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| {{short description|Species of fungus}} | |||
| {{user sandbox}} | |||
| {{Speciesbox | |||
| {{Taxobox | |||
| ⚫ | | |
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| | image = Aspergillus clavatus.jpg | | image = Aspergillus clavatus.jpg | ||
| | image_width = 240px | |||
| | image_caption = Conidial head of ''Aspergillus clavatus'' | | image_caption = Conidial head of ''Aspergillus clavatus'' | ||
| | |
| genus = Aspergillus | ||
| ⚫ | | species = clavatus | ||
| | divisio = ] | |||
| | authority = ] (1834) | |||
| | classis = ] | |||
| ⚫ | | synonyms = '''''Aspergillus pallidus''''' | ||
| | ordo = ] | |||
| | familia = ] | |||
| | genus = '']'' | |||
| | species = '''''A. clavatus''''' | |||
| | binomial = ''Aspergillus clavatus'' | |||
| | binomial_authority = ] (1834) | |||
| ⚫ | | synonyms = ''' |
||
| (Samson, 1979) | (Samson, 1979) | ||
| | synonyms_ref = | | synonyms_ref = | ||
| }} | }} | ||
| '''''Aspergillus clavatus''''' is a ] of fungus in the genus '']'' with ] dimensions 3–4.5 x 2.5–4.5 ]. It is found in ] and animal ]. The fungus was first ] scientifically in 1834 by the French mycologist ].<ref name=desmazieres1834/> | |||
| The fungus can produce the ] ], which may be associated with disease in humans and animals. This species is only occasionally ]ic. | |||
| Other sources have identified many species of ''Aspergillus'' as producing dry, hydrophobic ]s that are easily inhaled by humans and animals. Due to the small size of the spores, about 70% of spores of ''A. fumigatus'' are able to penetrate into the ] and primary ] and close to 1% into ]. Inhalation of spores of ''Aspergillus'' is a health risk. ''A. clavatus'' is allergenic, causing the occupational ] known as ]. | |||
| ==History and taxonomy== | ==History and taxonomy== | ||
| ''Aspergillus clavatus'' is a ] of '']'' and is characterized by elongated club-shaped vesicles, and blue-green uniseriate ].<ref name=howard2003 /> The ] was first described scientifically in 1834 by the French mycologist ].<ref name=desmazieres1834 /> It belongs to the ''Aspergillus'' section ''Clavati'', (formerly known as the ''Aspergillus clavatus'' group) recognized by ] and ] (1926), alongside two species, ''Aspergillus clavatus'' and '']''.<ref name=varga2007 /> In the succeeding years, four more species were discovered belonging to the ''Aspergillus section Clavati'', which included '']'', '']'', ''Neocarpenteles acanthosporus'' and '']''.<ref name=varga2007 /> Later, ''Aspergillus pallidus'' was concluded to be a white variant (]) of ''A. clavatus'' by Samson (1979), which was supported by the identical ] sequences of the two species.<ref name=samson2000 /> A sexual stage was described in 2018 with a ''Neocarpenteles'' teleomorph but under the one fungus-one name convention the original ''A. clavatus'' epithet was retained.<ref name=":0">{{Cite journal|last1=Ojeda-López|first1=M|last2=Chen W|last3=Eagle CE|last4=Gutiérrez G|last5=Jia WL|last6=Swilaiman SS|last7=Huang Z|last8=Park HS|last9=Yu JH|last10=Dyer PS|date=2018|editor9-last=Cánovas D|title=Evolution of asexual and sexual reproduction in the aspergilli|url=https://www.studiesinmycology.org/|journal=Studies in Mycology|volume=91|pages=37–59|doi=10.1016/j.simyco.2018.10.002|pmid=30425416|pmc=6231087}}</ref> | |||
| ==Growth and morphology== | ==Growth and morphology== | ||
| '' |
''Aspergillus clavatus'' undergoes rapid growth, resulting in the formation of a velvety and fairly dense felt that is observed to be bluish-grey green in colour.<ref name=onions1981 /> The emerging conidial heads are large and clavate when very young, quickly splitting into conspicuous and compact divergent columns.<ref name=raper1965 /> The conidia bearing conidiophores are generally coarse, smooth walled, uncoloured,<ref name=raper1965 /> hyaline and can grow to be very long.<ref name=onions1981 /> Elongated club-shaped vesicles<ref name=onions1981 /> clavate,<ref name=raper1965 /> and bear phialides (]: ]) over their entire-surface, contributing to its short and densely packed structure.<ref name=onions1981 /> The ] are usually found to be uniseriate, numerous and crowded.<ref name=raper1965 /> Conidia formed in them are elliptical, smooth and comparatively thick-walled.<ref name=raper1965 /> ''A. clavatus'' usually express conidiophores 1.5–3.00 mm in length, which arises from specialized and widened ] cells that eventually become the branching foot cells.<ref name=domsch1980 /> The conidia on ''A. clavatus'' has been measured up to 3.0 – 4.5 X 2.5 – 3.5 μm.<ref name=domsch1980 /> Cleistothecia are produced in crosses after approximately 4–10 weeks of incubation on suitable growth media at 25 °C. Cleistothecia are yellowish-brown (fawn) to dark brown in colour and range in diameter from 315-700 μm in diameter and have a relatively hard outer wall (peridium). At maturity the cleistothecia contain asci that themselves contain ascospores, which are clear, lenticular (with ridges evident) and between 6.0-7.0 μm in diameter.<ref name=":0" /> | ||
