Fusarium oxysporum f.sp. cubense: Difference between revisions

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Better {{anchor}}s in the race list. Vegetative compatibility group 01213/16, Formosana (GCTCV-218), GCTCV-119, from Molina et al 2009. Better format my Queensland/Dita PCR text. {{convert}}. Try wikilinking inoculum as Propagule.
Large transfer of text and citations to Panama disease. Almost the entire article did not pertain to Foc, but instead to the disease.
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'''''Fusarium oxysporum'' f. sp. ''cubense''''' {{Audio|Fusarium Oxysporum F.SP. Cubense Pronunciation.ogg|Pronunciation}} is a [[fungus|fungal]] [[plant pathogen]] that causes '''[[Panama disease]]''' of [[banana]] (''[[Musa (genus)|Musa]]'' spp.), also known as '''fusarium wilt of banana'''.
'''''Fusarium oxysporum'' f. sp. ''cubense''''' {{Audio|Fusarium Oxysporum F.SP. Cubense Pronunciation.ogg|Pronunciation}} is a [[fungus|fungal]] [[plant pathogen]] that causes '''[[Panama disease]]''' of [[banana]] (''[[Musa (genus)|Musa]]'' spp.), also known as '''fusarium wilt of banana'''.

==Overview==
Although fruits of the wild bananas (''Musa'' spp.) have large, hard seeds, most edible bananas are seedless. Banana plants are therefore propagated [[Asexual reproduction|asexually]] from offshoots. Because these rhizomes are usually free of symptoms even when the plant is infected by ''F. oxysporum ''f. sp.'' cubense'', they are a common means by which this pathogen is disseminated. It can also be spread in soil and running water, on farm implements or machinery.<ref name="Stover-1962">{{cite book |title=Fusarial Wilt (Panama Disease) of Bananas and other ''Musa'' species |series=Phytopathological Papers | volume=4 | last=Stover | first=R. H. | publisher=[[Royal Botanic Gardens, Kew]] - [[Commonwealth Mycological Institute]] | date=1962 | oclc=3494840 | pages=vi+117 | url=https://books.google.com/books?id=AnvgzAEACAAJ | isbn=978-0000000859}}</ref>

Panama disease is one of the most destructive plant diseases of modern times.<ref name="Stover-1962" /><ref name="Stover-Simmonds-1987" /> It is believed to have originated in Southeast Asia<ref name="Dita-et-al-2018" /> and was first reported in Australia in 1876.<ref name="PHP" /> By 1950 it had spread to all the banana-producing regions of the world with the exception of some islands in the South Pacific, the Mediterranean, Melanesia and Somalia.<ref name="PHP">{{Cite journal |last=Ploetz |first=R. C. |s2cid=12228953 |title=Panama Disease:A Classic and Destructive Disease of Banana |journal= Plant Health Progress|volume=1 |pages=10 |year=2000 |doi=10.1094/PHP-2000-1204-01-HM}}</ref>
[[File:Approximately 30 Gros Michel Bananas.jpg|thumb|200px|left|Gros Michel bananas]]
Panama disease affects a wide range of banana cultivars; however, it is best known for the damage it caused to a single cultivar in the early export plantations.<ref name="Stover-1962" /> Before 1960, a total reliance was put on the cultivar '[[Gros Michel banana|Gros Michel]]', and it supplied almost all the export trade. It proved susceptible to the disease<ref name="Dita-et-al-2018" /> and the use of infected rhizomes to establish new plantations caused widespread and severe losses. Some indication of the scale of the losses is demonstrated by the complete eradication of production on {{ convert | 30,000 | hectare | acre }} of plantation in the Ulua Valley of Honduras between 1940 and 1960. In Suriname, an entire operation of {{ convert | 4,000 | hectare | acre }} was out of business within eight years and in the Quepos area of Costa Rica, {{ convert | 6,000 | hectare | acre }} were destroyed in twelve years.<ref name="PHP"/> Overall [[fungal banana disease|fungal diseases]] - including most prominently ''Foc'' - are disproportionately important to [[small island developing states]].<ref name="Thomas-et-al-2020">{{cite journal | last1=Thomas | first1=Adelle | last2=Baptiste | first2=April | last3=Martyr-Koller | first3=Rosanne | last4=Pringle | first4=Patrick | last5=Rhiney | first5=Kevon | title=Climate Change and Small Island Developing States | journal=[[Annual Review of Environment and Resources]] | publisher=[[Annual Reviews (publisher)|Annual Reviews]] | volume=45 | issue=1 | date=2020-10-17 | issn=1543-5938 | doi=10.1146/annurev-environ-012320-083355 | pages=1–27| doi-access=free }}</ref>

