{{DISPLAYTITLE:''Fusarium oxysporum'' f.sp. ''cubense''}} {{Short description|Fungus, causes banana wilt/Panama disease}} {{Infraspeciesbox special | image = Fusarium oxysporum f. sp. cubense race 1 (Foc-1) (38100977215).jpg | image_caption = ''Fusarium oxysporum'' f.sp. ''cubense'' race 1 growing for several days on brown rice | image_alt = | genus = Fusarium | species = oxysporum | infraspecies_rank1_name = Forma specialis | infraspecies_rank1_abbrev = f.sp. | infraspecies_rank1 = ''cubense'' | authority = E.F.Sm., W.C.Snyder & H.N.Hansen (1940) | synonyms = *''Fusarium cubense'' <small>E.F.Sm. (1910)</small> *''Fusarium oxysporum'' var. ''cubense'' <small>(E.F.Sm.) Wollenw. (1935)</small> *''Fusarium oxysporum'' f. ''cubense'' <small>(E.F.Sm.) W.C.Snyder & H.N.Hansen (1940)</small> }} '''''Fusarium oxysporum'' f. sp. ''cubense''''' ({{Audio|Fusarium Oxysporum F.SP. Cubense Pronunciation.ogg|Pronunciation}}) is a fungal plant pathogen that causes '''Panama disease''' of banana (''Musa'' spp.), also known as '''Fusarium wilt'''. The fungi and the related disease are responsible for widespread pressure on banana growing regions, destroying the economic viability of several commercially important banana varieties.

==Description== ''Fusarium oxysporum'' is a common inhabitant of soil<ref name="Drenth-Guest-2016" /> and produces three types of asexual spores: macroconidia, microconidia and chlamydospores.<ref>{{cite web |url=http://www.padil.gov.au/viewPestDiagnosticImages.aspx?id=1043 |title=Fusarium Wilt of Chickpea |publisher=Pests and Diseases Image Library |accessdate=2011-01-03 |archive-url=https://web.archive.org/web/20110321072331/http://www.padil.gov.au/viewPestDiagnosticImages.aspx?id=1043 |archive-date=2011-03-21 |url-status=dead }}</ref> The macroconidia are nearly straight, slender and thin-walled. They usually have three or four septa, a foot-shaped basal cell and a curved and tapered apical cell. They are generally produced from phialides on conidiophores by basipetal division. They are important in secondary infection.<ref name=CA>Couteaudier, Y. and C. Alabouvette, 1990 Survival and inoculum potential of conidia and chlamydospores of ''Fusarium oxysporum'' f. sp. ''lini'' in soil. Can. J. Microbiol. 36:551-556</ref>

[[Image:Porto de Tabatinga, Amazonas, Brazil 01.jpg|thumb|A warning sign at the port of Tabatinga, Brazil announcing ''"Do not transport seedlings from countries with the presence of the pest to Brazil"'']] The microconidia are ellipsoidal and have either a single septum or none at all. They are formed from phialides in false heads by basipetal division. They are important in secondary infection.<ref name=CA/>

The chlamydospores are globose with thick walls. They are either formed from hyphae or by the modification of hyphal cells. They endure in soils for long periods and act as inocula in primary infection.<ref name=CA/>

The macroconidia and chlamydospores are normally only formed on dead or dying host plants. Chlamydospores are the most significant survival structures of this pathogen.<ref name="PHP">{{Cite journal |last=Ploetz |first=Randy C. |s2cid=12228953 |title=Panama Disease:A Classic and Destructive Disease of Banana |journal=Plant Health Progress |publisher=American Phytopathological Society | volume=1 |pages=10 |year=2000 |doi=10.1094/PHP-2000-1204-01-HM}}</ref>

The teleomorph or sexual reproductive stage of ''F. oxysporum'' is unknown.<ref>Leslie JF, Summerell BA (2006) The Fusarium Laboratory manual. (Blackwell Publishing: Iowa, USA)</ref>