| ===Growth on Czapek's solution agar=== | |||
| '' |
''Aspergillus clavatus'' colonies grow rapidly on ], reaching 3.0–3.5 cm, in 10 days at 24–26 °C.<ref name=raper1965 /> Growth is usually plane or moderately furrowed, with occasional appearance of floccose strains. But generally, a comparatively thin surface layer of ] felt is observed, which produces a copious number of erect conidiophores.<ref name=raper1965 /> The reverse is usually uncoloured but becomes brown with passing time in some strains.<ref name=raper1965 /> While odor is not prominent in some strains, it can be extremely unpleasant in others.<ref name=raper1965 /> Large conidial heads extend from 300 to 400 μm by 150 to 200 μm when young. However, with time, they split into two or more divergent and compressed cordial chains reaching 1.00 mm portraying a colour consisting of artemisia green to slate olive.<ref name=raper1965 /> The observed conidiophores grow up to 1.5–3.00 mm in length with 20–30 μm in diameter. They slowly and ultimately enlarge at the apex into a clavate vesicle, which consists of a fertile area, 200 to 250 μm in length and 40–60 μm wide.<ref name=raper1965 /> | ||
| The sterigmata usually ranges from 2.5 to 3.5 μm by 2.0 to 3.0 μm at the base of the vesicle, to 7.0 or 8.0 and occasionally 10 μm to 2.5 to 3.0 μm at the apex.<ref name=raper1965 /> The conidia are comparatively thick-walled and measures 3.0 to 4.5 μm by 2.5 to 3.5 μm. While they can be larger in some strains, in others their appearance may be irregular.<ref name=raper1965 /> |
The sterigmata usually ranges from 2.5 to 3.5 μm by 2.0 to 3.0 μm at the base of the vesicle, to 7.0 or 8.0 and occasionally 10 μm to 2.5 to 3.0 μm at the apex.<ref name=raper1965 /> The conidia are comparatively thick-walled and measures 3.0 to 4.5 μm by 2.5 to 3.5 μm. While they can be larger in some strains, in others their appearance may be irregular.<ref name=raper1965 /> | ||
| ===Growth on malt extract agar=== | |||
| ] | |||
| On malt extract agar, the structural morphology of ''A. clavatus'' appears to be different than in |
On malt extract agar, the structural morphology of ''A. clavatus'' appears to be different than in Czapek's solution agar.<ref name=raper1965 /> The typical strains extracted from malt media contain less abundant conidial structures, which could be larger in size.<ref name=raper1965 /> In other (non-typical) strains, the conidial heads increase in number but decrease in size. The conidiophores range from 300 to 500 μm and bear loose, columnar heads. Typical strains may be resembled by strong and unpleasant odor whereas non-typical strains are characterized being odorless.<ref name=raper1965 /> The colonies arising from one ] on malt extract agar, consisted of 25X10^7 conidia after being observed for six days.<ref name=domsch1980 /> | ||
| ===Examination=== | |||
| The phialide development and conidium formation in ''A. clavatus'' has been examined using ].<ref name=domsch1980 /> And by using ], it was discovered that the first-formed conidium and phialide share a continuous wall.<ref name=domsch1980 /> Additionally recombination with an albino ] led to the production of ] conidial heads with mixed conidial colours.<ref name=domsch1980 /> A ] of 52. |
The phialide development and conidium formation in ''A. clavatus'' has been examined using ].<ref name=domsch1980 /> And by using ], it was discovered that the first-formed conidium and phialide share a continuous wall.<ref name=domsch1980 /> Additionally recombination with an albino ] led to the production of ] conidial heads with mixed conidial colours.<ref name=domsch1980 /> A ] of 52.5–55% was also detected upon DNA analysis.<ref name=domsch1980 /> And its soluble wall carbohydrates consist of ] and ].<ref name=domsch1980 /> | ||
| ==Physiology== | ==Physiology== | ||
| Light stimulates the elongation of conidiophores in ''A. clavatus.'' And the more favourable C sources include ], ], ] and especially ].<ref name=domsch1980 /> Substantial degree of ] synthesis occurs, whereas ] and ] are degraded.<ref name=domsch1980 /> ''A. clavatus'' also produces ], ], acid phosphodiesterase and ] when in liquid culture.<ref name=domsch1980 /> |