[[File:Bananen Frucht.jpg|thumb|right|100px|A [[Cavendish bananas|Cavendish banana]]]]
By the middle of the 20th century, resistant cultivars in the '[[Cavendish banana|Cavendish]]' subgroup were being used as a substitute for 'Gros Michel' in the export trade.<ref name="Ploetz-Pegg-2000" /><ref name="Dita-et-al-2018" /> For a time these cultivars did prove resilient and grew well, and in some areas remain the clones on which the current export trade is based. Unfortunately, in several growing areas in the Eastern Hemisphere, these cultivars are falling to TR4.<ref name="Dita-et-al-2018" /> It was inevitable that this disease would spread to the Western Hemisphere,<ref name="FAO-DOI-10-17660-ActaHortic-2011-897-45">{{ cite web | url=http://www.fao.org/fileadmin/templates/banana/documents/Docs_Resources_2015/TR4/IN130262_pp.pdf | title=Raising Awareness of the Threat of Fusarium Wilt Tropical Race 4 in Latin America and the Caribbean }}</ref> and indeed in 2019 it was detected in [[Colombia]].<ref name="first-report-tr4-in-colombia">{{ Cite journal | title=First Report of Fusarium Wilt Tropical Race 4 in Cavendish Bananas Caused by ''Fusarium odoratissimum'' in Colombia | year=2020 | doi=10.1094/PDIS-09-19-1922-PDN | last1=García-Bastidas | first1=F. A. | last2=Quintero-Vargas | first2=J. C. | last3=Ayala-Vasquez | first3=M. | last4=Schermer | first4=T. | last5=Seidl | first5=M. F. | last6=Santos-Paiva | first6=M. | last7=Noguera | first7=A. M. | last8=Aguilera-Galvez | first8=C. | last9=Wittenberg | first9=A. | last10=Hofstede | first10=R. | last11=Sørensen | first11=A. | last12=Kema | first12=G. H. J. | journal=Plant Disease | volume=104 | issue=3 | page=994 | doi-access=free }}</ref> This poses a significant threat to production, because there are currently no acceptable replacement cultivars.<ref name="PHP"/> It is anticipated by experts that [[crop disease surveillance|disease surveillance]], [[integrated pest management]], breeding of resistant cultivars, and genetic engineering will yield worthwhile results.<ref name="Staver-et-al-2020" /> This conclusion comes from an economic analysis which examined these as investments which governments and international organizations may or may not choose to invest in.<ref name="Staver-et-al-2020" /> Their opinion was that there would be positive yield from these investments, taken as either [[net present value]] or the [[internal rate of return]].<ref name="Staver-et-al-2020" />

A variant cultivar called Formosana (GCTCV-218), a type of Taiwanese Cavendish, has been reported in the popular press as having some resistance to TR4,<ref>{{cite news |last=Gittleson |first=Kim |title=Battling to save the world's bananas |publisher=BBC |date=2018-02-01 |url= https://www.bbc.com/news/business-42777803 |accessdate=2019-09-30}}</ref> as well as by Molina et al 2009,<ref name="Molina-et-al-2009" /> although in 2015 Ploetz was of the opinion that is has been evaluated as having no resistance at all.<ref name="Ploetz-2015">{{cite journal | last=Ploetz | first=Randy C. | title=Management of ''Fusarium'' wilt of banana: A review with special reference to tropical race 4 | journal=[[Crop Protection (journal)|Crop Protection]] | publisher=[[International Association for the Plant Protection Sciences]] ([[Elsevier]]) | volume=73 | year=2015 | issn=0261-2194 | doi=10.1016/j.cropro.2015.01.007 | pages=7–15 | s2cid=86468883}}</ref> '[[GCTCV-119]]' is another Taiwanese Cavendish used commercially for its TR4 resistance.<ref name="Molina-et-al-2009" />

To make things worse, this variant of the pathogen also affects [[true plantain|plantain]]s, ''Musa acuminata × balbisiana'', which are an important [[staple food]] in tropical regions of the world.<ref name="PHP"/> The average American eats {{convert|26.2|lb|kg}} of the Cavendish banana each year, and the question is being asked as to whether this oft-consumed fruit is on course to extinction.<ref>{{cite web |last=Koeppel |first=Dan |title=Can this fruit be saved? |publisher=Popular Science |url=http://www.popsci.com/scitech/article/2008-06/can-fruit-be-saved |date=2005-06-19 |accessdate=2011-01-03}}</ref>

Apart from the export trade, 85% of banana production is for local consumption and many of the cultivars used for this purpose are also susceptible to infection.<ref name="PHP"/>


==Description==
==Description==
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The [[Conidium|chlamydospores]] are globose with thick walls. They are either formed from [[hypha]]e or by the modification of hyphal cells. They endure in soils for long periods and act as [[Inoculation|inocula]] in primary infection.<ref name=CA/>
The [[Conidium|chlamydospores]] are globose with thick walls. They are either formed from [[hypha]]e or by the modification of hyphal cells. They endure in soils for long periods and act as [[Inoculation|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/>
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, anamorph and holomorph|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>
The [[Teleomorph, anamorph and holomorph|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 [[Musa (genus)#Banana and plantain cultivar naming|banana cultivars]]:
Four races of this pathogen have been described which attack different [[Musa (genus)#Banana and plantain cultivar naming|banana cultivars]]:
* Race 1 attacks cultivars in the ''Musa'' (AAA group) 'Gros Michel' and caused the 20th century epidemic.<ref name="Dita-et-al-2018" /> It also attacks ''Musa'' (AAB group) 'Pome' and its subgroups, Musa (AAB group) 'Silk' and ''Musa'' (ABB group) 'Pisang Awak'. (See {{section link||Race 1}}.)
* Race 1 attacks cultivars in the ''Musa'' (AAA group) 'Gros Michel' 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 Media|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 ''Musa'' (AAB group) 'Pome' and its subgroups, Musa (AAB group) 'Silk' and ''Musa'' (ABB group) 'Pisang Awak'. (See {{section link||Race 1}}.)
* Race 2 attacks ''Musa'' (ABB group) 'Bluggoe' and its close relatives. (See {{section link||Race 2}}.)
* Race 2 attacks ''Musa'' (ABB group) 'Bluggoe' and its close relatives. (See {{section link||Race 2}}.)
* Race 3 attacks ''[[Heliconia]]'' spp.<ref name="Waite-1963" /> (See {{section link||Race 3}}.)
* 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> (See {{section link||Race 3}}.)
* Race 4 attacks ''Musa'' (AAA group) 'Dwarf Cavendish' as well as the hosts of races 1 and 2.<ref>Crop Protection Compendium 2005 Edition. Fusarium oxysporum f.sp. cubense (Panama disease of banana). (CAB International: Wallingford, UK).</ref><ref>Ploetz RC & Pegg KG (2000). Fusarium wilt. In Jones DR (ed.) Diseases of banana, abaca and enset. (CABI Publishing: Wallingford, UK). p.143-159.</ref> (See {{section link||Race 4}}.)
* Race 4 attacks ''Musa'' (AAA group) 'Dwarf Cavendish' as well as the hosts of races 1 and 2.<ref>Crop Protection Compendium 2005 Edition. Fusarium oxysporum f.sp. cubense (Panama disease of banana). (CAB International: Wallingford, UK).</ref><ref>Ploetz RC & Pegg KG (2000). Fusarium wilt. In Jones DR (ed.) Diseases of banana, abaca and enset. (CABI Publishing: Wallingford, UK). p.143-159.</ref> (See {{section link||Race 4}}.)