Four races of this pathogen have been described which attack different banana cultivars: * Race 1 attacks cultivars in the ''Musa'' (AAA group) 'Gros Michel'<ref name="Drenth-Guest-2016">{{cite journal | last1=Drenth | first1=André | last2=Guest | first2=David I. | title=Fungal and Oomycete Diseases of Tropical Tree Fruit Crops | journal=Annual Review of Phytopathology | publisher=Annual Reviews | volume=54 | issue=1 | date=2016-08-04 | issn=0066-4286 | doi=10.1146/annurev-phyto-080615-095944 | pages=373–395| pmid=27491435 }}</ref> and caused the 20th century epidemic.<ref name="Dita-et-al-2018">{{cite journal | last1=Dita | first1=Miguel | last2=Barquero | first2=Marcia | last3=Heck | first3=Daniel | last4=Mizubuti | first4=Eduardo S. G. | last5=Staver | first5=Charles P. | title=Fusarium Wilt of Banana: Current Knowledge on Epidemiology and Research Needs Toward Sustainable Disease Management | journal=Frontiers in Plant Science | publisher=Frontiers | volume=9 | date=2018-10-19 | page=1468 | issn=1664-462X | doi=10.3389/fpls.2018.01468| pmid=30405651 | pmc=6202804 | doi-access=free }}</ref> It also attacks 'Ducasse',<ref name="Drenth-Guest-2016" /> 'Lady Finger',<ref name="Drenth-Guest-2016" /> ''Musa'' (AAB group) 'Pome'{{cn|date=November 2021}} and its subgroups, Musa (AAB group) 'Silk'<ref name="Drenth-Guest-2016" /> and ''Musa'' (ABB group) 'Pisang Awak'.{{cn|date=November 2021}} (See {{section link||Race 1}}.) * Race 2 attacks ''Musa'' (ABB group) 'Bluggoe'<ref name="Drenth-Guest-2016" /> and its close relatives.{{cn|date=November 2021}} (See {{section link||Race 2}}.) * Race 3 attacks ''Heliconia'' spp.<ref name="Waite-1963">{{cite journal| last=Waite | first=B.H | date=1963 | title=Wilt of ''Heliconia'' spp. caused by ''Fusarium oxysporum'' f. sp. ''cubense'' Race 3 | journal=Tropical Agriculture (Trinidad) | volume=40 | pages=299–305}}</ref><ref name="Drenth-Guest-2016" /> (See {{section link||Race 3}}.) * Race 4 attacks ''Musa'' (AAA group) 'Dwarf Cavendish'<ref name="Crop-Prot-Comp" /><ref name="Ploetz-Pegg-2000" /> as well as the hosts of races 1 and 2.<ref name="Crop-Prot-Comp">Crop Protection Compendium 2005 Edition. ''Fusarium oxysporum'' f.sp. ''cubense'' (Panama disease of banana). (CAB International: Wallingford, UK).</ref><ref name="Ploetz-Pegg-2000" /><ref name="Drenth-Guest-2016" /> (See {{section link||Race 4}}.)

==Taxonomy== A ribosomal intergenic spacer analysis by Kurtz and Schouten 2009 failed to distinguish some ''F. oxysporum'' isolates merely endophytic on ''Musa'' from pathogenic ''Foc'' strains.<ref name="Schouten-2016" />

==Dispersal== Splash by rainfall, movement of contaminated soil, and movement of contaminated propagation materials are the major means of dispersal of ''Foc''. Dispersal by wind alone remains unproven and while animals can test positive for ''Foc'' on their outer surfaces, it remains unproven whether they can be effective vectors.<ref name="Dita-et-al-2018" /> Although it is a soil-borne pathogen, it does not compete well against other soil microbes for growth on dead buried tissue. It is nonetheless able to produce infection in living ''Musa'' hosts after a complete absence of hosts for 20 years - despite a population decline of 97% within the first three years. This is thought to be due to durable chlamydospores and due to persistence as an asymptomatic infection.<ref name="Gordon-2017" />