Light stimulates the elongation of conidiophores in ''A. clavatus.'' And the more favourable C sources include ], ], ] and especially ].<ref name=domsch1980 /> Substantial degree of ] synthesis occurs, whereas ] and ] are degraded.<ref name=domsch1980 /> ''A. clavatus'' also produces ], ], acid phosphodiesterase and ] when in liquid culture.<ref name=domsch1980 /> | ||
| ''A. clavatus'' has the properties to oxidize ] to ]. It can absorb and collect ] from fuel oil, incorporate ] and synthesize ], clavatol and ].<ref name=domsch1980 /> It is also responsible for the production of ] ] and ].<ref name=onions1981 /> And has extremely high capacity for alcohol fermentation.<ref name=harold1965 /> |
''A. clavatus'' has the properties to oxidize ] to ]. It can absorb and collect ] from fuel oil, incorporate ] and synthesize ], clavatol and ].<ref name=domsch1980 /> It is also responsible for the production of ] ] and ].<ref name=onions1981 /> And has extremely high capacity for alcohol fermentation.<ref name=harold1965 /> | ||
| When it comes to |
When it comes to genomics, bioinformatic analysis revealed that ''A. clavatus'' contains a full complement of identified euascomycete sex genes.<ref name=machida2010 /> A heterothallic sexual cycle involving outcrossing between ''MAT1-1'' and ''MAT1-2'' isolates was subsequently described ''.<ref name=":0"/> A. clavatus'' can also be a food source for ] and has been found to be parasitized by ''].''<ref name=domsch1980 /> | ||
| ==Habitat and ecology== | ==Habitat and ecology== | ||
| '' |
''Aspergillus clavatus'' is often described as a spoilage organism occurring on dung and in soil and can also grow in strong alkaline conditions.<ref name=onions1981 /> When it comes to geographical distribution, ''A. clavatus'' has been spotted in the tropical, subtropical and Mediterranean areas.<ref name=domsch1980 /> It has been accounted in low frequencies in the soils of India. And is also found in Bangladesh, Sri Lanka, Hong Kong, Jamaica, Brazil, Argentina, South Africa, the Ivory Coast, Egypt, Libya, Turkey, Greece, Italy, the United States of America, Japan, the USSR and Czechoslovakia.<ref name=domsch1980 /> It was tracked in rocks of a carst cave and stratigraphic core samples descending to 1200 m in Central Japan.<ref name=domsch1980 /> However, it is usually and solely collected from cultivated soils, including the ones that bear cotton, potatoes, sugar canes, legumes, paddy and '']''.<ref name=domsch1980 /> It has also been garnered from soil under burnt steppe vegetation, desert soils, the rhizospheres of banana, ground-nuts and wheat.<ref name=domsch1980 /> ''A. clavatus'' has also been detected in the ripe compost of municipal waste, and ] and NPK fertilizers are found to play an important role in its stimulation process.<ref name=domsch1980 /> | ||
| ''A. clavatus'' is also referred as a cosmopolitan fungus. Other than soil and dung, it can additionally be found in stored products with high levels of entrapped moisture. Such as stored cereals, rice, corn and millet.<ref name=varga2007 /> It has been further isolated from insects, especially from dead adult bees and |
''A. clavatus'' is also referred as a cosmopolitan fungus. Other than soil and dung, it can additionally be found in stored products with high levels of entrapped moisture. Such as stored cereals, rice, corn and millet.<ref name=varga2007 /> It has been further isolated from insects, especially from dead adult bees and honeycombs.<ref name=domsch1980 /> Moreover, it has been collected from the feathers and droppings from free-living birds.<ref name=domsch1980 /> ''A. clavatus'' is also common is decomposing materials.<ref name=raper1965 /> Their ability to resist strongly alkaline conditions, allows them to act as decomposition catalysts in situations where other fungus usually do not function.<ref name=raper1965 /> | ||
| ==Applications and medical uses== | ==Applications and medical uses== | ||
| ⚫ | Weisner in March 1942 first noted the production of an antibiotic by strains of ''A. clavatus'', and the active substance was known as ''clavatin.''<ref name=raper1965 /> Later the antibiotic was named ''clavacin'' in August 1942 by Waksman, Horning and Spencer. Clavacin is also known as patulin.<ref name=raper1965 /> Patulin is receiving significant attention in the world today because of its manifestations in apple juices.<ref name=machida2010 /> Clavacin was noted to be valuable in the treatment of common-cold and applies a fungistatic or fungicidal effect on certain dermatophytes.<ref name=raper1965 /> ''A. clavatus'' with '']'' in soil provided protection against damping of tomato seedlings, by decreasing the spreading of pathogens.