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Aspergillus niger 01.jpg
Conidial head of Aspergillus niger
Scientific classification Red Pencil Icon.png
Kingdom: Fungi
Division: Ascomycota
Class: Eurotiomycetes
Order: Eurotiales
Family: Trichocomaceae
Genus: Aspergillus
Micheli (1729)

See List of Aspergillus species

Aspergillus ( /ˌæspərˈɪləs/ ) is a genus consisting of several hundred mold species found in various climates worldwide.


Aspergillus was first catalogued in 1729 by the Italian priest and biologist Pier Antonio Micheli. Viewing the fungi under a microscope, Micheli was reminded of the shape of an aspergillum (holy water sprinkler), from Latin spargere (to sprinkle), and named the genus accordingly. [1] Aspergillum is an asexual spore-forming structure common to all Aspergillus species; around one-third of species are also known to have a sexual stage. [2] While some species of Aspergillus are known to cause fungal infections, others are of commercial importance.



Aspergillus consists of 837 species of fungi. [3]

Growth and distribution

Aspergillus on a tomato in detail Aspergillus on tomato.jpg
Aspergillus on a tomato in detail

Aspergillus is defined as a group of conidial fungi—that is, fungi in an asexual state. Some of them, however, are known to have a teleomorph (sexual state) in the Ascomycota. With DNA evidence, all members of the genus Aspergillus are members of the phylum Ascomycota.

Members of the genus possess the ability to grow where a high osmotic pressure exists (high concentration of sugar, salt, etc.). Aspergillus species are highly aerobic and are found in almost all oxygen-rich environments, where they commonly grow as molds on the surface of a substrate, as a result of the high oxygen tension. Commonly, fungi grow on carbon-rich substrates like monosaccharides (such as glucose) and polysaccharides (such as amylose). Aspergillus species are common contaminants of starchy foods (such as bread and potatoes), and grow in or on many plants and trees.[ citation needed ]

In addition to growth on carbon sources, many species of Aspergillus demonstrate oligotrophy where they are capable of growing in nutrient-depleted environments, or environments with a complete lack of key nutrients. Aspergillus niger is a prime example of this; it can be found growing on damp walls, as a major component of mildew.

Several species of Aspergillus, including A. niger and A. fumigatus , will readily colonise buildings, [4] favouring warm and damp or humid areas such as bathrooms and around window frames. [5]

Aspergillus are found in millions of pillows. [6]

Commercial importance

Various Penicillium, Aspergillus spp. and other fungi growing in axenic culture Ascomycetes.jpg
Various Penicillium, Aspergillus spp. and other fungi growing in axenic culture
Historical model of Aspergillus, Botanical Museum Greifswald Modell von Aspergillus (Giesskannenschimmel) -Osterloh- -Brendel 10f-.jpg
Historical model of Aspergillus, Botanical Museum Greifswald

Species of Aspergillus are important medically and commercially. Some species can cause infection in humans and other animals. Some infections found in animals have been studied for years, while other species found in animals have been described as new and specific to the investigated disease, and others have been known as names already in use for organisms such as saprophytes. More than 60 Aspergillus species are medically relevant pathogens. [7] For humans, a range of diseases such as infection to the external ear, skin lesions, and ulcers classed as mycetomas are found.

Other species are important in commercial microbial fermentations. For example, alcoholic beverages such as Japanese sake are often made from rice or other starchy ingredients (like manioc), rather than from grapes or malted barley. Typical microorganisms used to make alcohol, such as yeasts of the genus Saccharomyces , cannot ferment these starches. Therefore, koji mold such as Aspergillus oryzae is used to first break down the starches into simpler sugars. [8]

Members of the genus are also sources of natural products that can be used in the development of medications to treat human disease. [9] Aspergillus spp. are known to produce anthraquinone which has commercial importance due to its antibacterial and antifungal properties. [10]

Perhaps the largest application of Aspergillus niger is as the major source of citric acid; this organism accounts for over 99% of global citric acid production, or more than 1.4 million tonnes (>1.5 million US tons) per year.[ citation needed ]A. niger is also commonly used for the production of native and foreign enzymes, including glucose oxidase, lysozyme, and lactase. [11] In these instances, the culture is rarely grown on a solid substrate, although this is still common practice in Japan, but is more often grown as a submerged culture in a bioreactor. In this way, the most important parameters can be strictly controlled, and maximal productivity can be achieved. This process also makes it far easier to separate the chemical or enzyme of importance from the medium, and is therefore far more cost-effective.