==Infection process== Anigorufone is a phytoalexin produced by ''Musa''. It is a nematicidal compound and so infection with ''Foc'' induces an anti-nematode defense. Anigorufone is the only nematicidal or nematistatic compound known among the phytoalexins.<ref name="Schouten-2016">{{cite journal | last=Schouten | first=Alexander | title=Mechanisms Involved in Nematode Control by Endophytic Fungi | journal=Annual Review of Phytopathology | publisher=Annual Reviews | volume=54 | issue=1 | date=2016-08-04 | issn=0066-4286 | doi=10.1146/annurev-phyto-080615-100114 | pages=121–142| pmid=27296146 | doi-access=free }}</ref> ''Foc'' rapidly invades cortical cells as do many other ''Fo f.sp.''.<ref name="Gordon-2017" />

==Reproduction== There is some debate as to whether ''Foc'' is sexual and this is investigated by studying its history of recombination - or lack of it. One study of the linkage disequilibrium of gametes showed relatively high disequilibrium and another a high degree of correlation between independent genetic markers, both of which are diagnostic for a lack of recombination and thus a clonal population. Other post-sequencing data analysis performed by the disequilibrium study also ''failed to reject'' recombination however this could be consistent with horizontal transfer. Horizontal transfer has been experimentally induced and appears to have been proven in ''Foc''s past and so seems the more likely explanation.<ref name="Gordon-2017">{{cite journal | last=Gordon | first=Thomas R. | title=''Fusarium oxysporum'' and the ''Fusarium'' Wilt Syndrome | journal=Annual Review of Phytopathology | publisher=Annual Reviews | volume=55 | issue=1 | date=2017-08-04 | issn=0066-4286 | doi=10.1146/annurev-phyto-080615-095919 | pages=23–39| pmid=28489498 | doi-access=free }}</ref> Both ''Fo'' mating types have been observed in ''Foc'' and protoperithecia-like structures are produced, but not the sexual structures. This does not necessarily mean that the sexual process has degenerated however, instead this may be a defect of the experiment.<ref name="Gordon-2017" />

Spores germinate at a higher rate in the presence of ''Musa'' root secondary metabolites from susceptible cultivars than those from resistant cultivars. This suggests that inhibition of germination is an important part of host resistance.<ref name="Gordon-2017" />

==Tropical Race 1/TR1== {{for-text|the disease and its place in the world|{{section link|Panama disease|Tropical Race 1}}}} {{visible anchor|Tropical Race 1}}/{{visible anchor|TR1}} is also found in ''Paspalum fasciculatum'', ''Panicum purpurescens'', ''Ixophorus unisetus'', and ''Commelina diffusa'' in Central America.<ref name="Waite-Dunlap-1953">{{cite journal | last1=Waite | first1=B.H. | last2=Dunlap | first2=V.C. | date=1953 | title=Preliminary host range studies with ''Fusarium oxysporum'' f. sp. ''cubense'' | journal=Plant Disease Reporter | publisher=United States Department of Agriculture | volume=37 | pages=79–80 | url=http://www.cabdirect.org/cabdirect/abstract/20057008257}}</ref> These weeds may be acting as an inoculum source.<ref name="Dita-et-al-2018" />

==Tropical Race 2/TR2== {{for-text|the disease and its place in the world|{{section link|Panama disease|Tropical Race 2}}}}{{anchor|Tropical Race 2|TR2}}