==Dispersal==
==Dispersal==
Splash by rainfall, movement of contaminated soil, and movement of contaminated [[cutting (plant)|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 [[vector (epidemiology)|vector]]s.<ref name="Dita-et-al-2018" />
Splash by rainfall, movement of contaminated soil, and movement of contaminated [[cutting (plant)|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 [[vector (epidemiology)|vector]]s.<ref name="Dita-et-al-2018" />

==Symptoms==
[[File:Banana wilt (1919) (20166670479).jpg|thumb|200px|right|Gros Michel trees dying Costa Rica in 1919]]
Infection by ''F. oxysporum ''f. sp.'' cubense'' triggers the self-defense mechanisms of the host plant causing the secretion of a [[gel]]. This is followed by the formation of [[tylose]] in the [[vascular]] vessels which blocks the movement of water and nutrients to the upper parts of the plant.{{cn|date=November 2019}} The tips of the feeder roots are the initial sites of infection which then moves on to the [[rhizome]]. The signs of the disease are most noticeable as a dark stain where the [[Stele (biology)|stele]] joins the [[Cortex (anatomy)|cortex]]. As the disease develops, large portions of the [[xylem]] turn a reddish-brown colour. Externally, the oldest leaves start turning yellow and there is often a longitudinal splitting of the lower part of the outer [[leaf sheath]]s on the [[plant stem#Specialized terms for stems|pseudostem]]. The leaves begin to wilt and may buckle at the base of the petiole. As the disease progresses, younger leaves are affected, turn yellow and crumple and the whole canopy begins to consist of dead or dying leaves.<ref name=PHP/>
The leaf symptoms of Fusarium wilt can be confused with those of [[Banana Xanthomonas wilt|Xanthomonas wilt]]. In plants affected by Fusarium, yellowing and wilting of the leaves typically progresses from the older to the younger leaves.<ref>{{cite web|url=http://www.promusa.org/tiki-index.php?page=Fusarium+wilt|title=Fusarium wilt - Promusa - Mobilizing banana science for sustainable livelihoods|website=promusa.org}}</ref> The wilted leaves may also snap at the petiole and hang down the pseudostem. In plants affected by Xanthomonas, the wilting can begin with any leaf and the infected leaves tend to snap along the leaf blade.

==Management==
As [[fungicide]]s are largely ineffective, there are few options for managing Panama disease.<ref name="Ploetz-Pegg-2000" /> Chemical sterilisation of the soil with [[Bromomethane|methyl bromide]] significantly reduced incidence of the disease but was found to be effective for only three years after which the pathogen had recolonised the fumigated areas.<ref>Herbert, J. A., and Marx, D. 1990. Short-term control of Panama disease in South Africa. Phytophylactica 22:339-340.</ref> Injecting the host plants with [[carbendazim]] and potassium phosphonate appears to provide some control but results have been inconclusive. Heat treatment of soil has also been tried in the Philippines but the pathogen is likely to reinvade the treated area.<ref name=PHP/> The greatest hope for managing this disease in infested soils is the development of [[Genetic engineering|genetic modification]]s that will provide resistant cultivars.<ref>{{cite web|url=http://en.cnki.com.cn/Article_en/CJFDTOTAL-GSKK200903024.htm |title=Progress in protoplast culture and somatic hybridization in banana (''Musa'' spp.) |publisher=En.cnki.com.cn |date= |accessdate=2011-01-03}}</ref><ref>Ortiz, R., Ferris, R. S. B., and Vuylsteke, D. R. 1995. Banana and plantain breeding. Pages 110-146 in: Bananas and Plantains. Gowen, S., ed. Chapman & Hall. London.</ref>
Modified bananas developed in collaboration by Ugandan and Belgian scientists were reported in 2008 to be being grown experimentally in Uganda.<ref>{{cite web|last=Koeppel |first=Dan |url=http://www.the-scientist.com/news/display/54710/ |title=Banana: R.I.P.? |publisher=The Scientist |date=30 May 2008 |accessdate=2011-01-03}}</ref> In Australia the movement, sharing, and sale of [[propagation material]] is heavily restricted, especially between states, to slow down the combined threat of TR4, [[banana bunchy top virus|Bunchy Top]], and [[List of banana and plantain diseases|leaf spot]].<ref name="AusBanGrow-sharing">{{cite web | title=Biosecurity Risk: Sharing bits and suckers - Australian Banana Growers | website=Australian Banana Growers | date=2019-02-01 | url=http://abgc.org.au/biosecurity-risk-sharing-bits-and-suckers/ | access-date=2021-04-11}}</ref> The {{visible anchor|RGA2}} gene is ({{as of|2017|lc=yes}}) totally effective against TR4. It is unexpressed in Cavendish but has been found expressed in the diploid [[Musa acuminata ssp. malaccensis|''M. a.'' ssp. ''malaccensis'']]. It may be possible to produce an expressing Cavendish with [[CRISPR]].<ref name="ISAAA-Dale-2021" />

