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*''Acrothecium arenarium'' <small>Moreau & V. Moreau (1941)</small>}}
*''Acrothecium arenarium'' <small>Moreau & V. Moreau (1941)</small>}}


'''''Curvularia inaequalis''''' is a plant [[saprobe]]<ref name="14x" /> that resides in temperate and subtropical environments.<ref name="8x" /> It is commonly found in the soils of [[forage grasses]] and grains.<ref name="7x" /> The species has been observed in a broad distribution of countries including Turkey, France, Canada, The United States, Japan and India.<ref name="12x" /> The species belongs to the group of [[dematiaceous]] [[hyphomycetes]].<ref name="10x" />
'''''Curvularia inaequalis''''' is a plant [[saprobe]]<ref name="14x" /> that resides in temperate and subtropical environments.<ref name="8x" /> It is commonly found in the [[soil]]s of [[forage grass]]es and [[grain]]s.<ref name="7x" /> The species has been observed in a broad distribution of countries including Turkey, France, Canada, The United States, Japan and India.<ref name="12x" /> This species is [[dematiaceous]] and a [[hyphomycete]].<ref name="10x" />


==History and taxonomy==
==History and taxonomy==
The ''Curvularia'' genus can be identified by its spiral borne phaeophragmospores, which contain both [[hyaline]] end cells and disproportionately large cells.<ref name="9x" /> They possess [[conidia]] with differing curvature and number of septa.<ref name="9x" /> ''Curvularia'' species ''C. inaequalis'' was first described in 1907 by ecologist [[Cornelius Lott Shear]].<ref name="13x" /> The fungus was isolated from diseased New Jersey cranberry pulp<ref name="13x" /> and termed ''Helminthosporium inaequale''.<ref name="9x" /> It was later renamed as ''Curvularia inaequalis'', by [[Karel Bernard Boedijn|Karl Boedijin]].<ref name="9x" /> During his taxonomic organization and grouping of the ''Curvularia'' species, Boedijin recognized a similarity between the conidia of ''C. inaequalis'' and of the '''lunata''' ''Curvularia'' group.<ref name="9x" /> Recognition of the three-[[septate]] curved conidia motivated the introduction of the now popularized name.<ref name="9x" />
The ''Curvularia'' genus can be identified by its spiral borne phaeophragmospores, which contain both [[hyaline]] end cells and disproportionately large cells.<ref name="9x" /> They possess [[conidia]] with differing curvature and number of septa.<ref name="9x" /> ''C. inaequalis'' was first described in 1907 by ecologist [[Cornelius Lott Shear]].<ref name="13x" /> The fungus was isolated from diseased New Jersey cranberry pulp<ref name="13x" /> and termed ''Helminthosporium inaequale''.<ref name="9x" /> Later, during [[Karel Bernard Boedijn|Karl Boedijin]]'s taxonomic organization and grouping of this [[genus]], he recognized a similarity between them and ''H. inaequale''.<ref name="9x" /> He recognized a [[morphological]] similarity between its [[conidia]] and those of the [[Curvularia lunata|''lunata'' group within ''Curvularia'']], and so renamed it ''C. inaequalis''.<ref name="9x" /> Recognition of the three-[[septate]] curved conidia motivated the introduction of the now popularized name.<ref name="9x" />


==Growth and morphology==
==Growth and morphology==
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''Curvularia inaequalis'' is a filamentous fungus, with 3 to 12 densely packed filaments.<ref name="13x" /> The species is mostly brown in appearance, with pale brown end cells.<ref name="8x" /><ref name="13x" /> Conidia themselves, consist of 3-5 cells with thick cell walls and a larger central cell.<ref name="13x" /><ref name="8x" /> The diameter of the conidia ranges from 10 to 30 micrometers and have a slight leading curvature.<ref name="5x" /><ref name="2x" /> Overall the appearance of the species is described as looking "cottony" with clear branching cells.<ref name="8x" /><ref name="13x" />
''Curvularia inaequalis'' is a filamentous fungus, with 3 to 12 densely packed filaments.<ref name="13x" /> The species is mostly brown in appearance, with pale brown end cells.<ref name="8x" /><ref name="13x" /> Conidia themselves, consist of 3-5 cells with thick cell walls and a larger central cell.<ref name="13x" /><ref name="8x" /> The diameter of the conidia ranges from 10 to 30 micrometers and have a slight leading curvature.<ref name="5x" /><ref name="2x" /> Overall the appearance of the species is described as looking "cottony" with clear branching cells.<ref name="8x" /><ref name="13x" />


The species can be difficult to identify due to its similar appearance to both ''[[Curvularia lunata]]'' and ''Curvularia geniculate''.<ref name="8x" /> Instead, sequencing of nuclear rRNA [[internal transcribed spacer]] regions (ITS) can be done to achieve accurate identification.<ref name="8x" />
The species can be difficult to identify due to its similar appearance to both ''[[Curvularia lunata|C.]]'' and ''[[Curvularia geniculate|geniculate]]''.<ref name="8x" /> Instead, sequencing of nuclear rRNA [[internal transcribed spacer]] regions (ITS) can be done to achieve accurate identification.<ref name="8x" />


