Fusarium oxysporum f. sp.

cubense
Fusarium oxysporum f. sp. cubense (Foc) is one of
Foc at a glance
more than 100 formae speciales (special forms) of the
Fusarium oxysporum species complex of pathogenic as well
as non-pathogenic morphologically similar lamentous
fungi. The cubense special form comprises the pathogenic
strains that cause Fusarium wilt in cultivated bananas, as
well as strains that aect species in the Musaceae and
Heliconeaceae families. However, it does not follow that the
strains of F. oxysporum f. sp. cubense are necessarily
[1]
related genetically . F. oxysporum has no known sexual
Name of pathogen
stage. Variation in the fungus is thought to result from
Fusarium oxysporum f. sp. cubense
mutations.
Disease
The pathogenic strains are commonly classied into four
Fusarium wilt
races that are meant to reect dierences in the cultivars
Distribution
on which they cause disease. In reality, there are more
variants of the fungus than the number of races suggests. Race 1: pan-tropical
TR4: China, Philippines, Indonesia,
Progress in understanding the pathogen's diversity was
Malaysia, Australia, Jordan, Oman,
made possible with the development of a method to
Lebanon, Pakistan and Mozambique
classify Fusarium oxysporum strains into vegetative
compatibility groups (VCGs), based on the ability of their
hyphae to fuse and form stable heterokaryons (cells
containing two distinct nuclei)[2]. Each VCG has its own
characteristics in terms of aggressiveness and the banana
cultivars that it attacks most readily. The VCG associated
with tropical race 4 (TR4) VCG01213/16 was ranked as
the greatest threat to banana production because of its
lethal impact, wide host range and persistence in the soil[3].

Contents
History
Disease cycle
Races
Race 1
Race 2
Race 3
Race 4
Vegetative compatibility
References
See also on this website
Further reading
External links
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History
In 1876, the author of the rst report of the disease, J. Bancroft, speculated that it was caused by a
fungus[4]. Unaware of Bancroft's work, a horticulturist at the Hawaii Agricultural Experiment Station
[5]
also proposed in 1904 that a fungus caused the disease . The plant pathologist Erwin F. Smith
became the rst person to isolate the pathogen from banana tissues he had received from Cuba[6].
He reported his results at the rst meeting of the American Phytopathological Society held in
Boston in 1908[7]. Smith recognized that the fungus was in the genus Fusarium and named it
Fusarium cubense because of its Cuban source. The study conrming that the fungus was indeed
the causal agent was published in 1919[8]. Fusarium cubense was recognized as a variant of
Fusarium oxysporum and renamed Fusarium oxysporum f. sp. cubense in 1935[9].

Disease cycle
No sexual stage has been observed. Inside the plant, the fungus produces three types of asexual
spores: microconidia, macroconidia and chlamydospores.
Chlamydospores are round, thick-walled resting propagules that are produced by the dying
banana plant. They can persist in soil for indenite periods of time (as long as 30 years or more).
Infection is initiated when they germinate in response to exudates from the roots and hyphae (long
and branching lamentous structures collectively known as mycelium) penetrate the lateral roots.
Microconidia are one or two celled and oval- to kidney-shaped. They are the type of spore most
frequently produced within the vessels of infected plants.
Macroconidia are four to eight celled, sickle-shaped, thin-walled and delicate. These spores are
commonly found on the surface of plants killed by the fungus.
Australian scientists have established that 20 minutes in 65 C water is the minimum condition for
killing race 4 hyphae, microconidia and macroconidia in banana plant tissue, whereas
[10]
chlamydospores require autoclaving .
It's still not clear how the fungus kills the plant. One hypothesis is fungus releases toxins into the
plant, killing its cells. Another is that the fungus tricks the plant into killing its cells by triggering the
natural process of programmed cell death that eliminates unwanted, damaged or used cells. The
decaying plant tissues would then be used by the fungus as a food source[11].
The hypothesis is being tested in a banana that has been genetically modied to prevent the
fungus from co-opting the plants cell death pathways. Deprived of food, the fungus eventually
stops growing and fails to colonize and infect the tissues of the transgenic plant[12].

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Microconidia (Photo by Miguel Dita).

Macroconidia (Photo by Miguel Dita).

Chlamydospores (Photo by Miguel Dita).

