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'''''Aspergillus niger''''' is a mold classified within the ''Nigri'' section of the ''[[Aspergillus]]'' genus.<ref name=":4">{{cite journal | vauthors = Ellena V, Seekles SJ, Vignolle GA, Ram AF, Steiger MG | title = Genome sequencing of the neotype strain CBS 554.65 reveals the MAT1-2 locus of Aspergillus niger | journal = BMC Genomics | volume = 22 | issue = 1 | pages = 679 | date = September 2021 | pmid = 34548025 | pmc = 8454179 | doi = 10.1186/s12864-021-07990-8 }}</ref> 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.<ref name=":5">{{Citation | vauthors = Curtis L |title=Aspergillus |date=2020 |url= http://connection.ebscohost.com/c/articles/94416780 |encyclopedia=Salem Press Encyclopedia of Health |publisher=Salem Press |access-date=2022-10-18}}</ref> Species within this genus often grow quickly and can sporulate within a few days of [[germination]].<ref name=":5" /> 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 modification|post-translational modifications]] are responsible for ''A. niger's'' robust production of secondary metabolites.<ref name=":8">{{cite journal | vauthors = Kurt T, Marbà-Ardébol AM, Turan Z, Neubauer P, Junne S, Meyer V | title = Rocking Aspergillus: morphology-controlled cultivation of Aspergillus niger in a wave-mixed bioreactor for the production of secondary metabolites | journal = Microbial Cell Factories | volume = 17 | issue = 1 | pages = 128 | date = August 2018 | pmid = 30129427 | doi = 10.1186/s12934-018-0975-y | s2cid = 52053640 }}</ref> ''A. niger's'' capability to withstand extremely acidic conditions makes it especially important to the industrial production of [[citric acid]]. <ref name=":4" /><ref>{{cite journal | vauthors = Behera BC | title = Citric acid from <i>Aspergillus niger</i>: a comprehensive overview | journal = Critical Reviews in Microbiology | volume = 46 | issue = 6 | pages = 727–749 | date = November 2020 | pmid = 33044884 | doi = 10.1080/1040841X.2020.1828815 | s2cid = 222319687 }}</ref>
'''''Aspergillus niger''''' is a mold classified within the ''Nigri'' section of the ''[[Aspergillus]]'' genus.<ref name=":4">{{cite journal | vauthors = Ellena V, Seekles SJ, Vignolle GA, Ram AF, Steiger MG | title = Genome sequencing of the neotype strain CBS 554.65 reveals the MAT1-2 locus of Aspergillus niger | journal = BMC Genomics | volume = 22 | issue = 1 | pages = 679 | date = September 2021 | pmid = 34548025 | pmc = 8454179 | doi = 10.1186/s12864-021-07990-8 }}</ref> 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.<ref name=":5">{{Citation | vauthors = Curtis L |title=Aspergillus |date=2020 |url= http://connection.ebscohost.com/c/articles/94416780 |encyclopedia=Salem Press Encyclopedia of Health |publisher=Salem Press |access-date=2022-10-18}}</ref> Species within this genus often grow quickly and can sporulate within a few days of [[germination]].<ref name=":5" /> 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 modification|post-translational modifications]] are responsible for ''A. niger's'' robust production of secondary metabolites.<ref name=":8">{{cite journal | vauthors = Kurt T, Marbà-Ardébol AM, Turan Z, Neubauer P, Junne S, Meyer V | title = Rocking Aspergillus: morphology-controlled cultivation of Aspergillus niger in a wave-mixed bioreactor for the production of secondary metabolites | journal = Microbial Cell Factories | volume = 17 | issue = 1 | pages = 128 | date = August 2018 | pmid = 30129427 | doi = 10.1186/s12934-018-0975-y | pmc = 6102829 | s2cid = 52053640 }}</ref> ''A. niger's'' capability to withstand extremely acidic conditions makes it especially important to the industrial production of [[citric acid]]. <ref name=":4" /><ref>{{cite journal | vauthors = Behera BC | title = Citric acid from <i>Aspergillus niger</i>: a comprehensive overview | journal = Critical Reviews in Microbiology | volume = 46 | issue = 6 | pages = 727–749 | date = November 2020 | pmid = 33044884 | doi = 10.1080/1040841X.2020.1828815 | s2cid = 222319687 }}</ref>


''A. niger'' causes a disease known as "black mold" on certain fruits and vegetables such as grapes, apricots, onions, and peanuts, and is a common contaminant of food. It is ubiquitous in [[soil]] and is commonly found in indoor environments, where its black colonies can be confused with those of ''[[Stachybotrys]]'' (species of which have also been called "black mold").<ref name=Samson_2001>{{cite book |vauthors=Samson RA, Houbraken J, Summerbell RC, Flannigan B, Miller JD | chapter = Common and important species of fungi and actinomycetes in indoor environments|title=Microorganisms in Home and Indoor Work Environments | pages = 287–292 | publisher = CRC| year = 2001 | isbn= 978-0415268004}}</ref> ''A. niger'' is classified as [[Generally Recognized as Safe]] (GRAS) by the US [[Food and Drug Administration]] for use in food production,<ref>{{Citation |last=Singh |first=Nikita |title=GRAS Fungi: A New Horizon in Safer Food Product |date=2021 |url=https://doi.org/10.1007/978-3-030-64406-2_3 |work=Fungi in Sustainable Food Production |pages=27–37 |editor-last=Dai |editor-first=Xiaofeng |place=Cham |publisher=Springer International Publishing |language=en |doi=10.1007/978-3-030-64406-2_3 |isbn=978-3-030-64406-2 |access-date=2022-11-16 |last2=Gaur |first2=Smriti |editor2-last=Sharma |editor2-first=Minaxi |editor3-last=Chen |editor3-first=Jieyin}}</ref> although the microbe is capable of producing toxins that affect human health.<ref name=fris/>
''A. niger'' causes a disease known as "black mold" on certain fruits and vegetables such as grapes, apricots, onions, and peanuts, and is a common contaminant of food. It is ubiquitous in [[soil]] and is commonly found in indoor environments, where its black colonies can be confused with those of ''[[Stachybotrys]]'' (species of which have also been called "black mold").<ref name=Samson_2001>{{cite book |vauthors=Samson RA, Houbraken J, Summerbell RC, Flannigan B, Miller JD | chapter = Common and important species of fungi and actinomycetes in indoor environments|title=Microorganisms in Home and Indoor Work Environments | pages = 287–292 | publisher = CRC| year = 2001 | isbn= 978-0415268004}}</ref> ''A. niger'' is classified as [[Generally Recognized as Safe]] (GRAS) by the US [[Food and Drug Administration]] for use in food production,<ref>{{Citation |last1=Singh |first1=Nikita |title=GRAS Fungi: A New Horizon in Safer Food Product |date=2021 |url=https://doi.org/10.1007/978-3-030-64406-2_3 |work=Fungi in Sustainable Food Production |pages=27–37 |editor-last=Dai |editor-first=Xiaofeng |place=Cham |publisher=Springer International Publishing |language=en |doi=10.1007/978-3-030-64406-2_3 |isbn=978-3-030-64406-2 |access-date=2022-11-16 |last2=Gaur |first2=Smriti |s2cid=234175577 |editor2-last=Sharma |editor2-first=Minaxi |editor3-last=Chen |editor3-first=Jieyin}}</ref> although the microbe is capable of producing toxins that affect human health.<ref name=fris/>


== Taxonomy ==
== Taxonomy ==
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[[File:Aspergillus niger 2.jpg|alt=|thumb|''A. niger'' growing on [[potato dextrose agar]]]]
[[File:Aspergillus niger 2.jpg|alt=|thumb|''A. niger'' growing on [[potato dextrose agar]]]]