==Physiology==
==Physiology==
The optimal growth temperature for the species is 30°C.<ref name="13x" /> It is able to produce a multitude of chemical products with enzymatic properties. One [[enzyme]] produced is [[chloroperoxidase]], which can catalyze [[halogenation]] reactions.<ref name="3x" /> Chloroperoxidase secreted from ''C. inaequalis'' contains [[vanadium]] active site.<ref name="10x" /> The presence of the vanadium substrate [[vanadate]] is essential for the function of chloroperoxidase.<ref name="16x" /> The compound [[glucose]] however, acts as an [[Enzyme inhibitor|inhibitor]] for both enzyme function and production.<ref name="17x" /> In its active form, the enzyme is able to then produce [[hypochlorous acid]], a strong [[oxidizing agent]].<ref name="10x" /> It has been theorized that ''C. inaequalis'' utilizes chloroperoxidase and hypochlorous acid in combination to penetrate the host's cell wall.<ref name="10x" />
The optimal growth temperature for the species is 30°C.<ref name="13x" /> It is able to produce a multitude of chemical products with enzymatic properties. One [[enzyme]] produced is [[chloroperoxidase]], which can catalyze [[halogenation]] reactions.<ref name="3x" /> Chloroperoxidase secreted from ''C. inaequalis'' contains [[vanadium]] active site.<ref name="10x" /> The presence of the vanadium substrate [[vanadate]] is essential for the function of chloroperoxidase.<ref name="16x" /> The compound [[glucose]] however, acts as an [[Enzyme inhibitor|inhibitor]] for both enzyme function and production.<ref name="17x" /> In its active form, the enzyme is able to then produce [[hypochlorous acid]], a strong [[oxidizing agent]].<ref name="10x" /> It has been theorized that ''C. inaequalis'' utilizes chloroperoxidase and hypochlorous acid in combination to penetrate the host's cell wall.<ref name="10x" />


Other significant compounds produced include of [[B-galactosidase]], 4-hydroxyradianthin and Curvularone A. The species is able to produce large amounts of B-galactosidase, which can hydrolyze lactose in acid whey.<ref name="5x" /> ''C. inaequalis'' also contains 4-hydroxyradianthin and Curvularone A compounds which have been identified as potential anti-tumor agents.<ref name="11x" />
Other significant compounds produced include of [[B-galactosidase]], {{ Visible anchor |4-hydroxyradianthin}} and {{ Visible anchor |Curvularone A}}. The species is able to produce large amounts of [[β-galactosidase]], which can hydrolyze lactose in acid whey.<ref name="5x" /> ''C. inaequalis'' also contains 4-hydroxyradianthin and Curvularone A compounds which have been identified as potential anti-tumor agents.<ref name="11x" />


==Pathology and toxicology==
==Pathology and toxicology==


===Plant pathology and toxicology===
===Plant pathology and toxicology===
''Curvularia inaequalis'' is known to cause leaf spot, also known as [[Leaf blight|Leaf Bligh]].<ref name="2x" /><ref name="6x" /> Symptoms of infection by ''C. Inaequalis'' include the combination of oval shaped dark brown patches and leaf tip dieback.<ref name="2x" /> The infection slowly spreads causes necrosis until it has covered the entirety of the leaf.<ref name="2x" /> It results in the thinning of grass vegetation such as [[Zoysia]], [[Bent grass|Bent]], [[Bermuda-grass|Bermuda]] and [[Buffalograss]].<ref name="2x" /><ref name="6x" /> Blighting is believed to be caused by two ''C. inaequalis'' [[mycotoxins]], [[Pyrenocine|Pyrenocines]] A and B.<ref name="6x" /> Pyrenocines A is the more potent of the two, stunting growth and causing necrosis in vegetation.<ref name="6x" /> Both cause leaf tip die back in turf grass and leaf leakage of electrolytes in Bermuda grass.<ref name="6x" />
''Curvularia inaequalis'' is known to cause leaf spot, also known as [[Leaf blight|Leaf Blight]].<ref name="2x" /><ref name="Pyrenocine" /> Symptoms of infection by ''C. inaequalis'' include the combination of oval shaped dark brown patches and leaf tip dieback.<ref name="2x" /> The infection slowly spreads causes necrosis until it has covered the entirety of the leaf.<ref name="2x" /> It results in the thinning of grass vegetation such as [[Zoysia]]-, [[Bent grass|Bent-]], [[Bermuda-grass|Bermuda-]] and [[Buffalograss|Buffalo-]] grasses.<ref name="2x" /><ref name="Pyrenocine" /> Blighting is believed to be caused by two ''C. inaequalis'' [[mycotoxins]], [[Pyrenocine]]s {{ Visible anchor |Pyrenocine A|text=A}} and {{ Visible anchor |Pyrenocine B|text=B}}.<ref name="Pyrenocine" /> Pyrenocines A is the more potent of the two, stunting growth and causing necrosis in vegetation.<ref name="Pyrenocine" /> Both cause leaf tip die back in turf grass and leaf leakage of [[electrolyte]]s in Bermuda grass.<ref name="Pyrenocine" />