Races
The pathogenic strains of Foc are classied into races based on the dierential response of
cultivars. Traditionally, four races are recognized, although certain situations suggest that more
races may exist[13]. Extensive inoculation studies are needed to clearly dene the various
pathotypes, but these are expensive and time-consuming. The results may also be equivocal
because of variability in growing conditions and/or in planting material. The race concept has been
criticized for being an imperfect measure of pathogenic diversity and for not reecting genetic
relationships, but is nonetheless considered useful to describe host reaction and new disease
outbreaks.

Race 1
Race 1 strains cause disease in Gros Michel, Silk, Pome and Pisang awak cultivars, among others.

Race 2
Race 2 strains prey on Bluggoe and closely related cooking cultivars.

Race 3
Race 3 was reported to aect Heliconia species, and to a lesser extent Gros Michel and seedlings of
Musa balbisiana[14]. However, the disease has not been reported since[15].

Race 4
Race 4 was originally coined to designate the strains that attack Cavendish cultivars. Prior to the
1990s, symptoms of Fusarium wilt on Cavendish cultivars had been observed in the subtropical
growing areas of Australia, Canary Islands and South Africa[16], as well as in some tropical growing
areas such as Taiwan[17], Jamaica and Guadeloupe. Circumstantial evidence suggested that, with the
exception of Taiwan, the limited damage was due to predisposing factors: low temperatures in the
subtropics and edaphic factors in the tropics[18]. Originally classied as race 4, these pathogenic
isolates were later reclassied as subtropical race 4 (STR4) to distinguish them from the isolates
that cause Fusarium wilt in the tropics in the absence of predisposing factors, which then became
known as tropical race 4 (TR4). The 01213/16 VCG (see below) associated with TR4 was rst
identied in isolates from Taiwan[19]. In India, symptoms of Fusarium wilt have also been observed
on the Cavendish cultivar 'Grande Naine' in the absence of predisposing factors, except that the
isolate was not associated with either STR4 or TR4[20].
In addition to Taiwan, TR4 has been reported in peninsular Malaysia[21] and Sarawak, Indonesia
(Papua province[22], Kalimantan[23], Halmahera, Java, Sulawesi and Sumatra), mainland China
Page 3 / 7

[24]

[25]

[26]

(Guangdong , Hainan , Guangxi, Fujian and Yunnan), the Philippines and Australia (Northern
Territory in 1997[27] and Queensland in 2015[28]). In 2013, it was reported to be in Jordan where it
had been present since 2006 according to the disease report[29] and has since been observed in at
[30]
[31]
least one more production area and in Mozambique , where it might have arrived 2-3 years
earlier[32] and has since been observed in a second plantation[33]. In 2015, it was reported to be in
Lebanon and Pakistan[34][35]. There are also reports that it is in Oman[36].
TR4 also attacks cultivars susceptible to races 1 and 2, and additional cultivars such as Barangan
(Lakatan subgroup, AAA genome group)[23] and Pisang Mas. The wide host range of TR4 makes it
dicult to diagnose outbreaks. For example, while a Cavendish with Fusarium wilt would
immediately raise alarm, a Gros Michel infected with TR4 would not because the assumption would
be it is infected with a race 1 strain.

Vegetative compatibility
Vegetative compatibility is used to classify into
vegetative compatibility groups (VCGs) isolates that
share the same form (allele) of the genes that control the
formation of an heterokaryon (a cell with two distinct
[2]
nuclei) . Since alleles at each locus (the location of the
gene on the chromosome) must be identical in order for
isolates to be vegetatively compatible, isolates within a
VCG are assumed to be clonally derived. On the other hand,
since a mutation in one of those genes would make closely
related isolates vegetatively incompatible, isolates that
share a common ancestor could occur in dierent VCGs.