<ref name=raper1965 /> Reversely, ''A. clavatus'' with the addition of glucose, increased the pathogenicity of '']'' to tomatoes.<ref name=raper1965 /> ''A. clavatus'' also produces the following: Cytochalasin E, Cytochalasin K, Tryptoquivaline, Nortryptoquivalone, Nortryptoquivaline, Deoxytryptoquivaline, Deoxynortryptoquivaline, Tryptoquivaline E, and Tryptoquivaline N.<ref name=jarvis2003 /> Furthermore, ''A. clavatus'' isolates produce ], which can help develop ] processes for cancer.<ref name=machida2010 /> ''A.clavatus'' has also been used in the formation of extracellular bionanoparticles from ] solutions. These ] display ] properties, which work against ] and ].<ref name=saravanan2010 /> | ||
| ⚫ | Weisner in March |
||
| ==Pathogenicity== | ==Pathogenicity== | ||
| ⚫ | ''Aspergillus clavatus'' is known as an agent of allergic ]<ref name=hoog2000 /> and has been implicated in multiple pulmonary infections.<ref name=hoog2000 /> | ||
| ⚫ | '' |
||
| It has also been labelled as an opportunistic fungus, as it is responsible for causing aspergillosis in compromised patients.<ref name=al-doory1980 /> | It has also been labelled as an opportunistic fungus, as it is responsible for causing aspergillosis in compromised patients.<ref name=al-doory1980 /> | ||
| ''A.clavatus'' can also cause neurotoxicosis in sheep and ].<ref name=hoog2000 /> In Scotland and elsewhere, ''A. clavatus'' is reported for causing the mould allergy |
''A. clavatus'' can also cause neurotoxicosis in sheep and ].<ref name=hoog2000 /> In Scotland and elsewhere, ''A. clavatus'' is reported for causing the mould allergy "malster's lung" otherwise "maltster's lung".<ref name=ainsworth1986 /><ref>J E Smith, 1994: ''Aspergillus'' (Biotechnology Handbooks 7), p. 226. New York: Springer Science+Business Media</ref> | ||
| Extrinsic allergic alveolitis (]) is also caused by ''Aspergillus clavatus'' with a Type 1 immune reaction. It is described as a true hypersensitivity pneumonia, which usually occurs among malt workers, including symptoms of fever, chills, cough and dyspnea. In severe cases, ] are used. |
Extrinsic allergic alveolitis (]) is also caused by ''Aspergillus clavatus'' with a Type 1 immune reaction. It is described as a true hypersensitivity pneumonia, which usually occurs among malt workers, including symptoms of fever, chills, cough and dyspnea. In severe cases, ] are used.<ref name=holmberg1989 /> | ||
| Microgranulomatous hypersensitivity pneumonitis, where interstitial granulomatous infiltration occurs, usually in malt workers, is caused by allergy to antigens of ''Aspergillus clavatus''.<ref name=watts1987 /> | Microgranulomatous hypersensitivity pneumonitis, where interstitial granulomatous infiltration occurs, usually in malt workers, is caused by allergy to antigens of ''Aspergillus clavatus''.<ref name=watts1987 /> | ||
| EAA is caused by allergy to Aspergillus conidia, usually in the non-atopic individual.<ref name=rippon1982 /> Such individuals are usually exposed to organic dust heavily packed with conidia and mycelial debris.<ref name=rippon1982 /> This condition involves the lung parenchyma.<ref name=rippon1982 /> | EAA is caused by allergy to Aspergillus conidia, usually in the non-atopic individual.<ref name=rippon1982 /> Such individuals are usually exposed to organic dust heavily packed with conidia and mycelial debris.<ref name=rippon1982 /> This condition involves the lung parenchyma.<ref name=rippon1982 /> | ||
| A strain of ''A. clavatus'' has also caused ] in calves.<ref name=raper1965 /> Spore walls of a sputum-derived isolate of ''Aspergillus clavatus'' were extracted and treated with ethanol following alkaline hydrolysis. And it yielded mutagens.<ref name=blyth1982 /> The extracts were given to unimmunised mice, causing lung reaction |
A strain of ''A. clavatus'' has also caused ] in calves.<ref name=raper1965 /> Spore walls of a sputum-derived isolate of ''Aspergillus clavatus'' were extracted and treated with ethanol following alkaline hydrolysis. And it yielded mutagens.<ref name=blyth1982 /> The extracts were given to unimmunised mice, causing lung reaction and leading to cases of pulmonary mycotoxicosis. A rising incidence of lung tumours were also observed.<ref name=blyth1982 /> | ||
| This study revealed that an isolate of ''A. clavatus'', which is able to convert highly toxic metabolites in bacterial and mammalian cells, will cause inflammatory response in the lungs of unimmunized mice.<ref name=blyth1982 /> | This study revealed that an isolate of ''A. clavatus'', which is able to convert highly toxic metabolites in bacterial and mammalian cells, will cause inflammatory response in the lungs of unimmunized mice.<ref name=blyth1982 /> | ||