Four three-day-old Aspergillus colonies. Clockwise from top-left: an A. nidulans laboratory strain; a similar strain with a mutation in the yA marker gene involved in green pigmentation; an A. oryzae strain used in soy fermentation; A. oryzae RIB40 Four 3-day old Aspergillus colonies on a Petri dish.png
Four three-day-old Aspergillus colonies. Clockwise from top-left: an A. nidulans laboratory strain; a similar strain with a mutation in the yA marker gene involved in green pigmentation; an A. oryzae strain used in soy fermentation; A. oryzae RIB40
A scan of Aspergillus taken at 235 magnifications under a scanning electron microscope Aspergillus 235 mags 3X3 copy.jpeg
A scan of Aspergillus taken at 235 magnifications under a scanning electron microscope

A. nidulans (Emericella nidulans) has been used as a research organism for many years and was used by Guido Pontecorvo to demonstrate parasexuality in fungi. Recently, A. nidulans was one of the pioneering organisms to have its genome sequenced by researchers at the Broad Institute. As of 2008, a further seven Aspergillus species have had their genomes sequenced: the industrially useful A. niger (two strains), A. oryzae , and A. terreus , and the pathogens A. clavatus , A. fischerianus (Neosartorya fischeri), A. flavus , and A. fumigatus (two strains). [12] A. fischerianus is hardly ever pathogenic, but is very closely related to the common pathogen A. fumigatus; it was sequenced in part to better understand A. fumigatus pathogenicity. [13]

Sexual reproduction

Of the 250 species of aspergilli, about 64% have no known sexual state. [14] However, many of these species likely have an as yet unidentified sexual stage. [14] Sexual reproduction occurs in two fundamentally different ways in fungi. These are outcrossing (in heterothallic fungi) in which two different individuals contribute nuclei, and self-fertilization or selfing (in homothallic fungi) in which both nuclei are derived from the same individual. In recent years, sexual cycles have been discovered in numerous species previously thought to be asexual. These discoveries reflect recent experimental focus on species of particular relevance to humans.

A. fumigatus is the most common species to cause disease in immunodeficient humans. In 2009, A. fumigatus was shown to have a heterothallic, fully functional sexual cycle. [15] Isolates of complementary mating types are required for sex to occur.

A. flavus is the major producer of carcinogenic aflatoxins in crops worldwide. It is also an opportunistic human and animal pathogen, causing aspergillosis in immunocompromised individuals. In 2009, a sexual state of this heterothallic fungus was found to arise when strains of opposite mating types were cultured together under appropriate conditions. [16]

A. lentulus is an opportunistic human pathogen that causes invasive aspergillosis with high mortality rates. In 2013, A. lentulus was found to have a heterothallic functional sexual breeding system. [17]

A. terreus is commonly used in industry to produce important organic acids and enzymes, and was the initial source for the cholesterol-lowering drug lovastatin. In 2013, A. terreus was found to be capable of sexual reproduction when strains of opposite mating types were crossed under appropriate culture conditions. [18]

These findings with Aspergillus species are consistent with accumulating evidence, from studies of other eukaryotic species, that sex was likely present in the common ancestor of all eukaryotes. [19] [20]

A. nidulans, a homothallic fungus, is capable of self-fertilization. Selfing involves activation of the same mating pathways characteristic of sex in outcrossing species, i.e. self-fertilization does not bypass required pathways for outcrossing sex, but instead requires activation of these pathways within a single individual. [21]

Among those Aspergillus species that exhibit a sexual cycle, the overwhelming majority in nature are homothallic (self-fertilizing). [22] This observation suggests Aspergillus species can generally maintain sex though little genetic variability is produced by homothallic self-fertilization. A. fumigatus, a heterothallic (outcrossing) fungus that occurs in areas with widely different climates and environments, also displays little genetic variability either within geographic regions or on a global scale, [23] again suggesting sex, in this case outcrossing sex, can be maintained even when little genetic variability is produced.