===Human pathology===
===Human pathology===
''Curvularia inaequalis'' is typically a rare human pathogen.<ref name="8x" /> There are however, recorded medical cases that mention infection by the species. One such case is of an [[Eosinophilic]] fungal [[rhinosinusitis]] in an immunocompromised male.<ref name="1x" /> [[Endoscopic]] sinus surgery was required to remove a large [[polyposis]].<ref name="1x" /> ''C. inaequalis'' was found to have grown favorably in the eosinophilic mucus.<ref name="1x" /> Oral [[itraconazole]] and other [[corticosteroids]] successfully were administered to prevent reinfection.<ref name="1x" /> Another case of ''C. inaequalis'' causing disease includes [[peritonitis]] in an elderly patient.<ref name="8x" />
''Curvularia inaequalis'' is typically a rare human pathogen.<ref name="8x" /> There are however, recorded medical cases that mention infection by the species. One such case is of an [[Eosinophilic]] fungal [[rhinosinusitis]] in an immunocompromised male.<ref name="1x" /> [[Endoscopic]] sinus surgery was required to remove a large [[polyposis]].<ref name="1x" /> ''C. inaequalis'' was found to have grown favorably in the eosinophilic mucus.<ref name="1x" /> Oral [[itraconazole]] and other [[corticosteroids]] successfully were administered to prevent reinfection.<ref name="1x" /> Another case of ''C. inaequalis'' causing disease includes [[peritonitis]] in an elderly patient.<ref name="8x" />

It is suggested that contraction of the fungus occurs due to contact with soils.<ref name="8x" /> Furthermore, a case of recorded aerosolized ''C. inaequalis'' in one Canadian home supports airborne movement of spores as an important mode of transfer.<ref name="8x" /> While many cases of infection due to soil contact with the genus ''Curvularia'', connection with the specific species has not yet been confirmed.<ref name="8x" /> Further studies are required to determine its human pathogen potential.
It is suggested that contraction of the fungus occurs due to contact with soils.<ref name="8x" /> Furthermore, a case of recorded aerosolized ''C. inaequalis'' in one Canadian home supports airborne movement of spores as an important mode of transfer.<ref name="8x" /> While many cases of infection due to soil contact with the genus ''Curvularia'', connection with the specific species has not yet been confirmed.<ref name="8x" /> Further studies are required to determine its human pathogen potential.


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{{Reflist | 30em | refs =
{{Reflist | 30em | refs =


<ref name="1x">{{cite journal |last1=Posteraro |first1=B. |last2=Posteraro |first2=E. |last3=Sorda |first3=M. |last4=Torelli |first4=R. |last5=De Corso |first5=E. |title=Eosinophilic fungal rhinosinusitis due to the unusual pathogen Curvularia inaequalis |journal=Mycoses |date=2010 |volume=53 |issue=1 |pages=84–88 |doi=10.1111/j.1439-0507.2008.01671.x|pmid=19207840 }}</ref>
<ref name="1x">{{cite journal |last1=Posteraro |first1=B. |last2=Posteraro |first2=E. |last3=Sorda |first3=M. |last4=Torelli |first4=R. |last5=De Corso |first5=E. |title=Eosinophilic fungal rhinosinusitis due to the unusual pathogen ''Curvularia inaequalis'' |journal=[[Mycoses (journal)|Mycoses]] |date=2010 |volume=53 |issue=1 |pages=84–88 |doi=10.1111/j.1439-0507.2008.01671.x|pmid=19207840 }}</ref>


<ref name="2x">{{cite journal |last1=Amaradasa |first1=B. S. |last2=Amundsen |first2=K. |title=First Report of Curvularia inaequalis and Bipolaris spicifera Causing Leaf Blight of Buffalograss in Nebraska |journal=Plant Disease |date=February 2014 |volume=98 |issue=2 |page=279 |doi=10.1094/PDIS-05-13-0487-PDN|pmid=30708754 |doi-access=free }}</ref>
<ref name="2x">{{cite journal |last1=Amaradasa |first1=B. S. |last2=Amundsen |first2=K. |title=First Report of ''Curvularia inaequalis'' and ''Bipolaris spicifera'' Causing Leaf Blight of Buffalograss in Nebraska |journal=[[Plant Disease (journal)|Plant Disease]]|publisher=[[American Phytopathological Society]] (APS)|date=February 2014 |volume=98 |issue=2 |page=279 |doi=10.1094/PDIS-05-13-0487-PDN|pmid=30708754 |doi-access=free }}</ref>


<ref name="3x">{{cite book |last1=Lesk |first1=Arthur |title=Introduction to genomics |date=2012 |publisher=Oxford University Press |location=United Kingdom |isbn=978-0199564354 |page=339 |edition=2nd}}</ref>
<ref name="3x">{{cite book |last1=Lesk |first1=Arthur |title=Introduction to genomics |date=2012 |publisher=[[Oxford University Press]] (OUP)|location=[[United Kingdom]] |isbn=978-0199564354 |page=339 |edition=2nd}}</ref>