Protocols
Sampling infected plants
Isolating Foc from infected tissues
Determining vegetative
compatability group
Storing Foc isolates
Inoculating plants with Foc
Extraction of Foc DNA

Each VCG is given a four to ve digit code. The rst three

PCR diagnostic tests
numbers refer to the forma specialis to which the strain
Source: FAO
belongs, 012 in the case of the banana-specic f. sp.
cubense. The last number represents the order in which the
VCGs were identied (0120, 0121, 0122 etc.). The race 1
isolate used to designate the rst VCG0120 came from
Australia[2]. Some 24 VCGs have since been characterized
worlwide.
The region with the greatest diversity of VCGs is Asia. A series of surveys documented 11 VCGs in
the ve banana-producing regions of mainland China[37], 3 VCGs and 4 isolates of unknown VCG in
Indonesia's Lampung province[38], and 5 VCGs and 7 isolates of unknown VCG in Indonesia's West
Sumatera province[39]. A survey of 9 Asian countries reported 12 VCGs[40].
Tropical race 4 isolates belong to the VCG01213/16 complex, although other VCGs are also known
to cause Fusarium wilt on Cavendish cultivars: the so-called STR4 strains that attack Cavendish
bananas in the subtropics (0120, 0121, 0129 and 01211 in Australia; 0120 in South Africa and the
Canary Islands and 0122 in the Philippines)[41]. In India, VCG0124 isolates have also been extracted
from Cavendish bananas exhibiting Fusarium wilt symptoms[20]. In Latin America, VCG0124 isolates
are classied as race 1 because they infect Gros Michel but not Cavendish bananas.

Page 4 / 7

References
1. Ploetz, R.C. and Pegg, K.G. 1997. Fusarium wilt of banana and Wallace's line: was the disease originally
restricted to his Indo-Malayan region? Australasian Plant Pathology 26(4):239-249.
2. Puhalla, J.E. 1985. Classication of strains of Fusarium oxysporum on the basis of vegetative compatibility.
Canadian Journal of Botany, 63(2):179-183.
3. Ploetz, R.C. 2009. Assessing threats posed by destructive banana pathogens. Proceedings of the
International ISHS-ProMusa Symposium on Recent Advances in Banana Crop Protection for Sustainable
Production and Improved Livelihoods held in White River, South Africa, 10-14 September 2007. Jones, D.R. and
Van den Bergh, I. (eds). Acta Horticulturae 828:245-252.
4. Bancroft, J. 1876. Report of the board appointed to inquire into the cause of disease aecting livestock and
plants. In: Votes and Proceedings 1877, Vol 3, Queensland, pp. 1011-1038
5. Higgins, J.E. 1904. The banana in Hawaii. Bulletin No. 7, Hawaii Agricultural Experiment Station, University of
Hawaii. Honolulu, Hawaii.
6. Smith, E.F. 1910. A Cuban banana disease. Science 31: 754-755.
7. Ploetz, R.C. 2005. Panama disease: an old nemesis rears its ugly head: Part 1. The beginnings of the banana
export trades APSnet Feature:13.
8. Brandes, E.W. 1919. Banana wilt. Phytopathology 9: 339-389.
9. Wollenweber, H.W., Reinking, O.W. 1935. Die Fusarien. Berlin, DE. Paul Parey. 355 p.
10. Walduck, G. and Daly, A. 2007. Banana Tropical Race 4 Management Disease Management. Pp. 7-11 In
Northern Territory Department of Primary Industry, Fisheries and Mines. Australia. www.nt.gov.au
11. Dickman, M.B. 2004. Can model plants help banana improvement through biotechnology? InfoMusa,
France, 13(2):6-11.
12. Paul, J-Y., Becker, D., Dickman, M.B., Harding,R., Khanna, H. and Dale, J. 2011. Apoptosis-related genes
confer resistance to Fusarium wilt in transgenic 'Lady Finger' bananas. Plant Biotechnology Journal
9(9):1141-1148.
13. Ploetz, R.C., Pegg, K.G. 2000. Fusarium wilt. pp. 143-159. In: Jones, D.R. (ed.) Diseases of Banana, Abac
and Enset. CABI Publishing, Wallingford, UK.
14. Waite, B.H. 1963. Wilt of Heliconia spp. caused by Fusarium oxysporum f. sp. cubense Race 3. Tropical
Agriculture (Trinidad) 40:299-305.
15. Ploetz, R.C. and Pegg, K.G. 2000.Fungal diseases of the root, corm and pseudostem: Fusarium wilt. p.
143-159. In: Jones, D.R. (ed.) Diseases of Banana, Abac and Enset. CABI Publishing, Wallingford, UK.
16. Ploetz, R.C., Herbert, J., Sebasigari, K., Hernandez, J.H., Pegg, K.G., Ventura, J.A. and Mayato, L.S. 1990.
Importance of fusarium wilt in dierent banana-growing regions. p.9-26. In: Ploetz, R.C. (ed.). Proceedings of
International Conference on Fusarial Wilt of Banana, Miami (USA), 1989/08/27-30. Fusarium wilt of banana.
American Phytopathological Society (APS Press), St Paul (USA).
17. Sun, E.J., Su, H.J. and Ko, W.H. 1978. Identication of Fusarium oxysporum f. sp. cubense Race 4 from Soil
or Host Tissue by Cultural Characters. Phytopathology 68:1672-1673.
18. Ploetz, R.C. 2005. Panama disease: an old nemesis rears its ugly head: Part 2: the 'Cavendish' era and
beyond. APSnet Feature:13.
19. Buddenhagen, I. 2009. Understanding strain diversity in Fusarium oxysporum f. sp. cubense and history of
introduction of 'Tropical Race 4' to better manage banana production. Acta Horticulturae 828:193-204.
20. A new threat to Cavendish bananas? in the March 2011 issue of InfoMus@
21. Ong Kim Pin. 1996. Fusarium wilt of Cavendish banana in a commercial farm in Malaysia. p.211-217. In:
Frison, E.A., Horry, J. and De Waele, D. (eds.). Proceedings of New Frontiers in Resistance Breeding for
Nematode, Fusarium and Sigatoka, Kuala Lumpur (MYS), 1995/10/2-5. New frontiers in resistance breeding for
nematode, Fusarium and Sigatoka. INIBAP, Montpellier (FRA).
22. Davis, R.I., Moore, N.Y., Bentley, S., Gunua, T.G. and Rahamma, S. 2000. Further records of Fusarium
oxysporum f. sp. cubense from New Guinea. Australasian Plant Pathology 29(3):224.
23. Hermanto, C., Sutanto, A., HS, E., Daniells, J.W., O'Neill, W.T., Sinohin, V.G.O., Molina, A.B. and Taylor, P..
2011. Incidence and Distribution of Fusarium Wilt Disease of Banana in Indonesia. Proceedings of the
Page 5 / 7

