility of contamination of cereal grains, although efforts are usually made to reduce the ''Fusarium'' contamination of wheat itself during the preharvest period, so the likelihood of mycotoxins being present in consumer-ready cereal products is relatively low. In the 1940s, however, reported outbreaks of alimentary toxic aleukia (ATA) (the depletion of leukocytes, the causative agent of which was ingested during food consumption) in parts of what was then the USSR were thought to have occurred via the ingestion of ''Fusarium''-infected
millet Millets () are a highly varied group of small-seeded grasses, widely grown around the world as cereal crops or grains for fodder and human food. Most species generally referred to as millets belong to the tribe Paniceae, but some millets al ...
. ATA has a notably severe pathology and significantly different clinical manifestations in comparison to other mycotoxicoses, including immune suppression, necrosis, and hemorrhaging from the throat, nose, and skin. Although Snyder and Hansen classified the causative agent of the outbreak as ''F. tricinctum'', the mycotoxicologist
Abraham Joffe Abraham Z. Joffe (1909–2000) was Professor of Mycology and Mycotoxicology at the Hebrew University, Jerusalem. Joffe's professional interests were centered primarily in toxigenic fungi associated with production of mycotoxins (aflatoxins, trich ...
identified it as ''F. sporotrichioides'', a conclusion supported by several sources.


Fusarium mycotoxins as biological weapons

The use of trichothecene mycotoxins as biological weapons has been suspected in the case of the controversial Soviet air attacks on several Southeast Asian countries in the 1970s and 1980s, referred to as the
yellow rain Yellow rain was a 1981 political incident in which the United States Secretary of State Alexander Haig accused the Soviet Union of supplying T-2 mycotoxin to the Communist states in Vietnam, Laos and Cambodia for use in counterinsurgency warfare. ...
attacks. Although whether or not intentional biological warfare actually took place remains unclear, soil analyses of the affected areas indicate the presence of above-normal levels of tricothecenes, as well as types of tricothecenes that are not usually produced by naturally occurring microorganisms in the area.


Control and management

Considering the fact that ''Fusarium'' diseases jeopardize crop viability as well as releasing potentially hazardous mycotoxins, their management and control is relevant to agriculture and public health. Field management is a useful control measure, as alternating wheat cultivation with that of other crop types that are not susceptible to ''Fusarium'' diseases interferes with ''Fusarium ''species colonization. Moreover, appropriate ploughing techniques can be implemented to eliminate the layer of fungi that accumulates on soil and consequently prevent the propagation of fusaria. Irrigation control can also significantly limit water-mediated dispersion of pathogenic ''Fusarium'' species, ultimately reducing the likelihood of crop contamination.


References

{{Taxonbar, from=Q5114748 sporotrichioides Fungal tree pathogens and diseases Nut tree diseases Fungi described in 1915