==Tropical Race 3/TR3== {{for-text|the disease and its place in the world|{{section link|Panama disease|Tropical Race 3}}}} {{visible anchor|Tropical Race 3}}/{{visible anchor|TR3}} is a pest of ''Heliconia'' ornamental flowers.<ref name="Waite-1963" /><ref name="Edel-Hermann-Lecomte-2019">{{cite journal | last1=Edel-Hermann | first1=V. | last2=Lecomte | first2=C. | title=Current Status of ''Fusarium oxysporum'' Formae Speciales and Races | journal=Phytopathology | publisher=American Phytopathological Society | volume=109 | issue=4 | year=2019 | issn=0031-949X | doi=10.1094/phyto-08-18-0320-rvw | pages=512–530| pmid=30461350 | doi-access=free }}</ref> Formerly reported to be a lesser pest of ''Musa balbisiana'' seedlings and of Gros Michel, but that is no longer thought to be true.<ref name="Edel-Hermann-Lecomte-2019" /><ref name="Ploetz-Pegg-2000">{{cite book | editor-last=Jones | editor-first=D. R. | title=Diseases of Banana, Abacá, and Enset | year=2000 | isbn=978-0-85199-355-3 | oclc=41347037 | page=143-159 | publisher=CABI (Centre for Agriculture and Bioscience International) | publication-place=Wallingford, Oxfordshire, England, United Kingdom | last1=Ploetz | first1=R.C. | last2=Pegg | first2=K.G.}}</ref> Now renamed ''Fusarium oxysporum'' f. sp. ''heliconiae''.<ref name="Ploetz-2006">{{cite journal | last=Ploetz | first=Randy C. | title=Fusarium Wilt of Banana Is Caused by Several Pathogens Referred to as ''Fusarium oxysporum'' f. sp. ''cubense'' | journal=Phytopathology | publisher=American Phytopathological Society | volume=96 | issue=6 | year=2006 | issn=0031-949X | doi=10.1094/phyto-96-0653 | pages=653–656| pmid=18943184 | doi-access=free }}</ref><ref name="Edel-Hermann-Lecomte-2019" />