The simultaneous publication of three Aspergillus genome manuscripts in Nature in December 2005 established the genus as the leading filamentous fungal genus for comparative genomic studies. Like most major genome projects, these efforts were collaborations between a large sequencing centre and the respective community of scientists. For example, the Institute for Genome Research (TIGR) worked with the A. fumigatus community. A. nidulans was sequenced at the Broad Institute. A. oryzae was sequenced in Japan at the National Institute of Advanced Industrial Science and Technology. The Joint Genome Institute of the Department of Energy has released sequence data for a citric acid-producing strain of A. niger. TIGR, now renamed the J. Craig Venter Institute, is currently spearheading a project on the A. flavus genome. [24]

Aspergillus is characterized by high levels of genetic diversity and, using protostome divergence as a scale, is as diverse as the Vertebrates phylum although both inter and intra-specific genome structure is relatively plastic. [25] The genomes of some Aspergillus species, such as A. flavus and A. oryzae, are more rich and around 20% larger than others, such as A. nidulans and A. fumigatus. Several mechanisms could explain this difference, although the combination of segmental duplication, genome duplication, and horizontal gene transfer acting in a piecemeal fashion is well-supported. [26]

Genome sizes for sequenced species of Aspergillus range from about 29.3 Mb for A. fumigatus to 37.1 Mb for A. oryzae, while the numbers of predicted genes vary from about 9926 for A. fumigatus to about 12,071 for A. oryzae. The genome size of an enzyme-producing strain of A. niger is of intermediate size at 33.9 Mb. [1]


Some Aspergillus species cause serious disease in humans and animals. The most common pathogenic species are A. fumigatus and A. flavus , which produces aflatoxin which is both a toxin and a carcinogen, and which can contaminate foods such as nuts. The most common species causing allergic disease are A. fumigatus and A. clavatus . Other species are important as agricultural pathogens. Aspergillus spp. cause disease on many grain crops, especially maize, and some variants synthesize mycotoxins, including aflatoxin. Aspergillus can cause neonatal infections. [27]

A. fumigatus (the most common species) infections are primary pulmonary infections and can potentially become a rapidly necrotizing pneumonia with a potential to disseminate. The organism can be differentiated from other common mold infections based on the fact that it takes on a mold form both in the environment and in the host (unlike Candida albicans which is a dimorphic mold in the environment and a yeast in the body).


Pulmonary aspergillosis Pulmonary aspergillosis.jpg
Pulmonary aspergillosis

Aspergillosis is the group of diseases caused by Aspergillus. The most common species among paranasal sinus infections associated with aspergillosis is A. fumigatus . [28] The symptoms include fever, cough, chest pain, or breathlessness, which also occur in many other illnesses, so diagnosis can be difficult. Usually, only patients with already weakened immune systems or who suffer other lung conditions are susceptible.

In humans, the major forms of disease are: [29] [30]

Fungal infections from Aspergillus spores remain one theory of sickness and untimely death of some early Egyptologists and tomb explorers. Ancient spores which grew on the remains of food offerings and mummies sealed in tombs and chambers may have been blown around and inhaled by the excavators, ultimately linked to the notion of the curse of the pharaohs. [31]

Aspergillosis of the air passages is also frequently reported in birds, and certain species of Aspergillus have been known to infect insects. [7]

Most people inhale Aspergillus into their lungs everyday. [32] But it's generally only the immuno-compromised who get sick with Aspergillosis. [32]

See also

Related Research Articles

<span class="mw-page-title-main">Ascomycota</span> Division or phylum of fungi

Ascomycota is a phylum of the kingdom Fungi that, together with the Basidiomycota, forms the subkingdom Dikarya. Its members are commonly known as the sac fungi or ascomycetes. It is the largest phylum of Fungi, with over 64,000 species. The defining feature of this fungal group is the "ascus", a microscopic sexual structure in which nonmotile spores, called ascospores, are formed. However, some species of the Ascomycota are asexual, meaning that they do not have a sexual cycle and thus do not form asci or ascospores. Familiar examples of sac fungi include morels, truffles, brewers' and bakers' yeast, dead man's fingers, and cup fungi. The fungal symbionts in the majority of lichens such as Cladonia belong to the Ascomycota.

<i>Aspergillus niger</i> Species of fungus

Aspergillus niger is a mold classified within the Nigri section of the Aspergillus genus. The Aspergillus genus consists of common molds found throughout the environment within soil and water, on vegetation, in fecal matter, on decomposing matter, and suspended in the air. Species within this genus often grow quickly and can sporulate within a few days of germination. A combination of characteristics unique to A. niger makes the microbe invaluable to the production of many acids, proteins and bioactive compounds. Characteristics including extensive metabolic diversity, high production yield, secretion capability, and the ability to conduct post-translational modifications are responsible for A. niger's robust production of secondary metabolites. A. niger's capability to withstand extremely acidic conditions makes it especially important to the industrial production of citric acid.