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| {{Reflist|30em|refs= | {{Reflist|30em|refs= | ||
| ⚫ | <ref name=desmazieres1834>{{cite journal|last1=Desmazières|first1=JBHJ|title=Descriptions et figures de six hyphomycètes inédites à ajouter à la flore Française|journal=Annales des Sciences Naturelles Botanique|date=1834|volume=2|issue=2|pages=69–73|language=French}}</ref> | ||
| ⚫ | <ref name=howard2003>{{cite book|editor-last1=Howard|editor-first1=Dexter H.|title=Pathogenic fungi in humans and animals|url=https://archive.org/details/pathogenicfungih00howa_331|url-access=limited|date=2003|publisher=Dekker|location=New York |isbn=978-0-8247-0683-8|page=|edition=2.}}</ref> | ||
| ⚫ | <ref name=desmazieres1834>{{cite journal|last1=Desmazières|first1=JBHJ|title=Descriptions et figures de six hyphomycètes inédites à ajouter à la flore Française|journal=Annales des Sciences Naturelles Botanique|date=1834|volume=2|issue=2| |
||
| <ref name=varga2007>{{cite journal|last1=Varga|first1=J.|last2=Due|first2=M.|last3=Frisvad|first3=J.C.|last4=Samson|first4=R.A.|title=Taxonomic revision of Aspergillus section Clavati based on molecular, morphological and physiological data|journal=Studies in Mycology|date=2007|volume=59|pages=89–106|doi=10.3114/sim.2007.59.11|pmid=18490946|pmc=2275193}}</ref> | |||
| ⚫ | <ref name=howard2003>{{cite book|last1=Howard|first1= |
||
| <ref name= |
<ref name=samson2000>{{cite book|last1=Samson|first1=ed. by Robert A.|last2=Pitt|first2=John I.|title=Integration of modern taxonomic methods for penicillium and aspergillus classification|date=2000|publisher=Harwood Acad. Publ.|location=Amsterdam|isbn=978-90-5823-159-8}}</ref> | ||
| <ref name= |
<ref name=onions1981>{{cite book|last1=Onions|first1=A.H.S.|last2=Allsopp|first2=D.|last3=Eggins|first3=H.O.W.|title=Smith's introduction to industrial mycology|date=1981|publisher=Arnold|location=London, UK|isbn=978-0-7131-2811-6|edition=7th}}</ref> | ||
| <ref name= |
<ref name=raper1965>{{cite book|last1=Raper|first1=Kenneth B.|last2=Fennell|first2=Dorothy I.|title=The Genus Aspergillus|date=1965|publisher=The Williams and Wilkins Company|location=Baltimore|pages=137–146}}</ref> | ||
| <ref name= |
<ref name=domsch1980>{{cite book|last1=Domsch|first1=K.H.|last2=Anderson|first2=Traute-Heidi|last3=Gams|first3=W.|title=Compendium of Soil Fungi|date=1980|publisher=Academic Press|pages=86–88}}</ref> | ||
| <ref name=domsch1980>{{cite book|last1=Domsch|first1=K.H.|last2=Anderson|first2=Traute-Heidi|last3=Gams|first3=W.|title=Compendium of Soil Fungi|date=1980|publisher=Academic Press|pages=86-88}}</ref> | |||
| <ref name=machida2010>{{cite book|last1=Machida|first1=edited by Masayuki|last2=Gomi|first2=Katsuya|title=Aspergillus : molecular biology and genomics|date=2010|publisher=Caister Academic|location=Wymondham, Norfolk, UK|isbn=978-1-904455-53-0}}</ref> | <ref name=machida2010>{{cite book|last1=Machida|first1=edited by Masayuki|last2=Gomi|first2=Katsuya|title=Aspergillus : molecular biology and genomics|date=2010|publisher=Caister Academic|location=Wymondham, Norfolk, UK|isbn=978-1-904455-53-0}}</ref> | ||
| Line 87: | Line 82: | ||
| <ref name=harold1965>{{cite book|author1=Harold J. Blumenthal|editor1-last=Ainsworth|editor1-first=G.C.|editor2-last=Sussman|editor2-first=Alfred S.|title=THE FUNGI Volume 1 The Fungal Cell|date=1965|publisher=Academic Press|location=New York and London|page=251|chapter=10}}</ref> | <ref name=harold1965>{{cite book|author1=Harold J. Blumenthal|editor1-last=Ainsworth|editor1-first=G.C.|editor2-last=Sussman|editor2-first=Alfred S.|title=THE FUNGI Volume 1 The Fungal Cell|date=1965|publisher=Academic Press|location=New York and London|page=251|chapter=10}}</ref> | ||
| <ref name=jarvis2003>{{cite book|last1=Jarvis |
<ref name=jarvis2003>{{cite book|last1=Jarvis|first1=Richard J. Cole,... Milbra A. Schweikert,... Bruce B.|title=Handbook of secondary fungal metabolites|date=2003|publisher=Academic press|location=Amsterdam |isbn=978-0-12-179461-3}}</ref> | ||
| <ref name=saravanan2010>{{cite journal|last1=Saravanan|first1=M.|last2=Nanda|first2=Anima|title=Extracellular synthesis of silver bionanoparticles from Aspergillus clavatus and its antimicrobial activity against MRSA and MRSE|journal=Colloids and Surfaces B: Biointerfaces|date=June 2010|volume=77|issue=2|pages=214–218|doi=10.1016/j.colsurfb.2010.01.026}}</ref> | <ref name=saravanan2010>{{cite journal|last1=Saravanan|first1=M.|last2=Nanda|first2=Anima|title=Extracellular synthesis of silver bionanoparticles from Aspergillus clavatus and its antimicrobial activity against MRSA and MRSE|journal=Colloids and Surfaces B: Biointerfaces|date=June 2010|volume=77|issue=2|pages=214–218|doi=10.1016/j.colsurfb.2010.01.026|pmid=20189360}}</ref> | ||