''A. niger'' is a strict aerobe; therefore, it requires oxygen to grow.<ref name=":10">{{cite journal |vauthors=Costa CP, Gonçalves Silva D, Rudnitskaya A, Almeida A, Rocha SM |date=June 2016 |title=Shedding light on Aspergillus niger volatile exometabolome |journal=Scientific Reports |volume=6 |issue=1 |pages=27441 |doi=10.1038/srep27441 |pmid=27264696}}</ref> The fungus can grow in a range of environmental conditions; it can grow at temperatures ranging from 6 to 47ºC.<ref name=":9">{{cite journal |display-authors=6 |vauthors=Semova N, Storms R, John T, Gaudet P, Ulycznyj P, Min XJ, Sun J, Butler G, Tsang A |date=February 2006 |title=Generation, annotation, and analysis of an extensive Aspergillus niger EST collection |journal=BMC Microbiology |volume=6 |issue=1 |pages=7 |doi=10.1186/1471-2180-6-7 |pmc=1434744 |pmid=16457709}}</ref> As a mesophile,<ref name=":14">{{Cite web |title=Aspergillus niger |url=https://www.inspq.qc.ca/es/node/484 |access-date=2022-11-17 |website=INSPQ |language=es}}</ref> its optimal temperature range is 35-37ºC.<ref name=":10" /> It can tolerate pH ranging from 1.5 to 9.8.<ref name=":9" /> ''A. niger'' is xerophilic, meaning it can grow and reproduce in environments with very little water. It can also can also grow in humid conditions even tolerating environments with 90-100% relative humidity.<ref name=":14" /> The fungus is most commonly grown on potato dextrose agar (PDA), but it can grow on many different types of growth media including Czapek-Dox agar, lignocellulose agar, and several others. {{fact|date=November 2022}}
''A. niger'' is a strict aerobe; therefore, it requires oxygen to grow.<ref name=":10">{{cite journal |vauthors=Costa CP, Gonçalves Silva D, Rudnitskaya A, Almeida A, Rocha SM |date=June 2016 |title=Shedding light on Aspergillus niger volatile exometabolome |journal=Scientific Reports |volume=6 |issue=1 |pages=27441 |doi=10.1038/srep27441 |pmid=27264696|pmc=4893740 |bibcode=2016NatSR...627441C }}</ref> The fungus can grow in a range of environmental conditions; it can grow at temperatures ranging from 6 to 47ºC.<ref name=":9">{{cite journal |display-authors=6 |vauthors=Semova N, Storms R, John T, Gaudet P, Ulycznyj P, Min XJ, Sun J, Butler G, Tsang A |date=February 2006 |title=Generation, annotation, and analysis of an extensive Aspergillus niger EST collection |journal=BMC Microbiology |volume=6 |issue=1 |pages=7 |doi=10.1186/1471-2180-6-7 |pmc=1434744 |pmid=16457709}}</ref> As a mesophile,<ref name=":14">{{Cite web |title=Aspergillus niger |url=https://www.inspq.qc.ca/es/node/484 |access-date=2022-11-17 |website=INSPQ |language=es}}</ref> its optimal temperature range is 35-37ºC.<ref name=":10" /> It can tolerate pH ranging from 1.5 to 9.8.<ref name=":9" /> ''A. niger'' is xerophilic, meaning it can grow and reproduce in environments with very little water. It can also can also grow in humid conditions even tolerating environments with 90-100% relative humidity.<ref name=":14" /> The fungus is most commonly grown on potato dextrose agar (PDA), but it can grow on many different types of growth media including Czapek-Dox agar, lignocellulose agar, and several others. {{fact|date=November 2022}}


== Genome ==
== Genome ==
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''Aspergillus niger'' is cultured to facilitate the industrial production of many substances.<ref>{{cite journal | vauthors = Cairns TC, Nai C, Meyer V | title = How a fungus shapes biotechnology: 100 years of <i>Aspergillus niger</i> research | journal = Fungal Biology and Biotechnology | volume = 5 | pages = 13 | date = 2018 | pmid = 29850025 | pmc = 5966904 | doi = 10.1186/s40694-018-0054-5 }}</ref> Various strains of ''A. niger'' are used in the industrial preparation of [[citric acid]] (E330) and [[gluconic acid]] (E574); therefore, they have been deemed acceptable for daily intake by the [[World Health Organization]].<ref>{{cite journal | vauthors = Max B, Salgado JM, Rodríguez N, Cortés S, Converti A, Domínguez JM | title = Biotechnological production of citric acid | journal = Brazilian Journal of Microbiology | volume = 41 | issue = 4 | pages = 862–875 | date = October 2010 | pmid = 24031566 | pmc = 3769771 | doi = 10.1590/S1517-83822010000400005 }}</ref> ''A. niger'' fermentation is "generally recognized as safe" ([[GRAS]]) by the United States [[Food and Drug Administration (United States)|Food and Drug Administration]] under the [[Federal Food, Drug, and Cosmetic Act]].<ref>{{cite web|url=http://www.cfsan.fda.gov/~rdb/opa-gras.html |title=Inventory of GRAS Notices: Summary of all GRAS Notices |date=2008-10-22 |website=US FDA/CFSAN |access-date=2008-10-31 |archive-url=https://web.archive.org/web/20081011163648/http://www.cfsan.fda.gov/~rdb/opa-gras.html |archive-date=11 October 2008 |url-status=live }}</ref> ''A. niger'' is also being considered as a potential new source of natural food grade pigments.<ref>{{cite journal | vauthors = Toma MA, Nazir KH, Mahmud MM, Mishra P, Ali MK, Kabir A, Shahid MA, Siddique MP, Alim MA | display-authors = 6 | title = Isolation and Identification of Natural Colorant Producing Soil-Borne <i>Aspergillus niger</i> from Bangladesh and Extraction of the Pigment | journal = Foods | volume = 10 | issue = 6 | pages = 1280 | date = June 2021 | pmid = 34205202 | pmc = 8227025 | doi = 10.3390/foods10061280 | doi-access = free }}</ref>
''Aspergillus niger'' is cultured to facilitate the industrial production of many substances.<ref>{{cite journal | vauthors = Cairns TC, Nai C, Meyer V | title = How a fungus shapes biotechnology: 100 years of <i>Aspergillus niger</i> research | journal = Fungal Biology and Biotechnology | volume = 5 | pages = 13 | date = 2018 | pmid = 29850025 | pmc = 5966904 | doi = 10.1186/s40694-018-0054-5 }}</ref> Various strains of ''A. niger'' are used in the industrial preparation of [[citric acid]] (E330) and [[gluconic acid]] (E574); therefore, they have been deemed acceptable for daily intake by the [[World Health Organization]].<ref>{{cite journal | vauthors = Max B, Salgado JM, Rodríguez N, Cortés S, Converti A, Domínguez JM | title = Biotechnological production of citric acid | journal = Brazilian Journal of Microbiology | volume = 41 | issue = 4 | pages = 862–875 | date = October 2010 | pmid = 24031566 | pmc = 3769771 | doi = 10.1590/S1517-83822010000400005 }}</ref> ''A. niger'' fermentation is "generally recognized as safe" ([[GRAS]]) by the United States [[Food and Drug Administration (United States)|Food and Drug Administration]] under the [[Federal Food, Drug, and Cosmetic Act]].<ref>{{cite web|url=http://www.cfsan.fda.gov/~rdb/opa-gras.html |title=Inventory of GRAS Notices: Summary of all GRAS Notices |date=2008-10-22 |website=US FDA/CFSAN |access-date=2008-10-31 |archive-url=https://web.archive.org/web/20081011163648/http://www.cfsan.fda.gov/~rdb/opa-gras.html |archive-date=11 October 2008 |url-status=live }}</ref> ''A. niger'' is also being considered as a potential new source of natural food grade pigments.<ref>{{cite journal | vauthors = Toma MA, Nazir KH, Mahmud MM, Mishra P, Ali MK, Kabir A, Shahid MA, Siddique MP, Alim MA | display-authors = 6 | title = Isolation and Identification of Natural Colorant Producing Soil-Borne <i>Aspergillus niger</i> from Bangladesh and Extraction of the Pigment | journal = Foods | volume = 10 | issue = 6 | pages = 1280 | date = June 2021 | pmid = 34205202 | pmc = 8227025 | doi = 10.3390/foods10061280 | doi-access = free }}</ref>