<i>Aspergillus flavus</i> Species of fungus

Aspergillus flavus is a saprotrophic and pathogenic fungus with a cosmopolitan distribution. It is best known for its colonization of cereal grains, legumes, and tree nuts. Postharvest rot typically develops during harvest, storage, and/or transit. Its specific name flavus derives from the Latin meaning yellow, a reference to the frequently observed colour of the spores. A. flavus infections can occur while hosts are still in the field (preharvest), but often show no symptoms (dormancy) until postharvest storage or transport. In addition to causing preharvest and postharvest infections, many strains produce significant quantities of toxic compounds known as mycotoxins, which, when consumed, are toxic to mammals. A. flavus is also an opportunistic human and animal pathogen, causing aspergillosis in immunocompromised individuals.

<i>Aspergillus fumigatus</i> Species of fungus

Aspergillus fumigatus is a species of fungus in the genus Aspergillus, and is one of the most common Aspergillus species to cause disease in individuals with an immunodeficiency.

Heterothallic species have sexes that reside in different individuals. The term is applied particularly to distinguish heterothallic fungi, which require two compatible partners to produce sexual spores, from homothallic ones, which are capable of sexual reproduction from a single organism.

<i>Aspergillus oryzae</i> Filamentous fungus

Aspergillus oryzae, also known as kōji mold, is a filamentous fungus used in East Asia to saccharify rice, sweet potato, and barley in the making of alcoholic beverages such as sake and shōchū, and also to ferment soybeans for making soy sauce and miso. However, in the production of fermented foods of soybeans such as soy sauce and miso, Aspergillus sojae is sometimes used instead of A. oryzae. Incidentally, in China and Korea, the fungi used for fermented foods for a long time in the production of traditional alcoholic beverages were not A. oryzae but fungi belonging to Rhizopus and Mucor. A. oryzae is also used for the production of rice vinegars. Barley kōji (麦麹) or rice kōji (米麹) are made by fermenting the grains with A. oryzae hyphae.

<i>Aspergillus nidulans</i> Species of fungus

Aspergillus nidulans is one of many species of filamentous fungi in the phylum Ascomycota. It has been an important research organism for studying eukaryotic cell biology for over 50 years, being used to study a wide range of subjects including recombination, DNA repair, mutation, cell cycle control, tubulin, chromatin, nucleokinesis, pathogenesis, metabolism, and experimental evolution. It is one of the few species in its genus able to form sexual spores through meiosis, allowing crossing of strains in the laboratory. A. nidulans is a homothallic fungus, meaning it is able to self-fertilize and form fruiting bodies in the absence of a mating partner. It has septate hyphae with a woolly colony texture and white mycelia. The green colour of wild-type colonies is due to pigmentation of the spores, while mutations in the pigmentation pathway can produce other spore colours.

<span class="mw-page-title-main">Mating in fungi</span> Combination of genetic material between compatible mating types

Mating in fungi is a complex process governed by mating types. Research on fungal mating has focused on several model species with different behaviour. Not all fungi reproduce sexually and many that do are isogamous; thus, for many members of the fungal kingdom, the terms "male" and "female" do not apply. Homothallic species are able to mate with themselves, while in heterothallic species only isolates of opposite mating types can mate.

<span class="mw-page-title-main">Aspergillosis</span> Medical condition

Aspergillosis is a fungal infection of usually the lungs, caused by the genus Aspergillus, a common mould that is breathed in frequently from the air around, but does not usually affect most people. It generally occurs in people with lung diseases such as asthma, cystic fibrosis or tuberculosis, or those who have had a stem cell or organ transplant, and those who cannot fight infection because of medications they take such as steroids and some cancer treatments. Rarely, it can affect skin.

<span class="mw-page-title-main">Gliotoxin</span> Chemical compound

Gliotoxin is a sulfur-containing mycotoxin that belongs to a class of naturally occurring 2,5-diketopiperazines produced by several species of fungi, especially those of marine origin. It is the most prominent member of the epipolythiopiperazines, a large class of natural products featuring a diketopiperazine with di- or polysulfide linkage. These highly bioactive compounds have been the subject of numerous studies aimed at new therapeutics. Gliotoxin was originally isolated from Gliocladium fimbriatum, and was named accordingly. It is an epipolythiodioxopiperazine metabolite.