<ref name="5x">{{cite journal |last1=Laurila |first1=Heikki O. |last2=Nevalainen |first2=Helena |last3=Mikinen |first3=Veijo |title=Production of protoplasts from the fungi Curvularia inaequalis and Trichoderma reesei |journal=Applied Microbiology and Biotechnology |date=February 1985 |volume=21 |issue=3–4 |pages=210–212 |doi=10.1007/BF00295124}}</ref>
<ref name="5x">{{cite journal |last1=Laurila |first1=Heikki O. |last2=Nevalainen |first2=Helena |last3=Mikinen |first3=Veijo |title=Production of protoplasts from the fungi ''Curvularia inaequalis'' and ''Trichoderma reesei'' |journal=[[Applied Microbiology and Biotechnology]]|date=February 1985 |volume=21 |issue=3–4 |pages=210–212 |doi=10.1007/BF00295124}}</ref>


<ref name="Pyrenocine" >
⚫
<ref name="6x">{{cite journal |last1=Kim |first1=Jin-Cheol |last2=Choi |first2=Gyung Ja |last3=Kim |first3=Heung Tae |last4=Kim |first4=Hyun-Ju |last5=Cho |first5=Kwang Yun |title=Pathogenicity and Pyrenocine Production of Curvularia inaequalis Isolated from Zoysia Grass |journal=Plant Disease |date=June 2000 |volume=84 |issue=6 |pages=684–688 |doi=10.1094/PDIS.2000.84.6.684|pmid=30841112 |doi-access=free }}</ref>
This review...
{{ Cite book |
language=en|
year=2017|
edition=1|
location=[[Cham, Switzerland]]|
publisher=[[Springer International Publishing AG]]|
pages=xxv{{NBSP}}+{{NBSP}}411|
first2=Luis|
first1=Rosa|
last2=LLorca|
last1=Lopez|
title=Perspectives in Sustainable Nematode Management Through ''Pochonia chlamydosporia'' Applications for Root and Rhizosphere Health|
lccn=2017947835|
id={{ISBN|978-3-319-59222-0}}|
doi=10.1007/978-3-319-59224-4|
isbn=978-3-319-59224-4|
series=[[Sustainability in Plant and Crop Protection]]}}{{ RP |pages=154{{endash}}155}}
...cites this study:
⚫
{{cite journal |last1=Kim |first1=Jin-Cheol |last2=Choi |first2=Gyung Ja |last3=Kim |first3=Heung Tae |last4=Kim |first4=Hyun-Ju |last5=Cho |first5=Kwang Yun |title=Pathogenicity and Pyrenocine Production of ''Curvularia inaequalis'' Isolated from Zoysia Grass |journal=[[Plant Disease (journal)|Plant Disease]]|publisher=[[American Phytopathological Society]] (APS) |date=June 2000 |volume=84 |issue=6 |pages=684–688 |doi=10.1094/PDIS.2000.84.6.684|pmid=30841112 |doi-access=free }}
</ref>


<ref name="7x">{{cite book |last1=Sivanesan |first1=A. |title=Graminicolous species of bipolaris, curvularia, drechslera, exserohilum and their teleomorphs |date=1987 |publisher=C.A.B International |location=England |isbn=0851985874}}</ref>
<ref name="7x">{{cite book |last1=Sivanesan |first1=A. |title=Graminicolous species of ''Bipolaris'', ''Curvularia'', ''Drechslera'', ''Exserohilum'' and their teleomorphs |date=1987 |publisher=CABI ([[Centre for Agriculture and Bioscience International]])|location=[[England]] |isbn=0851985874}}</ref>


<ref name="8x">{{cite journal |last1=Pimentel |first1=J. D. |last2=Mahadevan |first2=K. |last3=Woodgyer |first3=A. |last4=Sigler |first4=L. |last5=Gibas |first5=C. |last6=Harris |first6=O. C. |last7=Lupino |first7=M. |last8=Athan |first8=E. |title=Peritonitis Due to Curvularia inaequalis in an Elderly Patient Undergoing Peritoneal Dialysis and a Review of Six Cases of Peritonitis Associated with Other Curvularia species. |journal=Journal of Clinical Microbiology |date=4 August 2005 |volume=43 |issue=8 |pages=4288–4292 |doi=10.1128/JCM.43.8.4288-4292.2005|pmid=16082004 |doi-access=free }}</ref>
<ref name="8x">{{cite journal |last1=Pimentel |first1=J. D. |last2=Mahadevan |first2=K. |last3=Woodgyer |first3=A. |last4=Sigler |first4=L. |last5=Gibas |first5=C. |last6=Harris |first6=O. C. |last7=Lupino |first7=M. |last8=Athan |first8=E. |title=Peritonitis Due to ''Curvularia inaequalis'' in an Elderly Patient Undergoing Peritoneal Dialysis and a Review of Six Cases of Peritonitis Associated with Other ''Curvularia'' species |journal=[[Journal of Clinical Microbiology]] |date=4 August 2005 |volume=43 |issue=8 |pages=4288–4292 |doi=10.1128/JCM.43.8.4288-4292.2005|pmid=16082004 |doi-access=free }}</ref>