The production of citric acid (CA) is achieved by growing strains of ''A. niger'' in a nutrient rich medium that includes high concentrations of sugar and mineral salts and an acidic pH of 2.5-3.5.<ref>{{cite journal | vauthors = Papagianni M | title = Advances in citric acid fermentation by Aspergillus niger: biochemical aspects, membrane transport and modeling | journal = Biotechnology Advances | volume = 25 | issue = 3 | pages = 244–263 | date = 2007-05-01 | pmid = 17337335 | doi = 10.1016/j.biotechadv.2007.01.002 }}</ref> Many microorganisms produce CA, but ''Aspergillus niger'' produces more than 1 million metric tons of CA annually via a fungal fermentation process.<ref name=":0">{{cite journal | vauthors = Baker SE | title = Aspergillus niger genomics: past, present and into the future | journal = Medical Mycology | volume = 44 | issue = 1 | pages = S17-S21 | date = September 2006 | pmid = 17050415 | doi = 10.1080/13693780600921037 | s2cid = 50631 }}</ref> CA is in high demand for applications such as the control of microorganism growth, food and beverage flavor enhancement, acidity manipulation, pharmaceuticals, etc.<ref>{{Cite book | vauthors = Sackett D |title=Citric Acid: Occurrence, Biochemistry, Applications and Processing |publisher=Nova Science Publishers Inc. |year=2014 |isbn=978-1-63117-237-3 |pages=119}}</ref>
The production of citric acid (CA) is achieved by growing strains of ''A. niger'' in a nutrient rich medium that includes high concentrations of sugar and mineral salts and an acidic pH of 2.5-3.5.<ref>{{cite journal | vauthors = Papagianni M | title = Advances in citric acid fermentation by Aspergillus niger: biochemical aspects, membrane transport and modeling | journal = Biotechnology Advances | volume = 25 | issue = 3 | pages = 244–263 | date = 2007-05-01 | pmid = 17337335 | doi = 10.1016/j.biotechadv.2007.01.002 }}</ref> Many microorganisms produce CA, but ''Aspergillus niger'' produces more than 1 million metric tons of CA annually via a fungal fermentation process.<ref name=":0">{{cite journal | vauthors = Baker SE | title = Aspergillus niger genomics: past, present and into the future | journal = Medical Mycology | volume = 44 | issue = 1 | pages = S17–S21 | date = September 2006 | pmid = 17050415 | doi = 10.1080/13693780600921037 | s2cid = 50631 }}</ref> CA is in high demand for applications such as the control of microorganism growth, food and beverage flavor enhancement, acidity manipulation, pharmaceuticals, etc.<ref>{{Cite book | vauthors = Sackett D |title=Citric Acid: Occurrence, Biochemistry, Applications and Processing |publisher=Nova Science Publishers Inc. |year=2014 |isbn=978-1-63117-237-3 |pages=119}}</ref>