==Race 4== ===Tropical Race 4/TR4=== {{Speciesbox | name = ''Fusarium odoratissimum'' | genus = Fusarium | species = odoratissimum | authority = Maryani ''et al.'', 2019{{nbsp}}<ref name="Maryani-et-al-2019-bundle" /> | synonyms = ''Foc'' strain TR4 | display_parents = 0 }} {{for-text|the disease and its place in the world|{{section link|Panama disease|Tropical Race 4}}}} <ref name="Pittaway-et-al-1999">{{cite journal | last1=Pittaway | first1=P. A. | last2=Nasir | first2=Nasril | last3=Pegg | first3=K. G. | title=Soil receptivity and host - pathogen dynamics in soils naturally infested with ''Fusarium oxysporum'' f. sp. ''cubense'', the cause of Panama disease in bananas | journal=Australian Journal of Agricultural Research | publisher=CSIRO Publishing (Commonwealth Scientific and Industrial Research Organisation) | volume=50 | issue=4 | year=1999 | issn=0004-9409 | doi=10.1071/a98152 | page=623}}</ref><ref name="Hennessy-et-al-2005">{{cite journal | last1=Hennessy | first1=Chelsea | last2=Walduck | first2=Geoff | last3=Daly | first3=Andrew | last4=Padovan | first4=Anna | title=Weed hosts of ''Fusarium oxysporum'' f. sp. ''cubense'' tropical race 4 in northern Australia | journal=Australasian Plant Pathology | publisher=Springer Science and Business Media LLC | volume=34 | issue=1 | year=2005 | issn=0815-3191 | doi=10.1071/ap04091 | page=115| bibcode=2005AuPP...34..115H | s2cid=13097833 }}</ref> {{visible anchor|Tropical Race 4}}/{{visible anchor|TR4}} belongs to vegetative compatibility group 01213/16. All cultivars which are susceptible to Race 1 and Race 2 are susceptible to TR4 (see {{section link||Race 1}} and {{section link||Race 2}}).<ref name="Molina-et-al-2009">{{cite journal | last1=Molina | first1=A.B. | last2=Fabregar | first2=E. | last3=Sinohin | first3=V.G. | last4=Yi | first4=G. | last5=Viljoen | first5=A. | title=Recent occurrence of ''Fusarium oxysporum'' f.sp. ''cubense'' tropical race 4 in Asia | journal=Acta Horticulturae | publisher=International Society for Horticultural Science (ISHS) | issue=828 | year=2009 | issn=0567-7572 | doi=10.17660/actahortic.2009.828.10 | pages=109–116 | hdl=2263/12191| hdl-access=free }}</ref> Starting in 2019 some authorities are following Maryani ''et al.'', 2019 in regarding this strain as ''{{visible anchor|Fusarium odoratissimum}}''.<ref name="Maryani-et-al-2019-bundle">{{Unbulleted list citebundle |{{cite journal | last1=Fisher | first1=Matthew C. | last2=Gurr | first2=Sarah J. | last3=Cuomo | first3=Christina A. | last4=Blehert | first4=David S. | last5=Jin | first5=Hailing | last6=Stukenbrock | first6=Eva H. | last7=Stajich | first7=Jason E. | last8=Kahmann | first8=Regine | last9=Boone | first9=Charles | last10=Denning | first10=David W. | last11=Gow | first11=Neil A. R. | last12=Klein | first12=Bruce S. | last13=Kronstad | first13=James W. | last14=Sheppard | first14=Donald C. | last15=Taylor | first15=John W. | last16=Wright | first16=Gerard D. | last17=Heitman | first17=Joseph | last18=Casadevall | first18=Arturo | last19=Cowen | first19=Leah E. | editor-last=Chowdhary | editor-first=Anuradha | title=Threats Posed by the Fungal Kingdom to Humans, Wildlife, and Agriculture | journal=mBio | publisher=American Society for Microbiology & American Academy of Microbiology | volume=11 | issue=3 | date=2020-06-30 | issn=2161-2129 | doi=10.1128/mbio.00449-20 | pmid=32371596 | pmc=7403777 | s2cid=218521623| doi-access=free }} |{{cite journal | last1=Cheek | first1=Martin | last2=Nic Lughadha | first2=Eimear | last3=Kirk | first3=Paul | last4=Lindon | first4=Heather | last5=Carretero | first5=Julia | last6=Looney | first6=Brian | last7=Douglas | first7=Brian | last8=Haelewaters | first8=Danny | last9=Gaya | first9=Ester | last10=Llewellyn | first10=Theo | last11=Ainsworth | first11=A. Martyn | last12=Gafforov | first12=Yusufjon | last13=Hyde | first13=Kevin | last14=Crous | first14=Pedro | last15=Hughes | first15=Mark | last16=Walker | first16=Barnaby E. | last17=Campostrini Forzza | first17=Rafaela | last18=Wong | first18=Khoon Meng | last19=Niskanen | first19=Tuula | title=New scientific discoveries: Plants and fungi | journal= Plants, People, Planet | publisher=New Phytologist Foundation (Wiley) | volume=2 | issue=5 | year=2020 | issn=2572-2611 | doi=10.1002/ppp3.10148 | pages=371–388 | s2cid=225177847| doi-access=free | hdl=1854/LU-8705210 | hdl-access=free }} |{{cite journal | last1=Drenth | first1=André | last2=McTaggart | first2=Alistair R. | last3=Wingfield | first3=Brenda D. | title=Fungal clones win the battle, but recombination wins the war | journal=IMA Fungus | publisher=International Mycological Association (Springer) | volume=10 | issue=1 | date=2019-10-29 | page=18 | issn=2210-6359 | doi=10.1186/s43008-019-0020-8 | s2cid=204941675 | pmid=32647622 | pmc=7325676 | doi-access=free }} |{{cite journal | last1=Lücking | first1=Robert | last2=Aime | first2=M. Catherine | last3=Robbertse | first3=Barbara | last4=Miller | first4=Andrew N. | last5=Aoki | first5=Takayuki | last6=Ariyawansa | first6=Hiran A. | last7=Cardinali | first7=Gianluigi | last8=Crous | first8=Pedro W. | last9=Druzhinina | first9=Irina S. | last10=Geiser | first10=David M. | last11=Hawksworth | first11=David L. | last12=Hyde | first12=Kevin D. | last13=Irinyi | first13=Laszlo | last14=Jeewon | first14=Rajesh | last15=Johnston | first15=Peter R. | last16=Kirk | first16=Paul M. | last17=Malosso | first17=Elaine | last18=May | first18=Tom W. | last19=Meyer | first19=Wieland | last20=Nilsson | first20=Henrik R. | last21=Öpik | first21=Maarja | last22=Robert | first22=Vincent | last23=Stadler | first23=Marc | last24=Thines | first24=Marco | last25=Vu | first25=Duong | last26=Yurkov | first26=Andrey M. | last27=Zhang | first27=Ning | last28=Schoch | first28=Conrad L. | title=Fungal taxonomy and sequence-based nomenclature | journal=Nature Microbiology | publisher=Nature Portfolio | volume=6 | issue=5 | date=2021-04-26 | issn=2058-5276 | doi=10.1038/s41564-021-00888-x | pages=540–548 | s2cid=233410673 | pmid=33903746| pmc=10116568 }} }}</ref> However, the validity of this taxonomic change has been challenged.<ref name="Torres-Bedoya-2021"> :{{cite book|year=2022|title=Regional strategy and action plan for the prevention, preparedness, response and recovery of Latin America and the Caribbean to Fusarium wilt of Musaceae tropical race 4|isbn=978-92-5-136494-9|s2cid=252425659|doi=10.4060/cb8674en|publisher=FAO (Food and Agriculture Organization of the United Nations)}} : :{{cite journal|issue=1|publisher=Wiley-VCH GmbH|volume=20|last1=Bragard|first1=Claude|last2=Baptista|first2=Paula|last3=Chatzivassiliou|first3=Elisavet|last4=Di Serio|first4=Francesco|last5=Gonthier|first5=Paolo|last6=Jaques Miret|first6=Josep Anton|last7=Justesen|first7=Annemarie Fejer|last8=MacLeod|first8=Alan|last9=Magnusson|first9=Christer Sven|last10=Milonas|first10=Panagiotis|last11=Navas-Cortes|first11=Juan A|last12=Parnell|first12=Stephen|last13=Potting|first13=Roel|last14=Stefani|first14=Emilio|last15=Thulke|first15=Hans-Hermann|last16=Van der Werf|first16=Wopke|last17=Civera|first17=Antonio Vicent|last18=Yuen|first18=Jonathan|last19=Zappalà|first19=Lucia|last20=Migheli|first20=Quirico|last21=Vloutoglou|first21=Irene|last22=Maiorano|first22=Andrea|last23=Streissl|first23=Franz|last24=Reignault|first24=Philippe Lucien|journal=EFSA Journal|issn=1831-4732|s2cid=246179769|doi=10.2903/j.efsa.2022.7092|title=Pest categorisation of ''Fusarium oxysporum'' f. sp. ''cubense'' Tropical Race 4|date=2022 |pages=e07092 |pmid=35079290 |pmc=8780018 |id=European Food Safety Authority (EFSA)|hdl=11380/1280747|hdl-access=free}} : :These reviews cite this research. : :{{cite journal|year=2021|first3=David|last1=Torres|first2=Daniel|issue=12|last3=Studholme|last2=Bebber|first1=Eliana|volume=111|pages=2141–2145|s2cid=235371340|publisher=American Phytopathological Society (APS)|journal=Phytopathology|issn=0031-949X|doi=10.1094/phyto-03-21-0089-le|title=Taxonomic Revision of the Banana Fusarium Wilt TR4 Pathogen Is Premature|pmid=34100303 |author2-link=Daniel Bebber|hdl=10871/126077|hdl-access=free}} </ref>