International ISHS-ProMusa Symposium on Global Perspectives on Asian Challenges held in Guangzhou, China,
14-18 September 2009. Van den Bergh, I., Smith, M. and Swennen, R. (eds). Acta Horticulturae 897:313-322.
24. Qi, P. 2001. Status report of banana Fusarium wilt disease in china. p.119-120. In: Molina, A.B., Nik Masdek,
N.H. and Liew, K.W. (eds.). Proceedings of International Workshop on the Banana Fusarium Wilt Disease,
Genting Highlands Resort, 1999/10/18-20. Banana Fusarium wilt management: Towards sustainable
cultivation. INIBAP, Los Banos, Philippines.
25. Qi, Y.X., Zhang, X., Pu, J.J., Xie, Y.X., Zhang, H.Q. and Huang, S.L. 2008. Race 4 identication of Fusarium
oxysporum f. sp. cubense from Cavendish cultivars in Hainan province, China. Australasian Plant Disease Notes
3(1):46-47
26. Molina, A., Fabregar, E., Sinohin, V.G., Herradura, L., Fourie, G. and Viljoen, A. 2008. Conrmation of
tropical race 4 of Fusarium oxysporum f. sp. cubense, infecting Cavendish bananas in the Philippines. Abstract
of presentation to the 2008 Centennial Meeting of the American Phytopathological Society.
27. Conde, B.D. and Pitkethley, R.N. 2001. The discovery, identication and management of banana fusarium
wilt outbreaks in the northern territory of Australia. p.260-265. In: Molina, A.B., Nik Masdek, N.H. and Liew,
K.W. (eds.). Proceedings of International Workshop on the Banana Fusarium Wilt Disease, Genting Highlands
Resort, 1999/10/18-20. Banana Fusarium wilt management: Towards sustainable cultivation. INIBAP, Los
Banos, Philippines.
28. ABGC release on suspected Panama TR4 case posted on 4 March 2015.
29. Garcia, F.A., Ordonez, N., Konkol, J., AlQasem, M., Naser, Z., Abdelwali, M., Salem, N.M., Waalwijk, C.,
Ploetz, R.C. and Kema, G. 2013. First Report of Fusarium oxysporum f. sp. cubense Tropical Race 4 associated
with Panama Disease of banana outside Southeast Asia. Plant Disease.
30. Ploetz, R. et al. 2015. Tropical race 4 of Panama disease in the Middle East. Phytoparasitica, 43:283-293.
31. New banana disease found in Mozambique, 13 Decembre 2013
32. Butler, D. 2013. Fungus threatens top banana. Nature 504(7479):195-196.
33. CNN news published 22 July 2015
34. Ordonez et al. 2015. First report of Fusarium oxysporum f. sp. cubense tropical race 4 causing Panama
disease in Cavendish bananas in Pakistan and Lebanon. Plant disease.
35. Syed, R.N. et al. 2015. First report of panama wilt disease of banana caused by Fusarium oxysporum f. sp.
cubense in Pakistan. Journal of Plant Pathology 1(1):213.
36. News on agricultural quarantine measures on bananas in the 5 September 2012 edition of
muscatdaily.com
37. Li, C.Y., Mostert, G., Zuo, C.W., Beukes, I., Yang, Q.S., Sheng, O., Kuang, R.B., Wei, Y.R., Hu, C.H., Rose, L.,
Karangwa, P., Yang, J., Deng, G.M., Liu, S.W., Gao, J., Viljoen, A. and Yi, G.J. 2013. Diversity and Distribution of
the Banana Wilt Pathogen Fusarium oxysporum F. Sp. cubense in China. Fungal Genomics and Biology 03(02).
38. Riska, J. and A. Soemargono. 2012. Identication and distribution of Fusarium oxysporum f. sp. cubense
isolates through analysis of vegetative compatibility group in Lampung province, Indonesia. Journal of
Engineering and Applied Sciences 7(4):279-284.
39. Riska, J., and C. Hermanto. 2012. Conrm the status of VCG Fusarium oxysporum f. sp. cubense in West
Sumatera Indonesia. Journal of Engineering and Applied Sciences 7(4):244-249.
40. Occurrence of various Vegetative Compatibility Groups (VCGs) of Foc in Asia, poster presented at the 10th
International Congress of Plant Pathology held in China in 2013.
41. Buddenhagen, I. 2009. Understanding strain diversity in Fusarium oxysporum f. sp. cubense and history of
introduction of 'Tropical Race 4' to better manage banana production. p.193-204. In: Jones, D.R. and Van den
Bergh, I. (eds.). Proceedings of International ISHS-ProMusa Symposium on Recent Advances in Banana Crop
Protection for Sustainable Production and Improved Livelihoods, White River, South Africa, 10-14 September
2007. Acta Horticulturae 828. ISHS, Leuven, Belgium.