''A. niger'' produces many useful [[enzyme]]s for the catabolism of biopolymers in order to obtain nutrients from its environment.<ref name=":3">{{cite journal | vauthors = Pel HJ, de Winde JH, Archer DB, Dyer PS, Hofmann G, Schaap PJ, Turner G, de Vries RP, Albang R, Albermann K, Andersen MR, Bendtsen JD, Benen JA, van den Berg M, Breestraat S, Caddick MX, Contreras R, Cornell M, Coutinho PM, Danchin EG, Debets AJ, Dekker P, van Dijck PW, van Dijk A, Dijkhuizen L, Driessen AJ, d'Enfert C, Geysens S, Goosen C, Groot GS, de Groot PW, Guillemette T, Henrissat B, Herweijer M, van den Hombergh JP, van den Hondel CA, van der Heijden RT, van der Kaaij RM, Klis FM, Kools HJ, Kubicek CP, van Kuyk PA, Lauber J, Lu X, van der Maarel MJ, Meulenberg R, Menke H, Mortimer MA, Nielsen J, Oliver SG, Olsthoorn M, Pal K, van Peij NN, Ram AF, Rinas U, Roubos JA, Sagt CM, Schmoll M, Sun J, Ussery D, Varga J, Vervecken W, van de Vondervoort PJ, Wedler H, Wösten HA, Zeng AP, van Ooyen AJ, Visser J, Stam H | display-authors = 6 | title = Genome sequencing and analysis of the versatile cell factory Aspergillus niger CBS 513.88 | journal = Nature Biotechnology | volume = 25 | issue = 2 | pages = 221–231 | date = February 2007 | pmid = 17259976 | doi = 10.1038/nbt1282 | s2cid = 19831590 }}</ref> The production of specific enzymes can be increased for industrial purposes.<ref>{{cite journal | vauthors = Ong LG, Abd-Aziz S, Noraini S, Karim MI, Hassan MA | title = Enzyme production and profile by Aspergillus niger during solid substrate fermentation using palm kernel cake as substrate | journal = Applied Biochemistry and Biotechnology | volume = 118 | issue = 1-3 | pages = 73–79 | date = 2004 | pmid = 15304740 | doi = 10.1385/ABAB:118:1-3:073 | s2cid = 19063403 }}</ref><ref name=":3" /> For example, ''A. niger'' [[glucoamylase]] ({{PDBe-KB|P69328}}) is used in the production of [[high-fructose corn syrup]] and [[pectinase]]s ([[Glycoside hydrolase family 28|GH28]]) are used in cider and [[wine clarification]]. [[Alpha-galactosidase]] ([[Glycoside hydrolase family 27|GH27]]), an enzyme that breaks down certain complex sugars, is a component of [[Beano (dietary supplement)|Beano]] and several other products that decrease [[flatulence]].<ref>{{cite journal | vauthors = Di Stefano M, Miceli E, Gotti S, Missanelli A, Mazzocchi S, Corazza GR | title = The effect of oral alpha-galactosidase on intestinal gas production and gas-related symptoms | journal = Digestive Diseases and Sciences | volume = 52 | issue = 1 | pages = 78–83 | date = January 2007 | pmid = 17151807 | doi = 10.1007/s10620-006-9296-9 | s2cid = 35435660 }}</ref> Another use for ''A. niger'' within the biotechnology industry is in the production of magnetic isotope-containing variants of biological [[macromolecule]]s for [[NMR spectroscopy|NMR]] analysis.<ref>{{cite journal | vauthors = MacKenzie DA, Spencer JA, Le Gal-Coëffet MF, Archer DB | title = Efficient production from Aspergillus niger of a heterologous protein and an individual protein domain, heavy isotope-labelled, for structure-function analysis | journal = Journal of Biotechnology | volume = 46 | issue = 2 | pages = 85–93 | date = April 1996 | pmid = 8672288 | doi = 10.1016/0168-1656(95)00179-4 }}</ref> ''Aspergillus niger'' is also cultured for the extraction of the enzyme, [[glucose oxidase]] ({{PDBe-KB|P13006}}), used in the design of glucose [[biosensor]]s, due to its high affinity for [[glucose|β-D-glucose]].<ref>{{cite journal | vauthors = Staiano M, Bazzicalupo P, Rossi M, D'Auria S | title = Glucose biosensors as models for the development of advanced protein-based biosensors | journal = Molecular BioSystems | volume = 1 | issue = 5-6 | pages = 354–362 | date = December 2005 | pmid = 16881003 | doi = 10.1039/b513385h }}</ref><ref name="ghoshdastider">{{cite journal|vauthors=Ghoshdastider U, Wu R, Trzaskowski B, Mlynarczyk K, Miszta P, Gurusaran M, Viswanathan S, Renugopalakrishnan V, Filipek S|s2cid=55816037|year=2015|title=Nano-Encapsulation of Glucose Oxidase Dimer by Graphene|journal=RSC Advances|volume=5|issue=18|pages=13570–78|doi=10.1039/C4RA16852F}}</ref>
''A. niger'' produces many useful [[enzyme]]s for the catabolism of biopolymers in order to obtain nutrients from its environment.<ref name=":3">{{cite journal | vauthors = Pel HJ, de Winde JH, Archer DB, Dyer PS, Hofmann G, Schaap PJ, Turner G, de Vries RP, Albang R, Albermann K, Andersen MR, Bendtsen JD, Benen JA, van den Berg M, Breestraat S, Caddick MX, Contreras R, Cornell M, Coutinho PM, Danchin EG, Debets AJ, Dekker P, van Dijck PW, van Dijk A, Dijkhuizen L, Driessen AJ, d'Enfert C, Geysens S, Goosen C, Groot GS, de Groot PW, Guillemette T, Henrissat B, Herweijer M, van den Hombergh JP, van den Hondel CA, van der Heijden RT, van der Kaaij RM, Klis FM, Kools HJ, Kubicek CP, van Kuyk PA, Lauber J, Lu X, van der Maarel MJ, Meulenberg R, Menke H, Mortimer MA, Nielsen J, Oliver SG, Olsthoorn M, Pal K, van Peij NN, Ram AF, Rinas U, Roubos JA, Sagt CM, Schmoll M, Sun J, Ussery D, Varga J, Vervecken W, van de Vondervoort PJ, Wedler H, Wösten HA, Zeng AP, van Ooyen AJ, Visser J, Stam H | display-authors = 6 | title = Genome sequencing and analysis of the versatile cell factory Aspergillus niger CBS 513.88 | journal = Nature Biotechnology | volume = 25 | issue = 2 | pages = 221–231 | date = February 2007 | pmid = 17259976 | doi = 10.1038/nbt1282 | s2cid = 19831590 }}</ref> The production of specific enzymes can be increased for industrial purposes.<ref>{{cite journal | vauthors = Ong LG, Abd-Aziz S, Noraini S, Karim MI, Hassan MA | title = Enzyme production and profile by Aspergillus niger during solid substrate fermentation using palm kernel cake as substrate | journal = Applied Biochemistry and Biotechnology | volume = 118 | issue = 1–3 | pages = 73–79 | date = 2004 | pmid = 15304740 | doi = 10.1385/ABAB:118:1-3:073 | s2cid = 19063403 }}</ref><ref name=":3" /> For example, ''A. niger'' [[glucoamylase]] ({{PDBe-KB|P69328}}) is used in the production of [[high-fructose corn syrup]] and [[pectinase]]s ([[Glycoside hydrolase family 28|GH28]]) are used in cider and [[wine clarification]]. [[Alpha-galactosidase]] ([[Glycoside hydrolase family 27|GH27]]), an enzyme that breaks down certain complex sugars, is a component of [[Beano (dietary supplement)|Beano]] and several other products that decrease [[flatulence]].<ref>{{cite journal | vauthors = Di Stefano M, Miceli E, Gotti S, Missanelli A, Mazzocchi S, Corazza GR | title = The effect of oral alpha-galactosidase on intestinal gas production and gas-related symptoms | journal = Digestive Diseases and Sciences | volume = 52 | issue = 1 | pages = 78–83 | date = January 2007 | pmid = 17151807 | doi = 10.1007/s10620-006-9296-9 | s2cid = 35435660 }}</ref> Another use for ''A. niger'' within the biotechnology industry is in the production of magnetic isotope-containing variants of biological [[macromolecule]]s for [[NMR spectroscopy|NMR]] analysis.<ref>{{cite journal | vauthors = MacKenzie DA, Spencer JA, Le Gal-Coëffet MF, Archer DB | title = Efficient production from Aspergillus niger of a heterologous protein and an individual protein domain, heavy isotope-labelled, for structure-function analysis | journal = Journal of Biotechnology | volume = 46 | issue = 2 | pages = 85–93 | date = April 1996 | pmid = 8672288 | doi = 10.1016/0168-1656(95)00179-4 }}</ref> ''Aspergillus niger'' is also cultured for the extraction of the enzyme, [[glucose oxidase]] ({{PDBe-KB|P13006}}), used in the design of glucose [[biosensor]]s, due to its high affinity for [[glucose|β-D-glucose]].<ref>{{cite journal | vauthors = Staiano M, Bazzicalupo P, Rossi M, D'Auria S | title = Glucose biosensors as models for the development of advanced protein-based biosensors | journal = Molecular BioSystems | volume = 1 | issue = 5–6 | pages = 354–362 | date = December 2005 | pmid = 16881003 | doi = 10.1039/b513385h }}</ref><ref name="ghoshdastider">{{cite journal|vauthors=Ghoshdastider U, Wu R, Trzaskowski B, Mlynarczyk K, Miszta P, Gurusaran M, Viswanathan S, Renugopalakrishnan V, Filipek S|s2cid=55816037|year=2015|title=Nano-Encapsulation of Glucose Oxidase Dimer by Graphene|journal=RSC Advances|volume=5|issue=18|pages=13570–78|doi=10.1039/C4RA16852F}}</ref>