===Subtropical Race 4/STR4=== {{for-text|the disease and its place in the world|{{section link|Panama disease|Subtropical Race 4}}}} {{visible anchor|Subtropical Race 4}}/{{visible anchor|STR4}} is a subtropical race and does not become symptomatic on Cavendish until the trees are stressed by cold.<ref name="QldAgDept">{{cite web | archive-url=https://web.archive.org/web/20181222213528/https://www.daf.qld.gov.au/business-priorities/biosecurity/plant/health-pests-diseases/a-z-list-of-emergency-plant-pests-and-diseases/panama-disease | archive-date=22 December 2018 | title=Panama disease tropical race 4 (TR4) - Business Queensland | website=Queensland Agriculture Department | date=2020-11-13 | url=http://www.daf.qld.gov.au/business-priorities/biosecurity/plant/health-pests-diseases/a-z-list-of-emergency-plant-pests-and-diseases/panama-disease | access-date=2021-02-06}}</ref><ref name="Drenth-Guest-2016" /> Also found in ''Paspalum'' spp. and ''Amaranthus'' spp. in Australia.<ref name="Pittaway-et-al-1999" /> These weeds may be acting as sources of inoculum.<ref name="Dita-et-al-2018" />

==Research== Much research is being undertaken because of the urgency in formulating effective control methods for Panama disease and breeding resistant banana cultivars. Researchers at University Sains Malaysia are examining variability in the genome of the pathogen and its genetic variability is being studied, as are the evolutionary relationships within vegetative compatibility groups of the pathogen.<ref name="Fourie-et-al-2009">{{Cite journal | last1 = Fourie | first1 = G. | last2 = Steenkamp | first2 = E. T. | last3 = Gordon | first3 = T. R. | last4 = Viljoen | first4 = A. | title = Evolutionary Relationships among the ''Fusarium oxysporum'' f. sp. ''cubense'' vegetative compatibility groups | doi = 10.1128/AEM.00370-09 | journal = Applied and Environmental Microbiology | volume = 75 | issue = 14 | pages = 4770–81 | year = 2009 | pmid = 19482953| pmc =2708428 | bibcode = 2009ApEnM..75.4770F }}</ref>

Research into the phylogenetic relationships among the different strains of ''F. oxysporum'' that cause wilt of banana has been undertaken to determine whether the strains that are specific to the banana have descended from a common ancestor or have developed independently. Results of this study show that it is not monophyletic and appears to have multiple evolutionary origins.<ref>{{cite journal|title=Multiple evolutionary origins of the fungus causing Panama disease of banana: Concordant evidence from nuclear and mitochondrial gene genealogies |journal=Proceedings of the National Academy of Sciences|volume=95 |issue=5 |pages=2044–2049 |date=1998-03-03 |doi=10.1073/pnas.95.5.2044 |pmid=9482835 |last1=O'Donnell |first1=K |last2=Kistler |first2=H. C |last3=Cigelnik |first3=E |last4=Ploetz |first4=R. C |pmc=19243 |bibcode=1998PNAS...95.2044O |doi-access=free}}</ref> The largest lineages of ''F. oxysporum ''f. sp.'' cubense'' ({{section link||Race 1}} and {{section link||Race 2}}) are genetically distinct from a lineage originating from East Africa ({{section link||Race 5}}) and developed pathogenicity for bananas independently from one another.<ref>{{cite journal | last1 = Koenig | first1 = R L | display-authors = etal | year = 1997 | title = ''Fusarium oxysporum'' f. sp. ''cubense'' consists of a small number of divergent and globally distributed clonal lineages | journal =Phytopathology| volume = 87 | issue = 9 | pages = 915–923 | doi = 10.1094/phyto.1997.87.9.915 | pmid = 18945062 | doi-access = free }}</ref>

Identification, differentiation, and usage of vegetative compatibility groups is useful and valid within ''Foc'' because there are relatively few VCGs.<ref name="Moore-et-al-2001">{{cite journal | last1=Moore | first1=N.Y. | last2=BENTLEY | first2=S. | last3=BUDDENHAGEN| first3=I.W. | last4=PEGG | first4=K.G. | title=Fusarium wilt of banana, a diverse clonal pathogen of a domesticated clonal host | website=Alianza SIDALC | url=http://www.sidalc.net/cgi-bin/wxis.exe/?IsisScript=ARTIC.xis&B1=Buscar&formato=1&cantidad=50&expresion=MOORE,%20N.Y. | language=en | access-date=2021-01-05 | date=2001 | location=Saint Paul, Minnesota | pages=212–224 | publisher=American Phytopathological Society Press |editor=B.A. Summerell |editor2=J.F. Leslie |editor3=D. Backhouse |editor4=W.L. Bryden |editor5=L.W. Burgess | series=Fusarium – Paul E. Nelson Memorial Symposium}}</ref>

