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''Aspergillus'' () 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. 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. While some species of ''Aspergillus'' are known to cause fungal infections, others are of commercial importance.


Taxonomy


Species

''Aspergillus'' consists of 837 species of fungi.


Growth and distribution

''Aspergillus'' is defined as a group of
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 fungi—that is, fungi in an asexual state. Some of them, however, are known to have a teleomorph (sexual state) in the
Ascomycota 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 def ...
. With DNA evidence, all members of the genus ''Aspergillus'' are members of the phylum
Ascomycota 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 def ...
. 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
monosaccharide Monosaccharides (from Greek ''monos'': single, '' sacchar'': sugar), also called simple sugars, are the simplest forms of sugar and the most basic units (monomers) from which all carbohydrates are built. They are usually colorless, water-solub ...
s (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. 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, favouring warm and damp or humid areas such as bathrooms and around window frames. ''Aspergillus'' are found in millions of pillows.


Commercial importance

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. 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 ''Saccharomyces'' is a genus of fungi that includes many species of yeasts. ''Saccharomyces'' is from Greek σάκχαρον (sugar) and μύκης (fungus) and means ''sugar fungus''. Many members of this genus are considered very important in f ...
'', cannot ferment these starches. Therefore, ''koji'' mold such as ''
Aspergillus oryzae ''Aspergillus oryzae'', also known as , is a filamentous fungus (a mold) 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 m ...
'' is used to first break down the starches into simpler sugars. Members of the genus are also sources of
natural product A natural product is a natural compound or substance produced by a living organism—that is, found in nature. In the broadest sense, natural products include any substance produced by life. Natural products can also be prepared by chemical syn ...
s that can be used in the development of medications to treat human disease. ''Aspergillus'' spp. are known to produce anthraquinone which has commercial importance due to its antibacterial and antifungal properties. 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. ''A. niger'' is also commonly used for the production of native and foreign
enzymes Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrate (chemistry), substrates, and the enzyme converts the substrates into different molecule ...
, including glucose oxidase, lysozyme, and lactase. 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 A bioreactor refers to any manufactured device or system that supports a biologically active environment. In one case, a bioreactor is a vessel in which a chemical reaction, chemical process is carried out which involves organisms or biochemistry, ...
. 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.


Research

'' 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). ''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.


Sexual reproduction

Of the 250 species of aspergilli, about 64% have no known sexual state. However, many of these species likely have an as yet unidentified sexual stage. 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 Immunodeficiency, also known as immunocompromisation, is a state in which the immune system's ability to fight infectious diseases and cancer is compromised or entirely absent. Most cases are acquired ("secondary") due to extrinsic factors that a ...
humans. In 2009, ''A. fumigatus'' was shown to have a heterothallic, fully functional sexual cycle. Isolates of complementary mating types are required for
sex Sex is the trait that determines whether a sexually reproducing animal or plant produces male or female gametes. Male plants and animals produce smaller mobile gametes (spermatozoa, sperm, pollen), while females produce larger ones (ova, oft ...
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. '' 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. ''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. 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
eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
s. ''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. Among those ''Aspergillus'' species that exhibit a sexual cycle, the overwhelming majority in nature are homothallic (self-fertilizing). 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, again suggesting sex, in this case outcrossing sex, can be maintained even when little genetic variability is produced.


Genomics

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 The J. Craig Venter Institute (JCVI) is a non-profit genomics research institute founded by J. Craig Venter, Ph.D. in October 2006. The institute was the result of consolidating four organizations: the Center for the Advancement of G ...
, is currently spearheading a project on the ''A. flavus'' genome. ''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. 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. 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.


Pathogens

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 infection Neonatal infections are infections of the neonate (newborn) acquired during prenatal development or in the first four weeks of life (neonatal period). Neonatal infections may be contracted by mother to child transmission, in the birth canal dur ...
s. ''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).


Aspergillosis

Aspergillosis is the group of diseases caused by ''Aspergillus''. The most common species among paranasal sinus infections associated with aspergillosis is '' A. fumigatus''. 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 The lungs are the primary organs of the respiratory system in humans and most other animals, including some snails and a small number of fish. In mammals and most other vertebrates, two lungs are located near the backbone on either side of t ...
conditions are susceptible. In humans, the major forms of disease are: * Acute invasive aspergillosis, a form that grows into surrounding tissue, more common in those with weakened immune systems such as
AIDS Human immunodeficiency virus infection and acquired immunodeficiency syndrome (HIV/AIDS) is a spectrum of conditions caused by infection with the human immunodeficiency virus (HIV), a retrovirus. Following initial infection an individual m ...
or chemotherapy patients * Allergic bronchopulmonary aspergillosis, which affects patients with respiratory diseases such as asthma,
cystic fibrosis Cystic fibrosis (CF) is a rare genetic disorder that affects mostly the lungs, but also the pancreas, liver, kidneys, and intestine. Long-term issues include difficulty breathing and coughing up mucus as a result of frequent lung infections. O ...
, and sinusitis * Aspergilloma, a "fungus ball" that can form within cavities such as the lung * Disseminated invasive aspergillosis, an infection spread widely through the body 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. Aspergillosis of the air passages is also frequently reported in birds, and certain species of ''Aspergillus'' have been known to infect insects. Most people inhale ''Aspergillus'' into their lungs everyday. But it's generally only the
immuno-compromised Immunodeficiency, also known as immunocompromisation, is a state in which the immune system's ability to fight infectious diseases and cancer is compromised or entirely absent. Most cases are acquired ("secondary") due to extrinsic factors that a ...
who get sick with Aspergillosis.


See also

* List of'' Aspergillus'' species *
Mold health issues Mold health issues refer to the harmful health effects of molds ("moulds" in British English) and their mycotoxins. However, recent research has shown these adverse health effects are caused not exclusively by molds, but also other microbial agen ...
*
Sick building syndrome Sick building syndrome (SBS) is an unsubstantiated diagnosis where health problems are attributed to buildings. The cause of the health problems are unknown. Symptoms attributed to SBS may or may not be a direct result of inadequate cleaning or ...


References


Further reading

* * * *


External links


Aspergillus Comparative Database
Comparative genomic resource at the Broad Institute
Aspergillus Genome Resources (NIH)

''Aspergillus'' surveillance project at a large tertiary-care hospital.
(
PDF Portable Document Format (PDF), standardized as ISO 32000, is a file format developed by Adobe in 1992 to present documents, including text formatting and images, in a manner independent of application software, hardware, and operating systems. ...
).
Central Aspergillus Data Repository

FungiDB: An integrated functional genomics database for fungi and oomycetes

Mold and Mildew

The Aspergillus Genome Database

The Aspergillus/Aspergillosis Website
An encyclopedia of ''Aspergillus'' for patients, doctors and scientists
The Fungal Genetics Stock Center
{{Authority control Parasitic fungi Eurotiomycetes genera