FUSARIUM WILT IN CAVENDISH BANANA CAUSED BY FOC TR4:

A THREAT TO PHILIPPINE BANANA INDUSTRY

CECIRLY G. PUIG
Graduate Student, University of the Philippines Los Baos, Los Baos, Laguna;
A term paper as requirement of the course PPTH 141 Plant Disease Epidemiology.

ABSTRACT

Fusarium wilt is responsible for severe losses of Cavendish

banana in Davao regions banana industry. The disease is also a global
threat to all banana growing areas around the world. With its continued
destruction, it resulted to more than 5,000 hectares abandoned in Davao del
Norte and ComVal alone. The causal organism Fusarium oxysporum f. sp.
cubense Tropical Race 4 (Foc TR4) affects not only the export Cavendish
banana but all banana cultivars growing in the region. Despite many
disease management strategies, control of Fusarium wilt in the region
remains unsuccessful just like the experience of other Asian countries,
Australia, South and Central America and Africa. Future research,
therefore, requires a highly disciplined effort whereby the current status of
the disease should be thoroughly analyzed and plant resistance responses
and pathogen virulence factors systematically investigated. An integrated
disease management program, built on enhanced plant resistance and
health, pathogen containment and improved field management strategies
should be advocated.

INTRODUCTION

The History and Global Distribution of the Disease

Diseases are a major constraint to banana production worldwide. A great number
of diseases affect banana (Jones, 2000), with one of the most important of these diseases
being Fusarium wilt of banana.
The first record of Fusarium wilt was in Australia in 1874 (Bancroft, 1876) and
the disease has since been found in all banana-growing regions of the world, except for
some of the countries bordering the Mediterranean Sea. Fusarium wilt has had a

particularly destructive history in the evolution of international banana trade. In the

period 1890-1960, some 40,000 hectares of the susceptible banana cultivar Gros Michel
(grown for export) were destroyed or abandoned in Central and South America and the
Caribbean because of race 1of Foc. In terms of crop destruction, Fusarium wilt then
ranked alongside the foremost devastating plant diseases such as wheat rust and potato
blight (Carefoot and Sprott, 1969).

Export industries were forced to replace the

susceptible Gros Michel variety with Cavendish cultivars, which continue to show
resistance to race 1 of Foc in these areas (Stover, 1990).
Cavendish cultivars remain the banana varieties of international trade. However,
these cultivars are not resistant to all strains of Foc. The subtropical race 4 strain of Foc
causes losses of Cavendish cultivars in the subtropical regions of the Canary Islands,
South Africa, Australia and Taiwan (Stover, 1990). More importantly, in the tropical
commercial and subsistence production regions of the Philippines, Indonesia, Taiwan,
Malaysia, and in the southern provinces of China, a new strain of Foc designated tropical
race 4 has caused widespread devastation (INIBAP, 2006). Alarmingly, the disease is
continuing to spread in these areas. Several incursions of this pathogen have also been
recorded in Australia (Northern Territory); however these outbreaks have all been
contained and have not reached the commercial growing regions situated on the eastcoast of the country (Walduck, 2002).
Tropical race 4 of Foc affects banana cultivars that comprise 80% of the worlds
banana production, including the important Cavendish and plantain subgroups (Ploetz,
2005). The tropical race 4 strain of Foc could cause significant damage to the major
world export production areas if introduced into Ecuador, Central America and
Colombia, which are based on Cavendish cultivars. As it stands, the tropical race 4 strain
poses a very real threat to the multi-billion dollar global banana trade, and the food
security of millions of subsistence farmers (Ploetz, 2005). Furthermore, the Cavendish
variety may risk the very same fate as Gros Michel, the cultivar it replaced nearly 50
years ago because of its susceptibility to race 1 of Foc.

Foc Recent Threat to Philippine Banana

Philippine Banana particularly the Cavendish type has gained popularity
worldwide making the country as the 3rd largest producer and exporter (in terms of
volume) in the world in 2012. Almost 50% of the total banana production is contributed
by Cavendish. It has been reported that Cavendish succumbed to Foc TR4 which has
already been reported in other Asian countries.

Here the most affected are the

independent growers (IG) since the multinationals can contain the disease. Now that IG
are the ones affected, the fast spread of the disease also scares the multinationals hence
everyone would like to put their efforts to address the disease. Region XI which grows
almost 60% of the total Cavendish started to be alarmed (Generalao, 2013).
Fusarium wilt, a soil-borne fungal disease in banana has been considered a threat
in the region, with Foc TR4 considered as the most devastating. It has already been
reported in China, Indonesia, Malaysia, and Taiwan. Recently, it was reported that more
than 5,000 ha Cavendish farms in Davao region has been infected with Fusarium wilt.