In the food industry, ''A. niger'' is also cultured to isolate the enzyme fructosyltransferase to produce [[Fructooligosaccharide|fructooligosaccharides (FOS)]].<ref name=":1">{{cite journal | vauthors = Mao S, Liu Y, Yang J, Ma X, Zeng F, Zhang Z, Wang S, Han H, Qin HM, Lu F | display-authors = 6 | title = Cloning, expression and characterization of a novel fructosyltransferase from <i>Aspergillus niger</i> and its application in the synthesis of fructooligosaccharides | journal = RSC Advances | volume = 9 | issue = 41 | pages = 23856–23863 | date = July 2019 | pmid = 35530578 | pmc = 9069702 | doi = 10.1039/C9RA02520K | bibcode = 2019RSCAd...923856M }}</ref> FOS are used to manufacture low-calorie and [[functional food]]s due to FOS characteristic ability to slow growth of pathogenic microorganisms in the intestines.<ref name=":1" /><ref name=":2">{{Cite journal | vauthors = Guo W, Yang H, Qiang S, Fan Y, Shen W, Chen X |date=2016-07-01 |title=Overproduction, purification, and property analysis of an extracellular recombinant fructosyltransferase |url=https://doi.org/10.1007/s00217-015-2620-x |journal=European Food Research and Technology |language=en |volume=242 |issue=7 |pages=1159–1168 |doi=10.1007/s00217-015-2620-x |s2cid=86927574 |issn=1438-2385}}</ref> These foods have [[Prebiotic (nutrition)|prebiotic]] fiber among other health promoting properties. ''A. niger'' is not the only organism to produce the enzyme fructosyltransferase, but it has been found to produce the enzyme at rates conducive to industrial production.<ref name=":1" /><ref name=":2" /> A specific use of A. niger within the food industry is its capability to product enzymes like carbohydrase and cellulase, which are commonly used in the seafood industry for removing the bellies of clams during processing and removing the tough external skin of shrimp from their edible internal tissue.<ref>{{Cite web |title=CFR - Code of Federal Regulations Title 21 |url=https://www.accessdata.fda.gov/scripts/cdrh/cfdocs/cfcfr/CFRSearch.cfm?fr=173.120 |access-date=2022-11-16 |website=www.accessdata.fda.gov}}</ref>
In the food industry, ''A. niger'' is also cultured to isolate the enzyme fructosyltransferase to produce [[Fructooligosaccharide|fructooligosaccharides (FOS)]].<ref name=":1">{{cite journal | vauthors = Mao S, Liu Y, Yang J, Ma X, Zeng F, Zhang Z, Wang S, Han H, Qin HM, Lu F | display-authors = 6 | title = Cloning, expression and characterization of a novel fructosyltransferase from <i>Aspergillus niger</i> and its application in the synthesis of fructooligosaccharides | journal = RSC Advances | volume = 9 | issue = 41 | pages = 23856–23863 | date = July 2019 | pmid = 35530578 | pmc = 9069702 | doi = 10.1039/C9RA02520K | bibcode = 2019RSCAd...923856M }}</ref> FOS are used to manufacture low-calorie and [[functional food]]s due to FOS characteristic ability to slow growth of pathogenic microorganisms in the intestines.<ref name=":1" /><ref name=":2">{{Cite journal | vauthors = Guo W, Yang H, Qiang S, Fan Y, Shen W, Chen X |date=2016-07-01 |title=Overproduction, purification, and property analysis of an extracellular recombinant fructosyltransferase |url=https://doi.org/10.1007/s00217-015-2620-x |journal=European Food Research and Technology |language=en |volume=242 |issue=7 |pages=1159–1168 |doi=10.1007/s00217-015-2620-x |s2cid=86927574 |issn=1438-2385}}</ref> These foods have [[Prebiotic (nutrition)|prebiotic]] fiber among other health promoting properties. ''A. niger'' is not the only organism to produce the enzyme fructosyltransferase, but it has been found to produce the enzyme at rates conducive to industrial production.<ref name=":1" /><ref name=":2" /> A specific use of A. niger within the food industry is its capability to product enzymes like carbohydrase and cellulase, which are commonly used in the seafood industry for removing the bellies of clams during processing and removing the tough external skin of shrimp from their edible internal tissue.<ref>{{Cite web |title=CFR - Code of Federal Regulations Title 21 |url=https://www.accessdata.fda.gov/scripts/cdrh/cfdocs/cfcfr/CFRSearch.cfm?fr=173.120 |access-date=2022-11-16 |website=www.accessdata.fda.gov}}</ref>
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== Toxicity ==
== Toxicity ==
''A. niger'' produces a wide variety of secondary metabolites,<ref name="fris">{{cite journal | vauthors = Frisvad JC, Møller LL, Larsen TO, Kumar R, Arnau J | title = Safety of the fungal workhorses of industrial biotechnology: update on the mycotoxin and secondary metabolite potential of Aspergillus niger, Aspergillus oryzae, and Trichoderma reesei | journal = Applied Microbiology and Biotechnology | volume = 102 | issue = 22 | pages = 9481–9515 | date = November 2018 | pmid = 30293194 | pmc = 6208954 | doi = 10.1007/s00253-018-9354-1 }}</ref> some of which are [[mycotoxin]]s called [[ochratoxin]]s,<ref name="Abarca_2004">{{cite journal | vauthors = Abarca ML, Bragulat MR, Castellá G, Cabañes FJ | title = Ochratoxin A production by strains of Aspergillus niger var. niger | journal = Applied and Environmental Microbiology | volume = 60 | issue = 7 | pages = 2650–2652 | date = July 1994 | pmid = 8074536 | pmc = 201698 | doi = 10.1128/AEM.60.7.2650-2652.1994 | bibcode = 1994ApEnM..60.2650A }}</ref> such as [[ochratoxin A]].<ref name="Samson_2001" /><ref>{{cite journal | vauthors = Schuster E, Dunn-Coleman N, Frisvad JC, Van Dijck PW | title = On the safety of Aspergillus niger--a review | journal = Applied Microbiology and Biotechnology | volume = 59 | issue = 4-5 | pages = 426–435 | date = August 2002 | pmid = 12172605 | doi = 10.1007/s00253-002-1032-6 | s2cid = 26113037 }}</ref> Contamination by filamentous fungi, such as ''A. niger,'' occurs frequently in grapes and grape based products resulting in contamination by ochratoxin A (OTA). OTA, a clinically relevant mycotoxin, can accumulate in human tissue and cause a variety of serious health conditions.<ref>{{cite journal | vauthors = Freire L, Guerreiro TM, Pia AK, Lima EO, Oliveira DN, Melo CF, Catharino RR, Sant'Ana AS | display-authors = 6 | title = A quantitative study on growth variability and production of ochratoxin A and its derivatives by A. carbonarius and A. niger in grape-based medium | journal = Scientific Reports | volume = 8 | issue = 1 | pages = 14573 | date = October 2018 | pmid = 30275502 | doi = 10.1038/s41598-018-32907-z }}</ref> Potential consequences of OTA poisoning include kidney damage, kidney failure and cancer but the United States Food and Drug Administration (FDA) has not set maximum permissible levels of OTA in food unlike the EU that set maximum permissible levels in a variety of food products. <ref>{{Cite journal |last=Ráduly |first=Zsolt |last2=Szabó |first2=László |last3=Madar |first3=Anett |last4=Pócsi |first4=István |last5=Csernoch |first5=László |date=2020 |title=Toxicological and Medical Aspects of Aspergillus-Derived Mycotoxins Entering the Feed and Food Chain |url=https://www.frontiersin.org/articles/10.3389/fmicb.2019.02908 |journal=Frontiers in Microbiology |volume=10 |doi=10.3389/fmicb.2019.02908/full |issn=1664-302X}}</ref>
''A. niger'' produces a wide variety of secondary metabolites,<ref name="fris">{{cite journal | vauthors = Frisvad JC, Møller LL, Larsen TO, Kumar R, Arnau J | title = Safety of the fungal workhorses of industrial biotechnology: update on the mycotoxin and secondary metabolite potential of Aspergillus niger, Aspergillus oryzae, and Trichoderma reesei | journal = Applied Microbiology and Biotechnology | volume = 102 | issue = 22 | pages = 9481–9515 | date = November 2018 | pmid = 30293194 | pmc = 6208954 | doi = 10.1007/s00253-018-9354-1 }}</ref> some of which are [[mycotoxin]]s called [[ochratoxin]]s,<ref name="Abarca_2004">{{cite journal | vauthors = Abarca ML, Bragulat MR, Castellá G, Cabañes FJ | title = Ochratoxin A production by strains of Aspergillus niger var. niger | journal = Applied and Environmental Microbiology | volume = 60 | issue = 7 | pages = 2650–2652 | date = July 1994 | pmid = 8074536 | pmc = 201698 | doi = 10.1128/AEM.60.7.2650-2652.1994 | bibcode = 1994ApEnM..60.2650A }}</ref> such as [[ochratoxin A]].<ref name="Samson_2001" /><ref>{{cite journal | vauthors = Schuster E, Dunn-Coleman N, Frisvad JC, Van Dijck PW | title = On the safety of Aspergillus niger--a review | journal = Applied Microbiology and Biotechnology | volume = 59 | issue = 4–5 | pages = 426–435 | date = August 2002 | pmid = 12172605 | doi = 10.1007/s00253-002-1032-6 | s2cid = 26113037 }}</ref> Contamination by filamentous fungi, such as ''A. niger,'' occurs frequently in grapes and grape based products resulting in contamination by ochratoxin A (OTA). OTA, a clinically relevant mycotoxin, can accumulate in human tissue and cause a variety of serious health conditions.<ref>{{cite journal | vauthors = Freire L, Guerreiro TM, Pia AK, Lima EO, Oliveira DN, Melo CF, Catharino RR, Sant'Ana AS | display-authors = 6 | title = A quantitative study on growth variability and production of ochratoxin A and its derivatives by A. carbonarius and A. niger in grape-based medium | journal = Scientific Reports | volume = 8 | issue = 1 | pages = 14573 | date = October 2018 | pmid = 30275502 | doi = 10.1038/s41598-018-32907-z | pmc = 6167359 | bibcode = 2018NatSR...814573F }}</ref> Potential consequences of OTA poisoning include kidney damage, kidney failure and cancer but the United States Food and Drug Administration (FDA) has not set maximum permissible levels of OTA in food unlike the EU that set maximum permissible levels in a variety of food products. <ref>{{Cite journal |last1=Ráduly |first1=Zsolt |last2=Szabó |first2=László |last3=Madar |first3=Anett |last4=Pócsi |first4=István |last5=Csernoch |first5=László |date=2020 |title=Toxicological and Medical Aspects of Aspergillus-Derived Mycotoxins Entering the Feed and Food Chain |journal=Frontiers in Microbiology |volume=10 |page=2908 |doi=10.3389/fmicb.2019.02908 |pmid=31998250 |pmc=6962185 |issn=1664-302X|doi-access=free }}</ref>