<ref name="9x">{{cite journal |last1=Kendrick |first1=W. B. |last2=Cole |first2=G. T. |title=Conidium ontogeny in hyphomycetes. The sympodulae of Beauveria and Curvularia |journal=Canadian Journal of Botany |date=October 1968 |volume=46 |issue=10 |pages=1297–1301 |doi=10.1139/b68-172}}</ref>
<ref name="9x">{{cite journal |last1=Kendrick |first1=W. B. |last2=Cole |first2=G. T. |title=Conidium ontogeny in hyphomycetes. The sympodulae of ''Beauveria'' and ''Curvularia'' |journal=[[Canadian Journal of Botany]] |date=October 1968 |volume=46 |issue=10 |pages=1297–1301 |doi=10.1139/b68-172}}</ref>


<ref name="10x">{{cite journal |last1=Messerschmidt |first1=A. |last2=Wever |first2=R. |title=X-ray structure of a vanadium-containing enzyme: chloroperoxidase from the fungus Curvularia inaequalis. |journal=Proceedings of the National Academy of Sciences |date=9 January 1996 |volume=93 |issue=1 |pages=392–396 |doi=10.1073/pnas.93.1.392|pmid=8552646 |pmc=40244 |doi-access=free }}</ref>
<ref name="10x">{{cite journal |last1=Messerschmidt |first1=A. |last2=Wever |first2=R. |title=X-ray structure of a vanadium-containing enzyme: chloroperoxidase from the fungus ''Curvularia inaequalis'' |journal=[[Proceedings of the National Academy of Sciences]] |date=9 January 1996 |volume=93 |issue=1 |pages=392–396 |doi=10.1073/pnas.93.1.392|pmid=8552646 |pmc=40244 |doi-access=free }}</ref>


<ref name="11x">{{cite journal |last1=Pang |first1=Yan-Wei |last2=Zhang |first2=Ling-Jian |last3=Fang |first3=Jia-Shuang |last4=Liu |first4=Qian-Feng |last5=Zhang |first5=Hui |last6=Xiang |first6=Wen-Sheng |last7=Wang |first7=Ji-Dong |last8=Wang |first8=Xiang-Jing |title=Two new antitumor constituents from a soil fungus Curvularia inaequalis (strain HS-FG-257) |journal=The Journal of Antibiotics |date=23 January 2013 |volume=66 |issue=5 |pages=287–289 |doi=10.1038/ja.2012.128|pmid=23340662 }}</ref>
<ref name="11x">{{cite journal |last1=Pang |first1=Yan-Wei |last2=Zhang |first2=Ling-Jian |last3=Fang |first3=Jia-Shuang |last4=Liu |first4=Qian-Feng |last5=Zhang |first5=Hui |last6=Xiang |first6=Wen-Sheng |last7=Wang |first7=Ji-Dong |last8=Wang |first8=Xiang-Jing |title=Two new antitumor constituents from a soil fungus ''Curvularia inaequalis'' (strain HS-FG-257) |journal=The [[Journal of Antibiotics]] |date=23 January 2013 |volume=66 |issue=5 |pages=287–289 |doi=10.1038/ja.2012.128|pmid=23340662 }}</ref>


<ref name="12x">{{cite journal |last1=Moubasher |first1=A.H. |last2=Zeinab |first2=Soliman |last3=Abdel-Sater |first3=M.A. |title=Contribution to the mycobiota of Egypt: 25 new records and interesting fungal taxa from citrus and grapevine plantations |journal=Journal of Basic & Applied Mycology (Egypt) |date=2018 |volume=9 |pages=43}}</ref>
<ref name="12x">{{cite journal |last1=Moubasher |first1=A.H. |last2=Zeinab |first2=Soliman |last3=Abdel-Sater |first3=M.A. |title=Contribution to the mycobiota of Egypt: 25 new records and interesting fungal taxa from citrus and grapevine plantations |journal=[[Journal of Basic & Applied Mycology (Egypt)]]{{Unreliable source inline|date=December 2022}} |date=2018 |volume=9 |pages=43}}</ref>


<ref name="13x">{{cite book |last1=Shear |first1=Cornelius Lott |last2=Stevens |first2=Neil Everett |last3=Bain |first3=Henry F. |title=Fungous diseases of the cultivated cranberry |date=1931 |publisher=United States Department of Agriculture |location=Champaign, Illinois |pages=1–58}}</ref>
<ref name="13x">{{cite book |last1=Shear |first1=Cornelius Lott |last2=Stevens |first2=Neil Everett |last3=Bain |first3=Henry F. |title=Fungous diseases of the cultivated cranberry |date=1931 |publisher=[[United States Department of Agriculture]] |location=[[Champaign, Illinois]], US |pages=1–58}}</ref>