REVIEW OF LITERATURE

Disease Symptoms
Fusarium wilt of banana can be described as a classic vascular wilt disease
causing disruption of water translocation, systemic foliar symptoms and plant collapse
(Jeger et al., 1995). It invades the vascular tissue (xylem) through the roots causing
discoloration and wilting. Externally, the first obvious signs of disease in most varieties
are wilting and a light yellow coloring of the lower older leaves, most prominent around
the margins. They eventually turn a bright yellow color with dead leaf margins (Fig. 1).
As the disease advances, more of the leaves become yellow and die. A skirt of dead
leaves often surrounds the pseudo-stem. In the advanced stages of disease, affected plants
may have a spiky appearance due to prominent upright apical leaves in contrast to the
skirt of dead lower leaves (Fig. 2).
Internally, symptoms first become obvious in the xylem (water conducting)
vessels of the roots and the rhizome. These turn a reddish-brown to maroon colour as the

fungus grows through the tissues (Fig

(Fig. 3). Occasionally, the discoloration
ration first appears
yellow in plants showing early stages of infection
infection. When a cross-section
section is cut,
c the
discoloration
ration appears in a circular pattern around the centre of the rhizome where the
infection concentrates due to the arrangement of the vessels (Fig
(Fig. 4).
4 As symptoms
progress into the pseudo
pseudo-stem, continuous lines of discoloration
ration are evident
eviden when the
plant is cut longitudinally
longitudinally.. The infection may travel all the way up to the top of the
pseudo-stem.
stem. In severe cases it may even enter the leaf petioles and the peduncle (bunch
stalk) of bunched plants. However, infection has not been shown to progress into the fruit
(Stover, 1962a).
Banana suckers that are less than about four months old do not develop visible
symptoms of Fusarium wilt. The lack of visible symptoms on suckers has assisted in the
movement of the pathogen to new regions through the movement of these asymptomatic
suckers to new areas as planting material. The fruit of the banana plant does not show any
specific disease symptoms.

Fig. 1. Plant showing early symptoms of

Fusarium wilt
wilt.

Fig. 2. Plant showing advanced symptoms

of Fusarium wilt.
wilt

Fig. 3. Reddish-brown to maroon discoloration of vessels in the pseudo-stem.

Fig. 4. Reddish-brown to maroon discoloration of vessels in rhizome.

Disease Development and Epidemiology

The disease cycle

Spores of Foc in the soil germinate and grow towards the nearby roots of banana
plants in response to chemical compounds exuded from the roots (Fig. 5). Infection takes
place on the secondary and finer roots and proceeds into the larger, primary roots through
the xylem vessels before entering the rhizome. The primary roots and the rhizome do not
appear to be infected directly by the pathogen. The xylem network consists of a series of

individual vessels with perforated end walls through which the sap flows. Movement of
the spores with the sap flow is blocked temporarily when they become lodged at the end
walls. The spores then germinate and hyphae grow through the perforations into
adjoining vessels where further spores are produced. The plant is often able to prevent
preven
infection from successfully travelling to and entering the rhizome by the production of
gels and tyloses (a resistance mechanism) to seal off the infection. However, multiple
infections occur during the life of a plant and invariably one or more lead to its complete
invasion. The virulence of tropical race 4 on Cavendish bananas suggests that the
resistance mechanisms employed by the plant against this strain are not as effective as for
sub-tropical
tropical race 4. This strain generally only causes serious losses in a plantation
where the plants are under stress.

Fig. 5.. Disease initiation and life cycle of Foc in a banana plant.

Spread of the pathogen

Foc is most commonly spread by the movement of infected planting material,
rhizomes and suckers (and the attached soil), to new uninfected areas. This infected
planting material often does not exhibit symptoms of Fusarium wilt (is asymptomatic).
The pathogen can also be effectively spread by the movement of soil, running water, and
farm machinery and implements. Once a site is infected, the pathogen can persist in the
soil as chlamydospores for more than 30 years (Stover 1962a; Waite and Dunlap 1953).
Also, it is likely that Foc can survive non-pathogenically on alternative hosts, such as
weed species (Hennessey et al.2005).