== Pathogenicity ==
== Pathogenicity ==
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=== Plant pathogen ===
=== Plant pathogen ===
''Aspergillus niger'' can cause black mold infections in certain fruits and vegetables like peanuts, grapes, and onions leading to the fungus being a common food contaminant. This filamentous [[Ascomycota|ascomycete]] has a tolerance to changes in pH, humidity, and heat, thriving in a temperature range 15°C–53°C.<ref>{{Cite journal | vauthors = Dania VO, Fajemisin AO, Azuh VO |date=2021-12-14 |title=Morphological and molecular characterization of Aspergillus niger causing postharvest rot of white yam (Dioscorea rotundata Poir) |url=https://doi.org/10.1080/03235408.2021.1983365 |journal=Archives of Phytopathology and Plant Protection |volume=54 |issue=19-20 |pages=2356–2374 |doi=10.1080/03235408.2021.1983365 |issn=0323-5408}}</ref> These characteristics make infections of ''A. niger'' a common cause of post-harvest decay in fruits and vegetables, which can lead to significant economic loss in the food industry.<ref name=":11">{{Cite journal | vauthors = Tawfik E, Alqurashi M, Aloufi S, Alyamani A, Baz L, Fayad E |date= January 2022 |title=Characterization of Mutant Aspergillus niger and the Impact on Certain Plants |journal=Sustainability |language=en |volume=14 |issue=3 |pages=1936 |doi=10.3390/su14031936 }}</ref> A. niger infection in plants can cause a reduction in seed germination, seedling emergence, root elongation, and shoot elongation, causing the plant to perish before maturation.<ref name=":11" /> Specifically, ''Aspergillus niger'' causes sooty mold on onions and ornamental plants.
''Aspergillus niger'' can cause black mold infections in certain fruits and vegetables like peanuts, grapes, and onions leading to the fungus being a common food contaminant. This filamentous [[Ascomycota|ascomycete]] has a tolerance to changes in pH, humidity, and heat, thriving in a temperature range 15°C–53°C.<ref>{{Cite journal | vauthors = Dania VO, Fajemisin AO, Azuh VO |date=2021-12-14 |title=Morphological and molecular characterization of Aspergillus niger causing postharvest rot of white yam (Dioscorea rotundata Poir) |url=https://doi.org/10.1080/03235408.2021.1983365 |journal=Archives of Phytopathology and Plant Protection |volume=54 |issue=19–20 |pages=2356–2374 |doi=10.1080/03235408.2021.1983365 |s2cid=244606795 |issn=0323-5408}}</ref> These characteristics make infections of ''A. niger'' a common cause of post-harvest decay in fruits and vegetables, which can lead to significant economic loss in the food industry.<ref name=":11">{{Cite journal | vauthors = Tawfik E, Alqurashi M, Aloufi S, Alyamani A, Baz L, Fayad E |date= January 2022 |title=Characterization of Mutant Aspergillus niger and the Impact on Certain Plants |journal=Sustainability |language=en |volume=14 |issue=3 |pages=1936 |doi=10.3390/su14031936 |doi-access= free }}</ref> A. niger infection in plants can cause a reduction in seed germination, seedling emergence, root elongation, and shoot elongation, causing the plant to perish before maturation.<ref name=":11" /> Specifically, ''Aspergillus niger'' causes sooty mold on onions and ornamental plants.