Several [[bacteria]]lly-derived [[volatile organic compound]]s have been found by Yuan et al 2012 to be selectively toxic to ''Foc'': Various [[alkylated benzene]]s, various [[phenol]]s, various [[naphthalene]]s, [[benzothiazole]], [[2-ethyl-1-hexanol]], [[2-undecanol]], [[2-nonanone]], [[2-decanone]], [[2-undecanone]], [[nonanal]], and [[decanal]].<ref name="Cellini-et-al-2021">{{cite journal | last1=Cellini | first1=Antonio | last2=Spinelli | first2=Francesco | last3=Donati | first3=Irene | last4=Ryu | first4=Choong-Min | last5=Kloepper | first5=Joseph W. | title=Bacterial volatile compound-based tools for crop management and quality | journal=[[Trends in Plant Science]] | publisher=[[Cell Press]] | volume=26 | issue=9 | year=2021 | issn=1360-1385 | doi=10.1016/j.tplants.2021.05.006 | pages=968–983}}</ref>


==Tropical Race 1/TR1==
==Tropical Race 1/TR1==
{{anchor|Tropical Race 1}}Race 1/{{visible anchor|TR1}} is also found in ''[[Paspalum]] [[Paspalum fasciculatum|fasciculatum]]'', ''[[Panicum]] [[Panicum purpurescens|purpurescens]]'', ''[[Ixophorus unisetus]]'', and ''[[Commelina diffusa]]'' in Central America.<ref name="Waite-Dunlap-1953" /> These weeds may be acting as an inoculum source.<ref name="Dita-et-al-2018" /> {{As of|2020|11|13}} TR1 is found in [[Queensland]].<ref name="BizQld-TR4" />
{{anchor|Tropical Race 1}}Race 1/{{visible anchor|TR1}} is also found in ''[[Paspalum]] [[Paspalum fasciculatum|fasciculatum]]'', ''[[Panicum]] [[Panicum purpurescens|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==
{{anchor|Tropical Race 2}}{{As of|2020|11|13}} {{visible anchor|TR2}} is found in [[Queensland]].<ref name="BizQld-TR4" />


==Tropical Race 3/TR3==
==Tropical Race 3/TR3==
{{anchor|Tropical Race 3}}Tropical Race 3/{{visible anchor|TR3}} is a pest of ''[[Heliconia]]'' ornamental flowers.<ref name="Waite-1963" /><ref name="Edel-Hermann-Lecomte-2019" /> 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" /> Now renamed ''Fusarium oxysporum'' f. sp. ''heliconiae''.<ref name="Ploetz-2006" /><ref name="Edel-Hermann-Lecomte-2019" />
{{anchor|Tropical Race 3}}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 (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=[[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 (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" />