The life cycle of Foc

The life cycle of Foc begins with the entry of the pathogen into the potential host
via the root tip. Substances produced by the host, in the region behind the zone of
elongation of the root meristem elicit a pathogen response which results in the
germination of chlamydospores. The germinated hyphae then infect the lateral roots and
progress to invade the xylem vessels. Further spread of most initial infections is usually
stopped in the xylem by the vascular occluding responses of the host, which include the
formation of gels, tyloses and the collapse of vessels. In susceptible cultivars, some of
these infections become established in the xylem and advance ahead of these defense
mechanisms. Microconidia are formed in the xylem vessels and are spread through the
vascular system of the plant, streaming to new sites where they germinate. Hyphae are
then produced, and this begins the invasion at this new site, thus repeating the cycle. In
resistant varieties, the initial pathogen-induced occlusion reaction is further enhanced by
the production of phenolic compounds, which lignifies these obstructions and limits the
pathogen to the infected vessels; no further colonization of the xylem vessels occurs
(Beckman 1969; Beckman 1987; Beckman 1990; Beckman and Keller 1977; Beckman
and Talboys 1981; VanderMolen et al. 1977).

Fig. 6. The life cycle of Foc in the soil and in its host.

Disease Control

Initially, control of Fusarium wilt was aimed at reducing

educing the pathogen population
in infested fields. However, fungicides, fumigants, flood fallowing, crop rotation (with
corn) and organic amendments have rarely provided long
long-term
term control in any production
pro
area.
Quarantine
ine and exclusion procedures are effective in controlling the spread of the
disease by restricting the movement of corms, suckers and soil that could carry Foc from
infested to non-infested areas. In the whole of Davao region,, legislation provided
provi
by the
President Decree of 1978
1978, which operates through the DA-RFU XII Quarantine Service
andd the Bureau of Plant Industry
Industry,, prohibits the transfer of planting material from certain
localities where the disease iiss prevalent or from any plantation not approved as a source
of planting material. Planting material is only approved if the source plantation has no
previous record of the occurrence of the disease, and is apparently free from Fusarium
wilt when subjected to row
row-by-row
row inspection by trained personnel preceding the

removal of the material. However, due to the presence of suppressive soils in which
microbial populations suppress the pathogen population, and since infected rhizomes or
suckers may not exhibit external symptoms, the pathogen may still be moved in approved
planting material. In recent years, the use of certified disease-free tissue-cultured
plantlets, from an approved Quality Banana Approved Nursery facility, has become a
cost effective way of maintaining disease-free production. Tissue-culture plants also have
the added benefits of providing uniform growth and harvest times.

It is now generally accepted that the most effective means of controlling Fusarium
wilt is by host resistance. Since banana is a clonally propagated, perennial crop,
resistance must be enduring. Natural sources of resistance exist in wild species and
cultivars of banana, and also in synthetic diploids developed by breeding programs.
Biotechnology, mutation breeding and somaclonal variations are also being used to
produce resistant genotypes. These programs are likely to provide replacement clones in
the future. Since the Philippines does not have a banana breeding program, close
collaboration with international breeding and plant improvement programs is essential for
obtaining replacement clones in the future.
Disease management

Management of this disease in Davao region is based around three key strategies:
the early detection and containment of outbreaks of Fusarium wilt and application of
protocols to prevent introduction of Foc into disease-free production districts; (2)
adoption of disease management strategies to enable farms with Fusarium wilt to
maintain production, and (3) research to evaluate and develop long-term disease
management strategies for the Mindanao banana industry.
Local quarantine measures are in place in banana production areas to restrict the
movement of banana planting material (particularly suckers and rhizome pieces). Other
banana growers require permits to move planting material to initiate new plantations and,
if vegetative material is used (e.g. suckers or rhizome pieces) this must be from a
certified source of clean planting material.

Result of the survey conducted by the PCAARRDs Socio-Economics Research

Division (SERD) for the project titled An Assessment of the Socio-economic Impact of
Fusarium Wilt Disease of Cavendish Banana (Aquino, 2013) at the farm level have
enumerated different methods used by small scale farmers to destroy Fusarium infected
plants. These are:
Single Method

Burning with rice hulls

Injecting chemicals into the pseudostem

Injecting biocontrol agents into the pseudostem

Putting chemical-impregnated sticks into the corm

Pouring chemicals on the candle/cigar leaf

Spraying chemicals on the whole plant

Removing leaves and chopping the plant
Combination of Methods

Burning with rice hulls
 and injecting chemicals into the pseudostem
 and putting chemical-impregnated sticks into the corm
 and injecting chemicals into the pseudostem and putting chemicalimpregnated sticks into the corm