<ref name="14x">{{cite journal |last1=Ayoubi |first1=Najmeh |last2=Soleimani |first2=Mohammad Javad |last3=Zare |first3=Rasoul |last4=Zafari |first4=Doustmorad |title=First report of Curvularia inaequalis and C. spicifera causing leaf blight and fruit rot of strawberry in Iran |journal=Nova Hedwigia |date=1 August 2017 |volume=105 |issue=1 |pages=75–85 |doi=10.1127/nova_hedwigia/2017/0402}}</ref>
<ref name="14x">{{cite journal |last1=Ayoubi |first1=Najmeh |last2=Soleimani |first2=Mohammad Javad |last3=Zare |first3=Rasoul |last4=Zafari |first4=Doustmorad |title=First report of ''Curvularia inaequalis'' and ''C. spicifera'' causing leaf blight and fruit rot of strawberry in Iran |journal=[[Nova Hedwigia]] |date=1 August 2017 |volume=105 |issue=1 |pages=75–85 |doi=10.1127/nova_hedwigia/2017/0402}}</ref>


<ref name="16x">{{cite journal |last1=van Schijndel |first1=Johannes W.P.M. |last2=Vollenbroek |first2=Esther G.M. |last3=Wever |first3=Ron |title=The chloroperoxidase from the fungus Curvularia inaequalis; a novel vanadium enzyme |journal=Biochimica et Biophysica Acta (BBA) - Protein Structure and Molecular Enzymology |date=February 1993 |volume=1161 |issue=2 |pages=249–256 |doi=10.1016/0167-4838(93)90221-C}}</ref>
<ref name="16x">{{cite journal |last1=van Schijndel |first1=Johannes W.P.M. |last2=Vollenbroek |first2=Esther G.M. |last3=Wever |first3=Ron |title=The chloroperoxidase from the fungus ''Curvularia inaequalis''; a novel vanadium enzyme |journal=[[Biochimica et Biophysica Acta (BBA) - Protein Structure and Molecular Enzymology]] |date=February 1993 |volume=1161 |issue=2 |pages=249–256 |doi=10.1016/0167-4838(93)90221-C}}</ref>


<ref name="17x">{{cite journal |last1=Barnett |first1=Philip |last2=Kruitbosch |first2=Danny L |last3=Hemrika |first3=Wieger |last4=Dekker |first4=Henk L |last5=Wever |first5=Ron |title=The regulation of the vanadium chloroperoxidase from Curvularia inaequalis |journal=Biochimica et Biophysica Acta (BBA) - Gene Structure and Expression |date=May 1997 |volume=1352 |issue=1 |pages=73–84 |doi=10.1016/S0167-4781(96)00238-2|pmid=9177485 }}</ref>
<ref name="17x">{{cite journal |last1=Barnett |first1=Philip |last2=Kruitbosch |first2=Danny L |last3=Hemrika |first3=Wieger |last4=Dekker |first4=Henk L |last5=Wever |first5=Ron |title=The regulation of the vanadium chloroperoxidase from ''Curvularia inaequalis'' |journal=[[Biochimica et Biophysica Acta (BBA) - Gene Structure and Expression]] |date=May 1997 |volume=1352 |issue=1 |pages=73–84 |doi=10.1016/S0167-4781(96)00238-2|pmid=9177485 }}</ref>


<ref name="18x">{{cite journal |last1=Nagy |first1=P. |last2=Fischl |first2=G. |title=Effect of static magnetic field on growth and sporulation of some plant pathogenic fungi |journal=Bioelectromagnetics |date=May 2004 |volume=25 |issue=4 |pages=316–318 |doi=10.1002/bem.20015|pmid=15114642 }}</ref>
<ref name="18x">{{cite journal |last1=Nagy |first1=P. |last2=Fischl |first2=G. |title=Effect of static magnetic field on growth and sporulation of some plant pathogenic fungi |journal=[[Bioelectromagnetics]] |date=May 2004 |volume=25 |issue=4 |pages=316–318 |doi=10.1002/bem.20015|pmid=15114642 }}</ref>


}}
}}

Revision as of 18:31, 16 December 2022

Curvularia inaequalis
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Fungi
Division: Ascomycota
Class: Dothideomycetes
Order: Pleosporales
Family: Pleosporaceae
Genus: Curvularia
Species:
C. inaequalis
Binomial name
Curvularia inaequalis
(Shear) Boedijn (1907)
Synonyms
  • Helminthosporium inaequale Shear (1907)
  • Acrothecium arenarium Moreau & V. Moreau (1941)

Curvularia inaequalis is a plant saprobe[1] that resides in temperate and subtropical environments.[2] It is commonly found in the soils of forage grasses and grains.[3] The species has been observed in a broad distribution of countries including Turkey, France, Canada, The United States, Japan and India.[4] This species is dematiaceous and a hyphomycete.[5]

History and taxonomy

The Curvularia genus can be identified by its spiral borne phaeophragmospores, which contain both hyaline end cells and disproportionately large cells.[6] They possess conidia with differing curvature and number of septa.[6] C. inaequalis was first described in 1907 by ecologist Cornelius Lott Shear.[7] The fungus was isolated from diseased New Jersey cranberry pulp[7] and termed Helminthosporium inaequale.[6] Later, during Karl Boedijin's taxonomic organization and grouping of this genus, he recognized a similarity between them and H. inaequale.[6] He recognized a morphological similarity between its conidia and those of the lunata group within Curvularia, and so renamed it C. inaequalis.[6] Recognition of the three-septate curved conidia motivated the introduction of the now popularized name.[6]