| <ref name=hoog2000>{{cite book|last1=Hoog|first1=G. S. de|last2=Guarro|first2=J.|last3=Gene|first3=J.|last4=Figueras|first4=M. J.|title=Atlas of clinical fungi|date=2000|publisher=Centraalbureau voor Schimmelcultures |location=Utrecht|isbn=90-7035-143- |
<ref name=hoog2000>{{cite book|last1=Hoog|first1=G. S. de|last2=Guarro|first2=J.|last3=Gene|first3=J.|last4=Figueras|first4=M. J.|title=Atlas of clinical fungi|date=2000|publisher=Centraalbureau voor Schimmelcultures |location=Utrecht|isbn=978-90-7035-143-4|edition=2.}}</ref> | ||
| <ref name=ainsworth1986>{{cite book|last1=Ainsworth|first1=G.C.|title=Introduction to the history of medical and veterinary mycology|date=1986|publisher=Cambridge University Press|location=Cambridge|isbn=0-521-30715- |
<ref name=ainsworth1986>{{cite book|last1=Ainsworth|first1=G.C.|title=Introduction to the history of medical and veterinary mycology|date=1986|publisher=Cambridge University Press|location=Cambridge|isbn=978-0-521-30715-4}}</ref> | ||
| <ref name=al-doory1980>{{cite book|last1=Al-Doory|first1=Yousef|title=Laboratory Medical Mycology|date=1980|publisher=Lea and Febiger|location=Philadelphia}}</ref> |
<ref name=al-doory1980>{{cite book|last1=Al-Doory|first1=Yousef|title=Laboratory Medical Mycology|date=1980|publisher=Lea and Febiger|location=Philadelphia}}</ref> | ||
| <ref name=holmberg1989>{{cite book|last1=Holmberg|first1=Kenneth|last2=Meyer|first2=Richard D.|title=Diagnosis and Therapy of Systemic Fungal Infections|date=1989|publisher=Raven Press|location=New York}}</ref> | <ref name=holmberg1989>{{cite book|last1=Holmberg|first1=Kenneth|last2=Meyer|first2=Richard D.|title=Diagnosis and Therapy of Systemic Fungal Infections|date=1989|publisher=Raven Press|location=New York}}</ref> | ||
| <ref name=watts1987>{{cite book|last1=Watts|first1=Francis W. Chandler, John C.|title=Pathologic diagnosis of fungal infections|date=1987|publisher=ASCP Press|location=Chicago|isbn=0-89189-252- |
<ref name=watts1987>{{cite book|last1=Watts|first1=Francis W. Chandler, John C.|title=Pathologic diagnosis of fungal infections|date=1987|publisher=ASCP Press|location=Chicago|isbn=978-0-89189-252-6}}</ref> | ||
| <ref name=rippon1982>{{cite book|last1=Rippon|first1=John Willard|title=Medical mycology : the pathogenic fungi and the pathogenic actinomycetes|date=1982|publisher=Saunders|location=Philadelphia|isbn=0-7216-7586- |
<ref name=rippon1982>{{cite book|last1=Rippon|first1=John Willard|title=Medical mycology : the pathogenic fungi and the pathogenic actinomycetes|date=1982|publisher=Saunders|location=Philadelphia|isbn=978-0-7216-7586-2|edition=2nd|url-access=registration|url=https://archive.org/details/medicalmycologype2ripp}}</ref> | ||
| <ref name=blyth1982>{{cite journal| |
<ref name=blyth1982>{{cite journal |vauthors=Blyth W, Hardy JC |title=Mutagenic and tumourigenic properties of the spores of Aspergillus clavatus |journal=Br. J. Cancer |volume=45 |issue=1 |pages=105–17 |year=1982 |pmid=7059453 |pmc=2010971 |doi=10.1038/bjc.1982.13}}</ref> | ||
| }} | }} | ||
| {{Taxonbar|from=Q138618}} | |||
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Latest revision as of 00:13, 31 December 2024
Species of fungus
| Aspergillus clavatus | |
|---|---|
| |
| Conidial head of Aspergillus clavatus | |
Scientific classification 
| |
| Domain: | Eukaryota |
| Kingdom: | Fungi |
| Division: | Ascomycota |
| Class: | Eurotiomycetes |
| Order: | Eurotiales |
| Family: | Aspergillaceae |
| Genus: | Aspergillus |
| Species: | A. clavatus |
| Binomial name | |
| Aspergillus clavatus Desm. (1834) | |
| Synonyms | |
|
Aspergillus pallidus (Samson, 1979) | |
Aspergillus clavatus is a species of fungus in the genus Aspergillus with conidia dimensions 3–4.5 x 2.5–4.5 μm. It is found in soil and animal manure. The fungus was first described scientifically in 1834 by the French mycologist John Baptiste Henri Joseph Desmazières.
The fungus can produce the toxin patulin, which may be associated with disease in humans and animals. This species is only occasionally pathogenic.
Other sources have identified many species of Aspergillus as producing dry, hydrophobic spores that are easily inhaled by humans and animals. Due to the small size of the spores, about 70% of spores of A. fumigatus are able to penetrate into the trachea and primary bronchi and close to 1% into alveoli. Inhalation of spores of Aspergillus is a health risk. A. clavatus is allergenic, causing the occupational hypersensitivity pneumonitis known as malt-worker's lung.