==Tropical Race 4/TR4==
==Tropical Race 4/TR4==
{{anchor|Tropical Race 4}}<ref name="Pittaway-et-al-1999" /><ref name="Hennessy-et-al-2005" /> Tropical Race 4/{{visible anchor|TR4}} belongs to [[vegetative compatibility group]] 01213/16. All [[banana cultivar|cultivars]] which are [[Plant disease susceptibility|susceptible]] to Race 1 and Race2 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}}</ref>
{{anchor|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+Business Media|Springer Science and Business Media LLC]] | volume=34 | issue=1 | year=2005 | issn=0815-3191 | doi=10.1071/ap04091 | page=115| s2cid=13097833 }}</ref> Tropical Race 4/{{visible anchor|TR4}} belongs to [[vegetative compatibility group]] 01213/16. All [[banana cultivar|cultivars]] which are [[Plant disease susceptibility|susceptible]] to Race 1 and Race2 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}}</ref>
===History of geographic spread===
====1997====
* The first detection in [[Australia]] occurs near [[Darwin, Northern Territory]].<ref name="BizQld-TR4" /><ref name="QldAgDept" />
====2005====
*''[[Chloris (plant)|Chloris]] [[Chloris inflata|inflata]]'', ''[[Euphorbia heterophylla]]'', ''[[Cyanthillium cinereum]]'', and ''[[Tridax procumbens]]'', in banana farms in Australia known to be infected with TR4.<ref name="Hennessy-et-al-2005" /> These weeds may be acting as sources of [[Propagule|inoculum]].<ref name="Dita-et-al-2018" />
====2010====
* Starting here TR4 spread from its origin in southeast Asia westward into [[Vietnam]], [[Laos]], [[Myanmar]], [[India]], [[Pakistan]], [[Oman]], [[Jordan]], [[Lebanon]], [[Israel]], and [[Mozambique]].<ref name="Dita-et-al-2018" />
====2015====
* March 26:<ref name="AusGrowers-false-pos" /> Sample taken from a suspect tree on a farm in [[Mareeba]],<ref name="AusGrowers-false-pos" /> [[Queensland]], [[Australia]].<ref name="Aus-biosecurity-awards-2020" />
* March 28: TR4 was detected in those samples.<ref name="AusGrowers-false-pos" /> The [[Far North Queensland|far north of Queensland]] producing 95% of the [[Australian dollar|$]]580 million of bananas Australia turns out every year, this was immediately taken seriously by [[Biosecurity Queensland]]'s [[Panama TR4 Program]] and the [[Australian Banana Growers' Council]].<ref name="Aus-biosecurity-awards-2020" />
:* However, the test used (the Dita PCR)<ref name="Dita-et-al-2010" /><ref name="Dita-et-al-2010-corrigendum" /> was questioned after this detection was reported, and is no longer used by BQ.<ref name="AusGrowers-false-pos" /> A dissection of the tree trunk showed none of the usual symptoms of ''[[Fusarium]]'' infiltration.<ref name="AusGrowers-false-pos" />
* May 18: Another 18 samples (of unspecified individual(s) from the same farm) tested negative.<ref name="AusGrowers-false-pos" />
:* BQ decided to only use [[vegetative compatibility group]]s and [[molecular diagnostics]] from then on.<ref name="AusGrowers-false-pos" />
* First legitimate detection later in the year in [[Tully, Queensland]].<ref name="BizQld-TR4" /><ref name="QldAgDept" />
====2017====
* [[Queensland University of Technology]] researchers located a gene ([[RGA2]]) which is present but [[gene expression|unexpressed]] in Cavendish, which they found expressed and effective against TR4 in the diploid [[Musa acuminata ssp. malaccensis|''M. a.'' ssp. ''malaccensis'']].<ref name="ISAAA-Dale-2021">{{cite web | title=Researchers Develop Cavendish Bananas Resistant to Panama Disease | website=ISAAA ([[International Service for the Acquisition of Agri-biotech Applications]]) Crop Biotech Update | date=2021-02-24 | url=http://www.isaaa.org/kc/cropbiotechupdate/article/default.asp?ID=18606 | access-date=2021-09-02}}</ref>

====2018====
* {{As of|2018|7|13}} TR4 is found widely throughout [[Southeast Asia]].<ref name="QldAgDept" /><ref name="Dita-et-al-2018" />

====2020====
* {{As of|November 2020}} TR4 was still only found on five plantations in the same area due to this government and industry response.<ref name="Aus-biosecurity-awards-2020" /> Queensland's response to TR4 has been of interest to countries in [[Latin America]] as TR4 moves into that region for the first time.<ref name="Aus-biosecurity-awards-2020">{{cite web|title=Australian Biosecurity Awards 2020 Round 2 award recipients|date=November 2020|website=Australian [[Department of Agriculture, Water and the Environment]]|url=http://www.agriculture.gov.au/sites/default/files/documents/biosecurity-awards-2020-round-2.pdf}}</ref>
* {{As of|2020|11|13}} TR4 is found in [[Laos]], [[Vietnam]], [[Taiwan]], [[Malaysia]], [[Borneo]], [[Indonesia]], [[mainland China]], [[Philippines]], [[Jordan]], [[Mozambique]], [[Pakistan]], [[Lebanon]], [[Oman]], [[India]], and [[North Queensland]] in Australia.<ref name="BizQld-TR4" />
====2021====
* In April the first detection in [[Peru]] occurred, in the [[Department of Piura]], and on the 11th the [[National Service for Agricultural Health (Peru)|National Service for Agricultural Health]] declared a phytosanitary emergency for the whole country. [[Ecuador]] had already been inspecting imports at the border due to Colombia's detection, and then increased inspections further in April.<ref name="first-in-Peru-IPPC">{{cite web | title=Devastating banana disease reported in Peru| website=[[International Plant Protection Convention]] (IPPC) | publisher=UN [[Food and Agriculture Organization|FAO]] | date=2021-04-28 | url=http://www.ippc.int/en/news/devastating-banana-disease-reported-in-peru/ | access-date=2021-05-10}}</ref><ref name="first-in-Peru-SENASA">{{cite web | title=SENASA confirma brote de Fusarium Raza 4 Tropical en Piura | website=[[National Service for Agricultural Health (Peru)|National Service for Agricultural Health]] | publisher=[[Government of Peru]] | date=1999-02-22 | url=http://www.gob.pe/institucion/senasa/noticias/429832-senasa-confirma-brote-de-fusarium-raza-4-tropical-en-piura | language=es | access-date=2021-05-10}}</ref> The appearance of TR4 in Peru threatens its {{convert|170,000|ha|acre}} of plantations.<ref name="first-in-Peru-IICA">{{cite web | title=Alarm bells sound in Peru and Ecuador amidst the banana "pandemic": experts call for public-private cooperation to battle the scourge | website=[[Inter-American Institute for Cooperation on Agriculture]] | date=2021-04-20 | url=http://www.iica.int/en/press/news/alarm-bells-sound-peru-and-ecuador-amidst-banana-pandemic-experts-call-public-private | access-date=2021-05-10}}</ref>