See also on this website

Photos on the symptoms of Fusarium wilt in the Musarama image bank
Video on the symptoms, transmission and prevention of Fusarium wilt in the Musarama video bank
News and blogs on Fusarium wilt:
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Australia's TR4 incursion one year on

The trap of extinction stories on bananas
Social and psychological impacts of the TR4 incursion in Queensland, Australia
Why screening protocols matter
TR4 quarantine lifted on second Queensland farm
More stories
Musapedia page on an INREF-funded research project managed by Wageningen University &
Research Centre - Panama disease: Multi-level solutions for a global problem

Further reading
Diagnostic manual and links to presentations given at a 2014 FAO-CARDI regional workshop on the
prevention and diagnostic of Fusarium wilt
Contingency plan (in Spanish) on TR4 for OIRSA countries
Fact sheet on Panama disease (8MB PDF) on the Plant Health Australia website
Fusarium wilt of banana laboratory diagnostics manual (1.8MB PDF) on the Plant Health Australia
website
Datasheet on Fusarium oxysporum f. sp. cubense in CABI's Invasive Species Compendium
Panama disease: an old nemesis rears its ugly head, Part 1: The beginnings of the banana export
trades Part 2: the Cavendish era and beyond

External links
Wikipedia page on Fusarium oxysporum f. sp. cubense
Website for the research projects on Fusarium wilt that are managed by Wageningen University &
Research Centre: panamadisease.org
Banana Fusarium wilt in Africa website (under development)
Contributors to this page: Anne Vzina .
Page last modied on Friday, 11 July 2014 11:39:45 CEST by Anne Vzina.
The original document is available at
http://www.promusa.org/Fusarium+oxysporum+f.+sp.+cubense

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