History and taxonomy
Aspergillus clavatus is a species of Aspergillus and is characterized by elongated club-shaped vesicles, and blue-green uniseriate conidia. The fungus was first described scientifically in 1834 by the French mycologist John Baptiste Henri Joseph Desmazières. It belongs to the Aspergillus section Clavati, (formerly known as the Aspergillus clavatus group) recognized by Charles Thom and Margaret Church (1926), alongside two species, Aspergillus clavatus and Aspergillus giganteus. In the succeeding years, four more species were discovered belonging to the Aspergillus section Clavati, which included Aspergillus rhizopodus, Aspergillus longivesica, Neocarpenteles acanthosporus and Aspergillus clavatonanicus. Later, Aspergillus pallidus was concluded to be a white variant (synonym) of A. clavatus by Samson (1979), which was supported by the identical DNA sequences of the two species. A sexual stage was described in 2018 with a Neocarpenteles teleomorph but under the one fungus-one name convention the original A. clavatus epithet was retained.
Growth and morphology
Aspergillus clavatus undergoes rapid growth, resulting in the formation of a velvety and fairly dense felt that is observed to be bluish-grey green in colour. The emerging conidial heads are large and clavate when very young, quickly splitting into conspicuous and compact divergent columns. The conidia bearing conidiophores are generally coarse, smooth walled, uncoloured, hyaline and can grow to be very long. Elongated club-shaped vesicles clavate, and bear phialides (singular: phialide) over their entire-surface, contributing to its short and densely packed structure. The sterigmata are usually found to be uniseriate, numerous and crowded. Conidia formed in them are elliptical, smooth and comparatively thick-walled. A. clavatus usually express conidiophores 1.5–3.00 mm in length, which arises from specialized and widened hyphal cells that eventually become the branching foot cells. The conidia on A. clavatus has been measured up to 3.0 – 4.5 X 2.5 – 3.5 μm. Cleistothecia are produced in crosses after approximately 4–10 weeks of incubation on suitable growth media at 25 °C. Cleistothecia are yellowish-brown (fawn) to dark brown in colour and range in diameter from 315-700 μm in diameter and have a relatively hard outer wall (peridium). At maturity the cleistothecia contain asci that themselves contain ascospores, which are clear, lenticular (with ridges evident) and between 6.0-7.0 μm in diameter.
Growth on Czapek's solution agar
Aspergillus clavatus colonies grow rapidly on Czapek's solution agar, reaching 3.0–3.5 cm, in 10 days at 24–26 °C. Growth is usually plane or moderately furrowed, with occasional appearance of floccose strains. But generally, a comparatively thin surface layer of mycelial felt is observed, which produces a copious number of erect conidiophores. The reverse is usually uncoloured but becomes brown with passing time in some strains. While odor is not prominent in some strains, it can be extremely unpleasant in others. Large conidial heads extend from 300 to 400 μm by 150 to 200 μm when young. However, with time, they split into two or more divergent and compressed cordial chains reaching 1.00 mm portraying a colour consisting of artemisia green to slate olive. The observed conidiophores grow up to 1.5–3.00 mm in length with 20–30 μm in diameter. They slowly and ultimately enlarge at the apex into a clavate vesicle, which consists of a fertile area, 200 to 250 μm in length and 40–60 μm wide. The sterigmata usually ranges from 2.5 to 3.5 μm by 2.0 to 3.0 μm at the base of the vesicle, to 7.0 or 8.0 and occasionally 10 μm to 2.5 to 3.0 μm at the apex. The conidia are comparatively thick-walled and measures 3.0 to 4.5 μm by 2.5 to 3.5 μm. While they can be larger in some strains, in others their appearance may be irregular.
Growth on malt extract agar
On malt extract agar, the structural morphology of A. clavatus appears to be different than in Czapek's solution agar. The typical strains extracted from malt media contain less abundant conidial structures, which could be larger in size. In other (non-typical) strains, the conidial heads increase in number but decrease in size. The conidiophores range from 300 to 500 μm and bear loose, columnar heads. Typical strains may be resembled by strong and unpleasant odor whereas non-typical strains are characterized being odorless. The colonies arising from one conidium on malt extract agar, consisted of 25X10^7 conidia after being observed for six days.
Examination
The phialide development and conidium formation in A. clavatus has been examined using TEM. And by using SEM, it was discovered that the first-formed conidium and phialide share a continuous wall. Additionally recombination with an albino mutant led to the production of heterokaryotic conidial heads with mixed conidial colours. A GC-content of 52.5–55% was also detected upon DNA analysis. And its soluble wall carbohydrates consist of mannitol and arabitol.
Physiology
Light stimulates the elongation of conidiophores in A. clavatus. And the more favourable C sources include starch, dextrin, glycogen and especially fructose. Substantial degree of lipid synthesis occurs, whereas cellulose and usnic acid are degraded. A. clavatus also produces riboflavin, ribonuclease, acid phosphodiesterase and acid phosphatase when in liquid culture.
A. clavatus has the properties to oxidize tryptamine to indole acetic acid. It can absorb and collect hydrocarbons from fuel oil, incorporate metaphosphate and synthesize ethylene, clavatol and kojic acid. It is also responsible for the production of mycotoxins Patulin and sterigmatocystin. And has extremely high capacity for alcohol fermentation.
When it comes to genomics, bioinformatic analysis revealed that A. clavatus contains a full complement of identified euascomycete sex genes. A heterothallic sexual cycle involving outcrossing between MAT1-1 and MAT1-2 isolates was subsequently described . A. clavatus can also be a food source for Collembola and has been found to be parasitized by Fusarium solani.