==Subtropical Race 4/STR4==
==Subtropical Race 4/STR4==
{{anchor|Subtropical Race 4}}Subtropical Race 4/{{visible anchor|STR4}} is a subtropical race and does not become symptomatic on [[Cavendish banana|Cavendish]] until the trees are stressed by cold.<ref name="QldAgDept" /> {{As of|2018|7|13}} quarantines for STR4 are imposed over [[South East Queensland]], northern [[New South Wales]], and [[Western Australia]].<ref name="QldAgDept" /> {{As of|2020|11|13}} STR4 is found in South East Queensland.<ref name="BizQld-TR4" /> 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" />
{{anchor|Subtropical Race 4}}Subtropical Race 4/{{visible anchor|STR4}} is a subtropical race and does not become symptomatic on [[Cavendish banana|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=[[Department of Agriculture and Fisheries (Queensland)|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> 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==
==Research==
Early research into ''Foc'' often was conducted by large companies with a financial interest in banana productivity, especially the [[United Fruit Company]]. Among UFC's notable pathologists was [[Frederick Wellman]] in the 1920s.<ref name="McCook-Peterson-2020">{{cite journal | last1=McCook | first1=Stuart | last2=Peterson | first2=Paul D. | title=The Geopolitics of Plant Pathology: Frederick Wellman, Coffee Leaf Rust, and Cold War Networks of Science | journal=[[Annual Review of Phytopathology]] | publisher=[[Annual Reviews (publisher)|Annual Reviews]] | volume=58 | issue=1 | date=2020-08-25 | issn=0066-4286 | doi=10.1146/annurev-phyto-082718-100109 | pages=181–199| pmid=32853100 }}</ref> 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 group]]s 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> The resistance of different banana cultivars to the pathogen is under scrutiny.<ref>{{cite journal |url=http://elfosscientiae.cigb.edu.cu/PDFs/BA/2006/23/2/BA002302RP153-157.pdf |journal=Biotecnología Aplicada |date=2006 |volume=23 |issue=2 |title=Differentiating resistance to ''Fusarium oxysporum'' f.sp. ''cubense'' strain 1 culture filtrates in banana leaves |accessdate=2011-01-03 |archiveurl=https://web.archive.org/web/20110718163159/http://elfosscientiae.cigb.edu.cu/PDFs/BA/2006/23/2/BA002302RP153-157.pdf |archivedate=2011-07-18 |vauthors=Companioni B, Mora N, Díaz L, Pérez A, Arzola M, Espinosa P, Hernández M, de la Caridad Ventura J, Pérez MC, Santos R, Lorenzo JC}}</ref>
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 group]]s 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 [[Phylogenetics|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 [[Monophyly|monophyletic]] and appears to have multiple [[evolution]]ary 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>
Research into the [[Phylogenetics|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 [[Monophyly|monophyletic]] and appears to have multiple [[evolution]]ary 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" />
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>


==See also==
==See also==
Line 119: Line 62:


==References==
==References==
{{Reflist|refs=
{{Reflist}}

<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>

<ref name="Staver-et-al-2020">{{cite journal | last1=Staver | first1=Charles | last2=Pemsl | first2=Diemuth E. | last3=Scheerer | first3=Lars | last4=Perez Vicente | first4=Luis | last5=Dita | first5=Miguel | title=Ex Ante Assessment of Returns on Research Investments to Address the Impact of Fusarium Wilt Tropical Race 4 on Global Banana Production | journal=[[Frontiers in Plant Science]] | publisher=[[Frontiers Media|Frontiers]] | volume=11 | date=2020-07-06 | page=844 | issn=1664-462X | doi=10.3389/fpls.2020.00844 | pmid=32733497 | pmc=7357546 | doi-access=free}}</ref>

<ref name="AusGrowers-false-pos">{{cite web | title=TR4 testing under the microscope - Australian Banana Growers | website=[[Australian Banana Growers Council]] | date=2015-10-03 | url=http://abgc.org.au/2015/10/03/tr4-testing-under-the-microscope/ | access-date=2021-02-06}}</ref>

<ref name="Dita-et-al-2010">{{cite journal | last1=Dita | first1=M. A. | last2=Waalwijk | first2=C. | last3=Buddenhagen | first3=I. W. | last4=Souza Jr | first4=M. T. | last5=Kema | first5=G. H. J. | title=A molecular diagnostic for tropical race 4 of the banana fusarium wilt pathogen | journal=[[Plant Pathology (journal)|Plant Pathology]] | publisher=[[Wiley (publisher)|Wiley]] | volume=59 | issue=2 | year=2010 | issn=0032-0862 | doi=10.1111/j.1365-3059.2009.02221.x | pages=348–357| doi-access=free }}</ref>

<ref name="Dita-et-al-2010-corrigendum">{{cite journal | last1=Dita | first1=M. A. | last2=Waalwijk | first2=C. | last3=Buddenhagen | first3=I. W. | last4=Souza Jr | first4=M. T. | last5=Kema | first5=G. H. J. | title=A molecular diagnostic for tropical race 4 of the banana fusarium wilt pathogen - Corrigendum | journal=[[Plant Pathology (journal)|Plant Pathology]] | publisher=[[Wiley (publisher)|Wiley]] | volume=60 | issue=2 | date=2011-03-01 | issn=0032-0862 | doi=10.1111/j.1365-3059.2010.02417.x | pages=384| doi-access=free }}</ref>