Growth and morphology

The species' spore producing cells take on a model of sympodial growth.[6] Conidia grow through successive apices which end in a terminal porospore.[6] Growth can be affected by static magnetic fields with field flux densities.[8] Under these conditions, the number of conidia are able to increase by a minimum of 68 percent.[8]

Curvularia inaequalis is a filamentous fungus, with 3 to 12 densely packed filaments.[7] The species is mostly brown in appearance, with pale brown end cells.[2][7] Conidia themselves, consist of 3-5 cells with thick cell walls and a larger central cell.[7][2] The diameter of the conidia ranges from 10 to 30 micrometers and have a slight leading curvature.[9][10] Overall the appearance of the species is described as looking "cottony" with clear branching cells.[2][7]

The species can be difficult to identify due to its similar appearance to both C. and geniculate.[2] Instead, sequencing of nuclear rRNA internal transcribed spacer regions (ITS) can be done to achieve accurate identification.[2]

Physiology

The optimal growth temperature for the species is 30°C.[7] It is able to produce a multitude of chemical products with enzymatic properties. One enzyme produced is chloroperoxidase, which can catalyze halogenation reactions.[11] Chloroperoxidase secreted from C. inaequalis contains vanadium active site.[5] The presence of the vanadium substrate vanadate is essential for the function of chloroperoxidase.[12] The compound glucose however, acts as an inhibitor for both enzyme function and production.[13] In its active form, the enzyme is able to then produce hypochlorous acid, a strong oxidizing agent.[5] It has been theorized that C. inaequalis utilizes chloroperoxidase and hypochlorous acid in combination to penetrate the host's cell wall.[5]

Other significant compounds produced include of B-galactosidase, 4-hydroxyradianthin and Curvularone A. The species is able to produce large amounts of β-galactosidase, which can hydrolyze lactose in acid whey.[9] C. inaequalis also contains 4-hydroxyradianthin and Curvularone A compounds which have been identified as potential anti-tumor agents.[14]

Pathology and toxicology

Plant pathology and toxicology

Curvularia inaequalis is known to cause leaf spot, also known as Leaf Blight.[10][15] Symptoms of infection by C. inaequalis include the combination of oval shaped dark brown patches and leaf tip dieback.[10] The infection slowly spreads causes necrosis until it has covered the entirety of the leaf.[10] It results in the thinning of grass vegetation such as Zoysia-, Bent-, Bermuda- and Buffalo- grasses.[10][15] Blighting is believed to be caused by two C. inaequalis mycotoxins, Pyrenocines A and B.[15] Pyrenocines A is the more potent of the two, stunting growth and causing necrosis in vegetation.[15] Both cause leaf tip die back in turf grass and leaf leakage of electrolytes in Bermuda grass.[15]

Human pathology

Curvularia inaequalis is typically a rare human pathogen.[2] There are however, recorded medical cases that mention infection by the species. One such case is of an Eosinophilic fungal rhinosinusitis in an immunocompromised male.[16] Endoscopic sinus surgery was required to remove a large polyposis.[16] C. inaequalis was found to have grown favorably in the eosinophilic mucus.[16] Oral itraconazole and other corticosteroids successfully were administered to prevent reinfection.[16] Another case of C. inaequalis causing disease includes peritonitis in an elderly patient.[2]

It is suggested that contraction of the fungus occurs due to contact with soils.[2] Furthermore, a case of recorded aerosolized C. inaequalis in one Canadian home supports airborne movement of spores as an important mode of transfer.[2] While many cases of infection due to soil contact with the genus Curvularia, connection with the specific species has not yet been confirmed.[2] Further studies are required to determine its human pathogen potential.