<i>Aspergillus terreus</i> Species of fungus

Aspergillus terreus, also known as Aspergillus terrestris, is a fungus (mold) found worldwide in soil. Although thought to be strictly asexual until recently, A. terreus is now known to be capable of sexual reproduction. This saprotrophic fungus is prevalent in warmer climates such as tropical and subtropical regions. Aside from being located in soil, A. terreus has also been found in habitats such as decomposing vegetation and dust. A. terreus is commonly used in industry to produce important organic acids, such as itaconic acid and cis-aconitic acid, as well as enzymes, like xylanase. It was also the initial source for the drug mevinolin (lovastatin), a drug for lowering serum cholesterol.

Pathogenic fungi are fungi that cause disease in humans or other organisms. Approximately 300 fungi are known to be pathogenic to humans. Markedly more fungi are known to be pathogenic to plant life than those of the animal kingdom. The study of fungi pathogenic to humans is called "medical mycology". Although fungi are eukaryotic, many pathogenic fungi are microorganisms. The study of fungi and other organisms pathogenic to plants is called plant pathology.

Homothallic refers to the possession, within a single organism, of the resources to reproduce sexually; i.e., having male and female reproductive structures on the same thallus. The opposite sexual functions are performed by different cells of a single mycelium.

<span class="mw-page-title-main">Fungi imperfecti</span> Fungal classification based on asexual characters when sexual reproduction is unidentified

The fungi imperfecti or imperfect fungi, are fungi which do not fit into the commonly established taxonomic classifications of fungi that are based on biological species concepts or morphological characteristics of sexual structures because their sexual form of reproduction has never been observed. They are known as imperfect fungi because only their asexual and vegetative phases are known. They have asexual form of reproduction, meaning that these fungi produce their spores asexually, in the process called sporogenesis.

<span class="mw-page-title-main">Chronic pulmonary aspergillosis</span> Fungal infection

Chronic pulmonary aspergillosis is a long-term fungal infection caused by members of the genus Aspergillus—most commonly Aspergillusfumigatus. The term describes several disease presentations with considerable overlap, ranging from an aspergilloma—a clump of Aspergillus mold in the lungs—through to a subacute, invasive form known as chronic necrotizing pulmonary aspergillosis which affects people whose immune system is weakened. Many people affected by chronic pulmonary aspergillosis have an underlying lung disease, most commonly tuberculosis, allergic bronchopulmonary aspergillosis, asthma, or lung cancer.

Myceliophthora thermophila is an ascomycete fungus that grows optimally at 45–50 °C (113–122 °F). It efficiently degrades cellulose and is of interest in the production of biofuels. The genome has recently been sequenced, revealing the full range of enzymes this organism uses for the degradation of plant cell wall material.

<i>Aspergillus candidus</i> Species of fungus

Aspergillus candidus is a white-spored species of fungus in the genus Aspergillus. Despite its lack of pigmentation, it is closely related to the most darkly-pigmented aspergilli in the Aspergillus niger group. It is a common soil fungus worldwide and is known as a contaminant of a wide array of materials from the indoor environment to foods and products. It is an uncommon agent of onychomycosis and aspergillosis. The species epithet candidus (L.) refers to the white pigmentation of colonies of this fungus. It is from the Candidi section. The fungi in the Candidi section are known for their white spores. It has been isolated from wheat flour, djambee, and wheat grain.

Aspergillus wentii is an asexual, filamentous, endosymbiotic fungus belonging to the mold genus, Aspergillus. It is a common soil fungus with a cosmopolitan distribution, although it is primarily found in subtropical regions. Found on a variety of organic materials, A. wentii is known to colonize corn, cereals, moist grains, peanuts and other ground nut crops. It is also used in the manufacture of biodiesel from lipids and is known for its ability to produce enzymes used in the food industry.

Aspergillus giganteus is a species of fungus in the genus Aspergillus that grows as a mold. It was first described in 1901 by Wehmer, and is one of six Aspergillus species from the Clavati section of the subgenus Fumigati. Its closest taxonomic relatives are Aspergillus rhizopodus and Aspergillus longivescia.


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Further reading