<ref name="BizQld-TR4">{{cite web | archive-url=https://web.archive.org/web/20200418105822/https://www.business.qld.gov.au/industries/farms-fishing-forestry/agriculture/crop-growing/priority-pest-disease/panama-disease | archive-date=18 April 2020 | title=Panama disease tropical race 4 (TR4) | website=Business Queensland, [[Queensland Government]] | date=2020-11-13 | url=http://www.business.qld.gov.au/industries/farms-fishing-forestry/agriculture/crop-growing/priority-pest-disease/panama-disease | access-date=2021-02-06}}</ref>

<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=[[Department of Agriculture and Fisheries (Queensland)|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="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 Media|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>

<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 (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>

<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 (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="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="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=[[Centre for Agriculture and Bioscience International]] | publication-place=[[Wallingford, Oxfordshire]], [[England]], [[United Kingdom]] | last1=Ploetz | first1=R.C. | last2=Pegg | first2=K.G.}}</ref>

<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>

<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+Business Media|Springer Science and Business Media LLC]] | volume=34 | issue=1 | year=2005 | issn=0815-3191 | doi=10.1071/ap04091 | page=115| s2cid=13097833 }}</ref>

<ref name="Stover-Simmonds-1987">{{cite book | editor-last1=Stover | editor-first1=Robert H. | editor-last2=Simmonds | editor-first2=N. W. | title=Bananas | series=Tropical Agriculture Series | publisher=Longman Scientific & Technical [[Wiley (publisher)|Wiley]] | publication-place=[[Harlow]], [[Essex]], [[England]]/[[New York City]] | date=August 10, 1987 | isbn=978-0-470-20684-3 | oclc=13008247 | language=en | page=1-468 | edition=3}}</ref>

}}


==Further reading==
==Further reading==

Revision as of 18:47, 24 November 2021

Fusarium oxysporum f.sp. cubense
Fusarium oxysporum f.sp. cubense race 1 growing for several days on brown rice
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Fungi
Division: Ascomycota
Class: Sordariomycetes
Order: Hypocreales
Family: Nectriaceae
Genus: Fusarium
Species:
F. oxysporum
Subspecies:
F. o. f.sp. cubense
Trinomial name
Fusarium oxysporum f.sp. cubense
Synonyms

Fusarium oxysporum f. sp. cubense Pronunciationⓘ is a fungal plant pathogen that causes Panama disease of banana (Musa spp.), also known as fusarium wilt of banana.

Description

Fusarium oxysporum is a common inhabitant of soil and produces three types of asexual spores; macroconidia, microconidia and chlamydospores.[1]

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.[2]

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.[2]

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.[2]

The macroconidia and chlamydospores are normally only formed on dead or dying host plants. Chlamydospores are the most significant survival structures of this pathogen.[3]

The teleomorph or sexual reproductive stage of F. oxysporum is unknown.[4]

Four races of this pathogen have been described which attack different banana cultivars:

  • Race 1 attacks cultivars in the Musa (AAA group) 'Gros Michel' and caused the 20th century epidemic.[5] It also attacks Musa (AAB group) 'Pome' and its subgroups, Musa (AAB group) 'Silk' and Musa (ABB group) 'Pisang Awak'. (See § Race 1.)
  • Race 2 attacks Musa (ABB group) 'Bluggoe' and its close relatives. (See § Race 2.)
  • Race 3 attacks Heliconia spp.[6] (See § Race 3.)
  • Race 4 attacks Musa (AAA group) 'Dwarf Cavendish' as well as the hosts of races 1 and 2.[7][8] (See § Race 4.)

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.[5]

Tropical Race 1/TR1

Race 1/TR1 is also found in Paspalum fasciculatum, Panicum purpurescens, Ixophorus unisetus, and Commelina diffusa in Central America.[9] These weeds may be acting as an inoculum source.[5]

Tropical Race 3/TR3

Tropical Race 3/TR3 is a pest of Heliconia ornamental flowers.[6][10] Formerly reported to be a lesser pest of Musa balbisiana seedlings and of Gros Michel, but that is no longer thought to be true.[10][11] Now renamed Fusarium oxysporum f. sp. heliconiae.[12][10]

Tropical Race 4/TR4

[13][14] Tropical Race 4/TR4 belongs to vegetative compatibility group 01213/16. All cultivars which are susceptible to Race 1 and Race2 are susceptible to TR4 (see § Race 1 and § Race 2).[15]

Subtropical Race 4/STR4

Subtropical Race 4/STR4 is a subtropical race and does not become symptomatic on Cavendish until the trees are stressed by cold.[16] Also found in Paspalum spp. and Amaranthus spp. in Australia.[13] These weeds may be acting as sources of inoculum.[5]

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.[17]

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.[18] The largest lineages of F. oxysporum f. sp. cubense (§ Race 1 and § Race 2) are genetically distinct from a lineage originating from East Africa (§ Race 5) and developed pathogenicity for bananas independently from one another.[19]

Identification, differentiation, and usage of vegetative compatibility groups is useful and valid within Foc because there are relatively few VCGs.[20]