Habitat and ecology
Aspergillus clavatus is often described as a spoilage organism occurring on dung and in soil and can also grow in strong alkaline conditions. When it comes to geographical distribution, A. clavatus has been spotted in the tropical, subtropical and Mediterranean areas. It has been accounted in low frequencies in the soils of India. And is also found in Bangladesh, Sri Lanka, Hong Kong, Jamaica, Brazil, Argentina, South Africa, the Ivory Coast, Egypt, Libya, Turkey, Greece, Italy, the United States of America, Japan, the USSR and Czechoslovakia. It was tracked in rocks of a carst cave and stratigraphic core samples descending to 1200 m in Central Japan. However, it is usually and solely collected from cultivated soils, including the ones that bear cotton, potatoes, sugar canes, legumes, paddy and Artemisia herba-alba. It has also been garnered from soil under burnt steppe vegetation, desert soils, the rhizospheres of banana, ground-nuts and wheat. A. clavatus has also been detected in the ripe compost of municipal waste, and Nitrogen and NPK fertilizers are found to play an important role in its stimulation process.
A. clavatus is also referred as a cosmopolitan fungus. Other than soil and dung, it can additionally be found in stored products with high levels of entrapped moisture. Such as stored cereals, rice, corn and millet. It has been further isolated from insects, especially from dead adult bees and honeycombs. Moreover, it has been collected from the feathers and droppings from free-living birds. A. clavatus is also common is decomposing materials. Their ability to resist strongly alkaline conditions, allows them to act as decomposition catalysts in situations where other fungus usually do not function.
Applications and medical uses
Weisner in March 1942 first noted the production of an antibiotic by strains of A. clavatus, and the active substance was known as clavatin. Later the antibiotic was named clavacin in August 1942 by Waksman, Horning and Spencer. Clavacin is also known as patulin. Patulin is receiving significant attention in the world today because of its manifestations in apple juices. Clavacin was noted to be valuable in the treatment of common-cold and applies a fungistatic or fungicidal effect on certain dermatophytes. A. clavatus with Phytophthora cryptogea in soil provided protection against damping of tomato seedlings, by decreasing the spreading of pathogens. Reversely, A. clavatus with the addition of glucose, increased the pathogenicity of Verticillium albo-atrum to tomatoes. A. clavatus also produces the following: Cytochalasin E, Cytochalasin K, Tryptoquivaline, Nortryptoquivalone, Nortryptoquivaline, Deoxytryptoquivaline, Deoxynortryptoquivaline, Tryptoquivaline E, and Tryptoquivaline N. Furthermore, A. clavatus isolates produce ribotoxins, which can help develop immunotherapy processes for cancer. A.clavatus has also been used in the formation of extracellular bionanoparticles from silver nitrate solutions. These nanoparticles display antimicrobial properties, which work against MRSA and MRSE.
Pathogenicity
Aspergillus clavatus is known as an agent of allergic aspergillosis and has been implicated in multiple pulmonary infections. It has also been labelled as an opportunistic fungus, as it is responsible for causing aspergillosis in compromised patients. A. clavatus can also cause neurotoxicosis in sheep and otomycosis. In Scotland and elsewhere, A. clavatus is reported for causing the mould allergy "malster's lung" otherwise "maltster's lung".
Extrinsic allergic alveolitis (EAA) is also caused by Aspergillus clavatus with a Type 1 immune reaction. It is described as a true hypersensitivity pneumonia, which usually occurs among malt workers, including symptoms of fever, chills, cough and dyspnea. In severe cases, glucocorticoids are used. Microgranulomatous hypersensitivity pneumonitis, where interstitial granulomatous infiltration occurs, usually in malt workers, is caused by allergy to antigens of Aspergillus clavatus. EAA is caused by allergy to Aspergillus conidia, usually in the non-atopic individual. Such individuals are usually exposed to organic dust heavily packed with conidia and mycelial debris. This condition involves the lung parenchyma.
A strain of A. clavatus has also caused hyperkeratosis in calves. Spore walls of a sputum-derived isolate of Aspergillus clavatus were extracted and treated with ethanol following alkaline hydrolysis. And it yielded mutagens. The extracts were given to unimmunised mice, causing lung reaction and leading to cases of pulmonary mycotoxicosis. A rising incidence of lung tumours were also observed. This study revealed that an isolate of A. clavatus, which is able to convert highly toxic metabolites in bacterial and mammalian cells, will cause inflammatory response in the lungs of unimmunized mice.
References
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- Howard, Dexter H., ed. (2003). Pathogenic fungi in humans and animals (2. ed.). New York : Dekker. p. 247. ISBN 978-0-8247-0683-8.
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{{cite book}}:|first1=has generic name (help) - ^ Ojeda-López, M; Chen W; Eagle CE; Gutiérrez G; Jia WL; Swilaiman SS; Huang Z; Park HS; Yu JH; Dyer PS (2018). "Evolution of asexual and sexual reproduction in the aspergilli". Studies in Mycology. 91: 37–59. doi:10.1016/j.simyco.2018.10.002. PMC 6231087. PMID 30425416.
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| Taxon identifiers | |
|---|---|
| Aspergillus clavatus | |