=== Human pathogen ===
=== Human pathogen ===
''A. niger'' is pathogenic. [[Aspergillosis]] is a fungal infection caused by spores of indoor and outdoor ''Aspergillus'' mold species.<ref name=":6">{{Cite web |date=2021-05-10 |title=Aspergillosis {{!}} Types of Fungal Diseases {{!}} Fungal Diseases {{!}} CDC |url=https://www.cdc.gov/fungal/diseases/aspergillosis/index.html |access-date=2022-10-26 |website=www.cdc.gov |language=en-us}}</ref> Due to the ubiquitous nature of A. niger, its spores are commonly inhaled by humans from their surrounding environment.<ref>{{Cite web |date=2022-07-11 |title=Information for Healthcare Professionals {{!}} Aspergillosis {{!}} Types of Fungal Diseases {{!}} Fungal Diseases {{!}} CDC |url=https://www.cdc.gov/fungal/diseases/aspergillosis/health-professionals.html |access-date=2022-10-26 |website=www.cdc.gov |language=en-us}}</ref> Aspergillosis infection customarily occurs in people with compromised immune systems or pre-existing lung conditions like [[asthma]] and [[cystic fibrosis]].<ref name=":6" /> Types of aspergillosis include allergic bronchopulmonary aspergillosis (ABPA), allergic aspergillus sinusitis, azole-resistant aspergillus fumigatus, cutaneous (skin) aspergillosis, and chronic pulmonary aspergillosis.<ref name=":6" /> Out of the approximated 180 species of ''aspergillus'' molds, roughly 40 species have been found to cause health concern in [[Immunodeficiency|immunocompromised]] humans.<ref name=":6" /> Aspergillosis is particularly frequent among [[horticulture|horticultural]] workers who often inhale [[peat]] dust, which can be rich in ''Aspergillus niger'' spores. The fungus has also been found in ancient Egyptian mummies and can be inhaled when they are disturbed.<ref>Handwerk, Brian (May 6, 2005) [http://news.nationalgeographic.com/news/2005/05/0506_050506_mummycurse.html Egypt's "King Tut Curse" Caused by Tomb Toxins?]. National Geographic.</ref> [[Otomycosis]], which is a superficial fungal infection of the ear canal, is another disorder that can be caused by overgrowth of ''Aspergillus'' molds like ''A. niger.''<ref name=":7">{{cite journal | vauthors = Javidnia J, Ghotbi Z, Ghojoghi A, Solhjoo K, Alshahni MM, Jeddi SA, Ahmadi B, Nouripour-Sisakht S, Ansari S, Shokoohi G | display-authors = 6 | title = Otomycosis in the South of Iran with a High Prevalence of Tympanic Membrane Perforation: A Hospital-Based Study | journal = Mycopathologia | volume = 187 | issue = 2-3 | pages = 225–233 | date = June 2022 | pmid = 35347533 | doi = 10.1007/s11046-022-00626-9 | s2cid = 247776123 }}</ref> Otomycosis caused by ''A. niger'' is frequently associated with mechanical damage of the ear canal's external skin barrier and often presents itself in patients living in tropical climates''.''<ref name=":7" /><ref>{{cite journal | vauthors = Schuster E, Dunn-Coleman N, Frisvad JC, Van Dijck PW | title = On the safety of Aspergillus niger--a review | journal = Applied Microbiology and Biotechnology | volume = 59 | issue = 4-5 | pages = 426–435 | date = August 2002 | pmid = 12172605 | doi = 10.1007/s00253-002-1032-6 | s2cid = 26113037 }}</ref> ''A. niger'' is rarely reported to cause [[pneumonia]] compared to other ''Aspergillus'' species, such as ''Aspergillus flavus, Aspergillus fumigatus'', and ''Aspergillus terreus''.<ref>{{cite journal | vauthors = Person AK, Chudgar SM, Norton BL, Tong BC, Stout JE | title = Aspergillus niger: an unusual cause of invasive pulmonary aspergillosis | journal = Journal of Medical Microbiology | volume = 59 | issue = Pt 7 | pages = 834–838 | date = July 2010 | pmid = 20299503 | pmc = 3052473 | doi = 10.1099/jmm.0.018309-0 }}</ref>
''A. niger'' is pathogenic. [[Aspergillosis]] is a fungal infection caused by spores of indoor and outdoor ''Aspergillus'' mold species.<ref name=":6">{{Cite web |date=2021-05-10 |title=Aspergillosis {{!}} Types of Fungal Diseases {{!}} Fungal Diseases {{!}} CDC |url=https://www.cdc.gov/fungal/diseases/aspergillosis/index.html |access-date=2022-10-26 |website=www.cdc.gov |language=en-us}}</ref> Due to the ubiquitous nature of A. niger, its spores are commonly inhaled by humans from their surrounding environment.<ref>{{Cite web |date=2022-07-11 |title=Information for Healthcare Professionals {{!}} Aspergillosis {{!}} Types of Fungal Diseases {{!}} Fungal Diseases {{!}} CDC |url=https://www.cdc.gov/fungal/diseases/aspergillosis/health-professionals.html |access-date=2022-10-26 |website=www.cdc.gov |language=en-us}}</ref> Aspergillosis infection customarily occurs in people with compromised immune systems or pre-existing lung conditions like [[asthma]] and [[cystic fibrosis]].<ref name=":6" /> Types of aspergillosis include allergic bronchopulmonary aspergillosis (ABPA), allergic aspergillus sinusitis, azole-resistant aspergillus fumigatus, cutaneous (skin) aspergillosis, and chronic pulmonary aspergillosis.<ref name=":6" /> Out of the approximated 180 species of ''aspergillus'' molds, roughly 40 species have been found to cause health concern in [[Immunodeficiency|immunocompromised]] humans.<ref name=":6" /> Aspergillosis is particularly frequent among [[horticulture|horticultural]] workers who often inhale [[peat]] dust, which can be rich in ''Aspergillus niger'' spores. The fungus has also been found in ancient Egyptian mummies and can be inhaled when they are disturbed.<ref>Handwerk, Brian (May 6, 2005) [http://news.nationalgeographic.com/news/2005/05/0506_050506_mummycurse.html Egypt's "King Tut Curse" Caused by Tomb Toxins?]. National Geographic.</ref> [[Otomycosis]], which is a superficial fungal infection of the ear canal, is another disorder that can be caused by overgrowth of ''Aspergillus'' molds like ''A. niger.''<ref name=":7">{{cite journal | vauthors = Javidnia J, Ghotbi Z, Ghojoghi A, Solhjoo K, Alshahni MM, Jeddi SA, Ahmadi B, Nouripour-Sisakht S, Ansari S, Shokoohi G | display-authors = 6 | title = Otomycosis in the South of Iran with a High Prevalence of Tympanic Membrane Perforation: A Hospital-Based Study | journal = Mycopathologia | volume = 187 | issue = 2–3 | pages = 225–233 | date = June 2022 | pmid = 35347533 | doi = 10.1007/s11046-022-00626-9 | s2cid = 247776123 }}</ref> Otomycosis caused by ''A. niger'' is frequently associated with mechanical damage of the ear canal's external skin barrier and often presents itself in patients living in tropical climates''.''<ref name=":7" /><ref>{{cite journal | vauthors = Schuster E, Dunn-Coleman N, Frisvad JC, Van Dijck PW | title = On the safety of Aspergillus niger--a review | journal = Applied Microbiology and Biotechnology | volume = 59 | issue = 4–5 | pages = 426–435 | date = August 2002 | pmid = 12172605 | doi = 10.1007/s00253-002-1032-6 | s2cid = 26113037 }}</ref> ''A. niger'' is rarely reported to cause [[pneumonia]] compared to other ''Aspergillus'' species, such as ''Aspergillus flavus, Aspergillus fumigatus'', and ''Aspergillus terreus''.<ref>{{cite journal | vauthors = Person AK, Chudgar SM, Norton BL, Tong BC, Stout JE | title = Aspergillus niger: an unusual cause of invasive pulmonary aspergillosis | journal = Journal of Medical Microbiology | volume = 59 | issue = Pt 7 | pages = 834–838 | date = July 2010 | pmid = 20299503 | pmc = 3052473 | doi = 10.1099/jmm.0.018309-0 }}</ref>


==Gallery==
==Gallery==

Revision as of 07:25, 21 November 2022

Aspergillus niger
Photomicrograph showing the conidial head of Aspergillus niger
Details of the head
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Fungi
Division: Ascomycota
Class: Eurotiomycetes
Order: Eurotiales
Family: Aspergillaceae
Genus: Aspergillus
Species:
A. niger
Binomial name
Aspergillus niger
Synonyms

Aspergillus niger var. niger
Aspergillopsis nigra (Tiegh.) Speg.
Rhopalocystis nigra (Tiegh.) Grove
Sterigmatocystis nigra (Tiegh.) Sacc. (1877)

Aspergillus niger is a mold classified within the Nigri section of the Aspergillus genus.[1] 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.[2] Species within this genus often grow quickly and can sporulate within a few days of germination.[2] 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.[3] A. niger's capability to withstand extremely acidic conditions makes it especially important to the industrial production of citric acid. [1][4]

A. niger causes a disease known as "black mold" on certain fruits and vegetables such as grapes, apricots, onions, and peanuts, and is a common contaminant of food. It is ubiquitous in soil and is commonly found in indoor environments, where its black colonies can be confused with those of Stachybotrys (species of which have also been called "black mold").[5] A. niger is classified as Generally Recognized as Safe (GRAS) by the US Food and Drug Administration for use in food production,[6] although the microbe is capable of producing toxins that affect human health.[7]

Taxonomy

Aspergillus niger is included in Aspergillus subgenus Circumdati, section Nigri. The section Nigri includes 15 related black-spored species that may be confused with A. niger, including A. tubingensis, A. foetidus, A. carbonarius, and A. awamori.[8][9] In 2004, a number of morphologically similar species were described by Samson et al.[9]

In 2007, the strain of ATCC 16404 Aspergillus niger was reclassified as Aspergillus brasiliensis (refer to publication by Varga et al.[10]). This required an update to the U.S. Pharmacopoeia and the European Pharmacopoeia, which commonly use this strain throughout the pharmaceutical industry.