See also

References

  1. ^ "Fusarium Wilt of Chickpea". Pests and Diseases Image Library. Archived from the original on 2011-03-21. Retrieved 2011-01-03.
  2. ^ a b c 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
  3. ^ Ploetz, Randy C. (2000). "Panama Disease:A Classic and Destructive Disease of Banana". Plant Health Progress. 1. American Phytopathological Society: 10. doi:10.1094/PHP-2000-1204-01-HM. S2CID 12228953.
  4. ^ Leslie JF, Summerell BA (2006) The Fusarium Laboratory manual. (Blackwell Publishing: Iowa, USA)
  5. ^ a b c d Dita, Miguel; Barquero, Marcia; Heck, Daniel; Mizubuti, Eduardo S. G.; Staver, Charles P. (2018-10-19). "Fusarium Wilt of Banana: Current Knowledge on Epidemiology and Research Needs Toward Sustainable Disease Management". Frontiers in Plant Science. 9. Frontiers: 1468. doi:10.3389/fpls.2018.01468. ISSN 1664-462X. PMC 6202804. PMID 30405651.
  6. ^ a b Waite, B.H (1963). "Wilt of Heliconia spp. caused by Fusarium oxysporum f. sp. cubense Race 3". Tropical Agriculture (Trinidad). 40: 299–305.
  7. ^ Crop Protection Compendium 2005 Edition. Fusarium oxysporum f.sp. cubense (Panama disease of banana). (CAB International: Wallingford, UK).
  8. ^ Ploetz RC & Pegg KG (2000). Fusarium wilt. In Jones DR (ed.) Diseases of banana, abaca and enset. (CABI Publishing: Wallingford, UK). p.143-159.
  9. ^ Waite, B.H.; Dunlap, V.C. (1953). "Preliminary host range studies with Fusarium oxysporum f. sp. cubense". Plant Disease Reporter. 37. United States Department of Agriculture: 79–80.
  10. ^ a b c Edel-Hermann, V.; Lecomte, C. (2019). "Current Status of Fusarium oxysporum Formae Speciales and Races". Phytopathology. 109 (4). American Phytopathological Society: 512–530. doi:10.1094/phyto-08-18-0320-rvw. ISSN 0031-949X. PMID 30461350.
  11. ^ Ploetz, R.C.; Pegg, K.G. (2000). Jones, D. R. (ed.). Diseases of Banana, Abacá, and Enset. Wallingford, Oxfordshire, England, United Kingdom: Centre for Agriculture and Bioscience International. p. 143-159. ISBN 978-0-85199-355-3. OCLC 41347037.
  12. ^ Ploetz, Randy C. (2006). "Fusarium Wilt of Banana Is Caused by Several Pathogens Referred to as Fusarium oxysporum f. sp. cubense". Phytopathology. 96 (6). American Phytopathological Society: 653–656. doi:10.1094/phyto-96-0653. ISSN 0031-949X. PMID 18943184.
  13. ^ a b Pittaway, P. A.; Nasir, Nasril; Pegg, K. G. (1999). "Soil receptivity and host - pathogen dynamics in soils naturally infested with Fusarium oxysporum f. sp. cubense, the cause of Panama disease in bananas". Australian Journal of Agricultural Research. 50 (4). CSIRO Publishing (Commonwealth Scientific and Industrial Research Organisation): 623. doi:10.1071/a98152. ISSN 0004-9409.
  14. ^ Hennessy, Chelsea; Walduck, Geoff; Daly, Andrew; Padovan, Anna (2005). "Weed hosts of Fusarium oxysporum f. sp. cubense tropical race 4 in northern Australia". Australasian Plant Pathology. 34 (1). Springer Science and Business Media LLC: 115. doi:10.1071/ap04091. ISSN 0815-3191. S2CID 13097833.
  15. ^ Molina, A.B.; Fabregar, E.; Sinohin, V.G.; Yi, G.; Viljoen, A. (2009). "Recent occurrence of Fusarium oxysporum f.sp. cubense tropical race 4 in Asia". Acta Horticulturae (828). International Society for Horticultural Science (ISHS): 109–116. doi:10.17660/actahortic.2009.828.10. hdl:2263/12191. ISSN 0567-7572.
  16. ^ "Panama disease tropical race 4 (TR4) - Business Queensland". Queensland Agriculture Department. 2020-11-13. Archived from the original on 22 December 2018. Retrieved 2021-02-06.
  17. ^ Fourie, G.; Steenkamp, E. T.; Gordon, T. R.; Viljoen, A. (2009). "Evolutionary Relationships among the Fusarium oxysporum f. sp. cubense vegetative compatibility groups". Applied and Environmental Microbiology. 75 (14): 4770–81. Bibcode:2009ApEnM..75.4770F. doi:10.1128/AEM.00370-09. PMC 2708428. PMID 19482953.
  18. ^ O'Donnell, K; Kistler, H. C; Cigelnik, E; Ploetz, R. C (1998-03-03). "Multiple evolutionary origins of the fungus causing Panama disease of banana: Concordant evidence from nuclear and mitochondrial gene genealogies". Proceedings of the National Academy of Sciences. 95 (5): 2044–2049. Bibcode:1998PNAS...95.2044O. doi:10.1073/pnas.95.5.2044. PMC 19243. PMID 9482835.
  19. ^ Koenig, R L; et al. (1997). "Fusarium oxysporum f. sp. cubense consists of a small number of divergent and globally distributed clonal lineages". Phytopathology. 87 (9): 915–923. doi:10.1094/phyto.1997.87.9.915. PMID 18945062.
  20. ^ Moore, N.Y.; BENTLEY, S.; BUDDENHAGEN, I.W.; PEGG, K.G. (2001). B.A. Summerell; J.F. Leslie; D. Backhouse; W.L. Bryden; L.W. Burgess (eds.). "Fusarium wilt of banana, a diverse clonal pathogen of a domesticated clonal host". Alianza SIDALC. Fusarium – Paul E. Nelson Memorial Symposium. Saint Paul, Minnesota: American Phytopathological Society Press: 212–224. Retrieved 2021-01-05.

Further reading

External links

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