References

  1. ^ Ayoubi, Najmeh; Soleimani, Mohammad Javad; Zare, Rasoul; Zafari, Doustmorad (1 August 2017). "First report of Curvularia inaequalis and C. spicifera causing leaf blight and fruit rot of strawberry in Iran". Nova Hedwigia. 105 (1): 75–85. doi:10.1127/nova_hedwigia/2017/0402.
  2. ^ a b c d e f g h i j k Pimentel, J. D.; Mahadevan, K.; Woodgyer, A.; Sigler, L.; Gibas, C.; Harris, O. C.; Lupino, M.; Athan, E. (4 August 2005). "Peritonitis Due to Curvularia inaequalis in an Elderly Patient Undergoing Peritoneal Dialysis and a Review of Six Cases of Peritonitis Associated with Other Curvularia species". Journal of Clinical Microbiology. 43 (8): 4288–4292. doi:10.1128/JCM.43.8.4288-4292.2005. PMID 16082004.
  3. ^ Sivanesan, A. (1987). Graminicolous species of Bipolaris, Curvularia, Drechslera, Exserohilum and their teleomorphs. England: CABI (Centre for Agriculture and Bioscience International). ISBN 0851985874.
  4. ^ Moubasher, A.H.; Zeinab, Soliman; Abdel-Sater, M.A. (2018). "Contribution to the mycobiota of Egypt: 25 new records and interesting fungal taxa from citrus and grapevine plantations". Journal of Basic & Applied Mycology (Egypt)[unreliable source?]. 9: 43.
  5. ^ a b c d Messerschmidt, A.; Wever, R. (9 January 1996). "X-ray structure of a vanadium-containing enzyme: chloroperoxidase from the fungus Curvularia inaequalis". Proceedings of the National Academy of Sciences. 93 (1): 392–396. doi:10.1073/pnas.93.1.392. PMC 40244. PMID 8552646.
  6. ^ a b c d e f g h Kendrick, W. B.; Cole, G. T. (October 1968). "Conidium ontogeny in hyphomycetes. The sympodulae of Beauveria and Curvularia". Canadian Journal of Botany. 46 (10): 1297–1301. doi:10.1139/b68-172.
  7. ^ a b c d e f g Shear, Cornelius Lott; Stevens, Neil Everett; Bain, Henry F. (1931). Fungous diseases of the cultivated cranberry. Champaign, Illinois, US: United States Department of Agriculture. pp. 1–58.
  8. ^ a b Nagy, P.; Fischl, G. (May 2004). "Effect of static magnetic field on growth and sporulation of some plant pathogenic fungi". Bioelectromagnetics. 25 (4): 316–318. doi:10.1002/bem.20015. PMID 15114642.
  9. ^ a b Laurila, Heikki O.; Nevalainen, Helena; Mikinen, Veijo (February 1985). "Production of protoplasts from the fungi Curvularia inaequalis and Trichoderma reesei". Applied Microbiology and Biotechnology. 21 (3–4): 210–212. doi:10.1007/BF00295124.
  10. ^ a b c d e Amaradasa, B. S.; Amundsen, K. (February 2014). "First Report of Curvularia inaequalis and Bipolaris spicifera Causing Leaf Blight of Buffalograss in Nebraska". Plant Disease. 98 (2). American Phytopathological Society (APS): 279. doi:10.1094/PDIS-05-13-0487-PDN. PMID 30708754.
  11. ^ Lesk, Arthur (2012). Introduction to genomics (2nd ed.). United Kingdom: Oxford University Press (OUP). p. 339. ISBN 978-0199564354.
  12. ^ van Schijndel, Johannes W.P.M.; Vollenbroek, Esther G.M.; Wever, Ron (February 1993). "The chloroperoxidase from the fungus Curvularia inaequalis; a novel vanadium enzyme". Biochimica et Biophysica Acta (BBA) - Protein Structure and Molecular Enzymology. 1161 (2): 249–256. doi:10.1016/0167-4838(93)90221-C.
  13. ^ Barnett, Philip; Kruitbosch, Danny L; Hemrika, Wieger; Dekker, Henk L; Wever, Ron (May 1997). "The regulation of the vanadium chloroperoxidase from Curvularia inaequalis". Biochimica et Biophysica Acta (BBA) - Gene Structure and Expression. 1352 (1): 73–84. doi:10.1016/S0167-4781(96)00238-2. PMID 9177485.
  14. ^ Pang, Yan-Wei; Zhang, Ling-Jian; Fang, Jia-Shuang; Liu, Qian-Feng; Zhang, Hui; Xiang, Wen-Sheng; Wang, Ji-Dong; Wang, Xiang-Jing (23 January 2013). "Two new antitumor constituents from a soil fungus Curvularia inaequalis (strain HS-FG-257)". The Journal of Antibiotics. 66 (5): 287–289. doi:10.1038/ja.2012.128. PMID 23340662.
  15. ^ a b c d e This review... Lopez, Rosa; LLorca, Luis (2017). Perspectives in Sustainable Nematode Management Through Pochonia chlamydosporia Applications for Root and Rhizosphere Health. Sustainability in Plant and Crop Protection (1 ed.). Cham, Switzerland: Springer International Publishing AG. pp. xxv + 411. doi:10.1007/978-3-319-59224-4. ISBN 978-3-319-59224-4. LCCN 2017947835. ISBN 978-3-319-59222-0.: 154–155  ...cites this study: Kim, Jin-Cheol; Choi, Gyung Ja; Kim, Heung Tae; Kim, Hyun-Ju; Cho, Kwang Yun (June 2000). "Pathogenicity and Pyrenocine Production of Curvularia inaequalis Isolated from Zoysia Grass". Plant Disease. 84 (6). American Phytopathological Society (APS): 684–688. doi:10.1094/PDIS.2000.84.6.684. PMID 30841112.
  16. ^ a b c d Posteraro, B.; Posteraro, E.; Sorda, M.; Torelli, R.; De Corso, E. (2010). "Eosinophilic fungal rhinosinusitis due to the unusual pathogen Curvularia inaequalis". Mycoses. 53 (1): 84–88. doi:10.1111/j.1439-0507.2008.01671.x. PMID 19207840.
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