==Management== Segura-Mena ''et al.'', 2021 finds that ''Foc'' {{Slink||R1}} and {{Slink||TR4}} are highly sensitive to pH.<ref name="Evaluating" /> They find that this is a potential management method in this disease.<ref name="Evaluating"> :{{cite conference|year=2023|publisher=IEEE|conference=2023 4th International Conference on Computing, Mathematics and Engineering Technologies (iCoMET)|last1=Memon|first1=Komal|last2=Umrani|first2=Fahim Aziz|last3=Baqai|first3=Attiya|last4=Syed|first4=Zafi Sherhan|doi=10.1109/icomet57998.2023.10099182|s2cid=258260664|title=A Review Based On Comparative Analysis of Techniques Used in Precision Agriculture}} : :This review cites this research. : :{{cite journal|issue=2|year=2021|publisher=Springer Science and Business Media LLC|first6=J.|first5=G.|first4=M.|first3=F.|first2=J.|first1=R.|last6=Sandoval|last5=Kema|last4=Salacinas-Niez|last3=García-Bastidas|last2=Stoorvogel|last1=Segura-Mena|volume=160|pages=441–455|journal=European Journal of Plant Pathology|issn=0929-1873|s2cid=233648418|doi=10.1007/s10658-021-02255-2|title=Evaluating the potential of soil management to reduce the effect of ''Fusarium oxysporum'' f. sp. ''cubense'' in banana (''Musa'' AAA)|id=Royal Netherlands Society of Plant Pathology (KNPV)+European Foundation for Plant Pathology (EFPP)|doi-access=free|bibcode=2021EJPP..160..441S }} </ref>

==See also== *List of banana and plantain diseases

==References== {{Reflist}}

==Further reading== * {{cite journal | last1=Ploetz | first1=Randy | last2=Pegg | first2=Kenneth | title=Fusarium wilt of banana and Wallace's line: Was the disease originally restricted to his Indo-Malayan region? | journal=Australasian Plant Pathology | publisher=Springer Science and Business Media LLC | volume=26 | issue=4 | year=1997 | issn=0815-3191 | doi=10.1071/ap97039 | page=239| bibcode=1997AuPP...26..239P | s2cid=36822018 }} * {{cite web | title=Information | website=FRAC (Fungicide Resistance Action Committee) | author=Banana Working Group | date=2020-01-31 | url=http://www.frac.info/frac-teams/working-groups/banana-group/information | access-date=2021-05-21}} * {{cite web | title=Recommendations for Bananas | website=FRAC (Fungicide Resistance Action Committee) | author=Banana Working Group | date=2020-01-31 | url=http://www.frac.info/frac-teams/working-groups/banana-group/recommendations-for-bananas | access-date=2021-05-21}}

==External links== * [http://www.promusa.org/Fusarium+wilt Information on Fusarium wilt on Musapedia] * {{cite web |title=Q&A on Tropical Race 4 |website=Dole Food Company |url=https://www.dole.com/en/blog/healthy-living/tropical-race-4 |access-date=2020-12-11}} * {{cite web | title=Emergencia Fusarium R4T | website=SENASA Perú | date=1999-02-22 | url=http://www.gob.pe/institucion/senasa/campa%C3%B1as/4295-emergencia-fusarium-r4t | language=es | access-date=2021-08-06}} * {{cite web | title=Panama TR4 Protect | website=Biosecurity Queensland | url=http://panamatr4protect.com.au/ | access-date=2021-09-21}}

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Category:Fungal plant pathogens and diseases Category:Banana diseases oxysporum f.sp. cubense Category:Forma specialis taxa Category:Banana Wars