Cultivation

A. niger growing on potato dextrose agar

A. niger is a strict aerobe; therefore, it requires oxygen to grow.[11] The fungus can grow in a range of environmental conditions; it can grow at temperatures ranging from 6 to 47ºC.[12] As a mesophile,[13] its optimal temperature range is 35-37ºC.[11] It can tolerate pH ranging from 1.5 to 9.8.[12] A. niger is xerophilic, meaning it can grow and reproduce in environments with very little water. It can also can also grow in humid conditions even tolerating environments with 90-100% relative humidity.[13] The fungus is most commonly grown on potato dextrose agar (PDA), but it can grow on many different types of growth media including Czapek-Dox agar, lignocellulose agar, and several others. [citation needed]

Genome

Aspergillus niger has a genome consisting of roughly 34 megabases (Mb) organized into eight chromosomes.[14] The DNA contains 10,785 genes which are transcribed and translated into 10,593 proteins.[14]

Genomic information
NCBI genome ID429
Ploidyhaploid
Genome size34 Mb
Number of chromosomes8

Two strains of A. niger have been sequenced. Strain CBS 513.88 produces enzymes used in industrial applications while strain ATCC 1015 is the wildtype strain of ATCC 11414 used to produce industrial citric acid (CA).[15][16][17] The A. niger ATCC 1015 genome was sequenced by the Joint Genome Institute in a collaboration with other institutions.[18] Completed sequences have been used to uncover orthologous genes and pathways involved in fungal metabolism, specifically the catabolism of monosaccharides.[19] The ability of A. niger to change its metabolism depending on the carbon sources and other nutrients present in its environment has enabled the microorganism to survive and be found in almost all ecosystems. Further research is being done to study these mechanisms for all fungi using the complete sequenced genome of A. niger.[19]

Industrial uses

There are two ways in which Aspergillus niger can be grown for industrial purposes: solid state fermentation (SSF) and submerged fermentation (SmF).[20] SSF uses a solid substrate with nutrients and minimal moisture to grow microorganisms. Nutrients such as nitrogen and carbon come from agricultural byproducts such as wheat bran, sugar pulp, rice husks, and corn flour.[21] SSF gives better yield of microbe products and is more cost effective than SmF due to using agricultural byproducts.[22] SSF is predominantly used over SmF.[22] In SmF, microbes are grown in a liquid medium inside large aseptic fermentation vessels.[21][22] These vessels are expensive pieces of equipment that provide more water for growth and allow for tight control of environmental factors, such as temperature and pH, that affects microbial growth.[22]

Aspergillus niger is cultured to facilitate the industrial production of many substances.[23] Various strains of A. niger are used in the industrial preparation of citric acid (E330) and gluconic acid (E574); therefore, they have been deemed acceptable for daily intake by the World Health Organization.[24] A. niger fermentation is "generally recognized as safe" (GRAS) by the United States Food and Drug Administration under the Federal Food, Drug, and Cosmetic Act.[25] A. niger is also being considered as a potential new source of natural food grade pigments.[26]

The production of citric acid (CA) is achieved by growing strains of A. niger in a nutrient rich medium that includes high concentrations of sugar and mineral salts and an acidic pH of 2.5-3.5.[27] Many microorganisms produce CA, but Aspergillus niger produces more than 1 million metric tons of CA annually via a fungal fermentation process.[28] CA is in high demand for applications such as the control of microorganism growth, food and beverage flavor enhancement, acidity manipulation, pharmaceuticals, etc.[29]

A. niger produces many useful enzymes for the catabolism of biopolymers in order to obtain nutrients from its environment.[30] The production of specific enzymes can be increased for industrial purposes.[31][30] For example, A. niger glucoamylase (P69328) is used in the production of high-fructose corn syrup and pectinases (GH28) are used in cider and wine clarification. Alpha-galactosidase (GH27), an enzyme that breaks down certain complex sugars, is a component of Beano and several other products that decrease flatulence.[32] Another use for A. niger within the biotechnology industry is in the production of magnetic isotope-containing variants of biological macromolecules for NMR analysis.[33] Aspergillus niger is also cultured for the extraction of the enzyme, glucose oxidase (P13006), used in the design of glucose biosensors, due to its high affinity for β-D-glucose.[34][35]

In the food industry, A. niger is also cultured to isolate the enzyme fructosyltransferase to produce fructooligosaccharides (FOS).[36] FOS are used to manufacture low-calorie and functional foods due to FOS characteristic ability to slow growth of pathogenic microorganisms in the intestines.[36][37] These foods have prebiotic fiber among other health promoting properties. A. niger is not the only organism to produce the enzyme fructosyltransferase, but it has been found to produce the enzyme at rates conducive to industrial production.[36][37] A specific use of A. niger within the food industry is its capability to product enzymes like carbohydrase and cellulase, which are commonly used in the seafood industry for removing the bellies of clams during processing and removing the tough external skin of shrimp from their edible internal tissue.[38]

Aspergillus niger can grow in gold-mining solutions containing cyano-metal complexes with gold, silver, copper, iron, and zinc. The fungus also plays a role in the solubilization of heavy-metal sulfides.[39] A. niger has also been shown to remediate acid mine drainage through biosorption of copper and manganese.[40]

Toxicity

A. niger produces a wide variety of secondary metabolites,[7] some of which are mycotoxins called ochratoxins,[41] such as ochratoxin A.[5][42] Contamination by filamentous fungi, such as A. niger, occurs frequently in grapes and grape based products resulting in contamination by ochratoxin A (OTA). OTA, a clinically relevant mycotoxin, can accumulate in human tissue and cause a variety of serious health conditions.[43] Potential consequences of OTA poisoning include kidney damage, kidney failure and cancer but the United States Food and Drug Administration (FDA) has not set maximum permissible levels of OTA in food unlike the EU that set maximum permissible levels in a variety of food products. [44]

Pathogenicity

A. niger growing on onion

Plant pathogen

Aspergillus niger can cause black mold infections in certain fruits and vegetables like peanuts, grapes, and onions leading to the fungus being a common food contaminant. This filamentous ascomycete has a tolerance to changes in pH, humidity, and heat, thriving in a temperature range 15°C–53°C.[45] These characteristics make infections of A. niger a common cause of post-harvest decay in fruits and vegetables, which can lead to significant economic loss in the food industry.[46] A. niger infection in plants can cause a reduction in seed germination, seedling emergence, root elongation, and shoot elongation, causing the plant to perish before maturation.[46] Specifically, Aspergillus niger causes sooty mold on onions and ornamental plants.

Human pathogen

A. niger is pathogenic. Aspergillosis is a fungal infection caused by spores of indoor and outdoor Aspergillus mold species.[47] Due to the ubiquitous nature of A. niger, its spores are commonly inhaled by humans from their surrounding environment.[48] Aspergillosis infection customarily occurs in people with compromised immune systems or pre-existing lung conditions like asthma and cystic fibrosis.[47] Types of aspergillosis include allergic bronchopulmonary aspergillosis (ABPA), allergic aspergillus sinusitis, azole-resistant aspergillus fumigatus, cutaneous (skin) aspergillosis, and chronic pulmonary aspergillosis.[47] Out of the approximated 180 species of aspergillus molds, roughly 40 species have been found to cause health concern in immunocompromised humans.[47] Aspergillosis is particularly frequent among horticultural workers who often inhale peat dust, which can be rich in Aspergillus niger spores. The fungus has also been found in ancient Egyptian mummies and can be inhaled when they are disturbed.[49] Otomycosis, which is a superficial fungal infection of the ear canal, is another disorder that can be caused by overgrowth of Aspergillus molds like A. niger.[50] Otomycosis caused by A. niger is frequently associated with mechanical damage of the ear canal's external skin barrier and often presents itself in patients living in tropical climates.[50][51] A. niger is rarely reported to cause pneumonia compared to other Aspergillus species, such as Aspergillus flavus, Aspergillus fumigatus, and Aspergillus terreus.[52]

See also

References

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  2. ^ a b Curtis L (2020), "Aspergillus", Salem Press Encyclopedia of Health, Salem Press, retrieved 2022-10-18
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