Injecting chemicals into the pseudostem and pouring chemicals on the
candle/cigar leaf

Putting chemical-impregnated sticks into the corm or putting pure salt into the
pseudostem

CRITICAL REVIEW

The following are the summarized methods and strategies of control that
individual growers, plantation groups, joint effects of government/private/nongovernmental institutions employed or are currently employing against Fusarium wilt
disease in their banana plantation and their corresponding epidemiologic effects (Table
1):
Table 1. Methods/strategies of disease control in Fusarium wilt of banana and their
epidemiological effects
METHODS/STRATEGIES OF CONTROL
MAJOR EFFECT ON
A. Avoidance of the Pathogen
1. Use of disease-free tissue cultured planting materials
Xo
2. Modification of cultural practices
r
B. Exclusion of the Pathogen
1. Treatment of planting materials
Xo
2. Inspection and certification
Xo
3. Exclusion or restriction by plant quarantine
Xo
r
C. Eradication of the Pathogen
1. Biological control of plant pathogens
Xo
r
2. Crop rotation
Xo
3. Removal or destruction of diseased plants
Xo
r
4. Dipping to fungicide of the planting stocks
Xo
5. Drenching with fungicide on the planting holes as basal
Xo
treatment
D. Protection of the Plant
1. Modification of the environment
r
2. Modification of nutrition (fertilization)
r
E. Development of Resistant Host
1. Selection and breeding for resistance
a. Vertical resistance
Xo
b. Horizontal resistance
r
2. Somaclonal variation (multilines)
r
3. Resistance through nutrition
r

Critical Analysis in Relation to Epidemiology

The problem is far from over!
The appearance of a virulent strain of Foc in Davao regions Cavendish banana
and in other Asian countries renewed the nightmare brought about by the Panama
Disease epidemic during the Gros Michel era. Foc TR4 infects not only Cavendish type
but other cultivars including the local variety Lakatan which is even more susceptible
than Cavendish. The disease is really a threat to our export banana and a threat to the
small-growers in their livelihood and food.
Although Fusarium wilt and its diversed pathogen forms have been in this region
before the 90s, the disease destructive potential was not realized until the demise of a lot
of commercial Cavendish plantations.
Why is it so dreadful? It is because the pathogen kills the plant completely! It can
survive and remains infective for decades. Until now, there was no effective fungicide to
eradicate the pathogen. It affects many important cultivars and potentially cause severe
epidemics especially the perennial mono-cropping system employed in banana
plantations in the region.
Why the fast spread of the disease? The following are identified factors in the
spread of the disease: infected rhizome/suckers, infected sheaths used to cushion fruit
crates, infested soil, contaminated irrigation or running water and infested farm
implements or machinery.
There are still banana growers that are traditionally using rhizome (bull heads)
and suckers as planting materials for banana. As previously discussed, young suckers
though infected or coming from an infected mother would show asymptomatic
expression of the disease. The farmer might have chosen an infected one and successfully
planted and distributed Fusarium in his farm. This is the same scenario with the use of
infected rhizome as planting stocks.
Local variety Lakatan is marketed from Mindanao to Visayas and to Luzon local
markets. The process of packing banana for transport might have contributed to the long
spread of the disease. Bananas are put in wooden crates but cushioned with banana bract
sheaths. The problem arises when they accidentally used infected sheaths to cushion the
fruit crates. Anywhere in the destination of the product, they will threw away the

infected sheaths and there goes the pathogen propagule will be able to land in a new area
and would just wait for the right time (host, environment) to cause new destruction.
There are a lot of workers in a banana plantation and even visitors. There in and
out of the area will contribute to the pathogens spread. If they will be able to step in a
Foc-infested soil then they are carrying with them the pathogens propagule. As to where
they are tracking in the area, there they already have sown the pathogen. This is really
very critical in the workers in the farm. I suggest there should only be one entry and exit
point. They should step (by wearing rubber boots) in a disinfectant before entering the
banana area and whenever they go out, step again to a disinfectant before going to
another parcel of the banana plantation. Big plantations are doing this. But independent
banana growers are mostly not.
During drought season or when there is scarcity of water that runs through the
drip irrigations in the plantation floors, there is an augmentation of boom sprayed
irrigation. The fear of this is that when the source of water is coming from a nearby
plantation river which in some way have catched some propagules of the pathogen.
There will be massive distribution of the propagules then when sprayed in the area.
The use of farm implements/machinery should be taken with utmost care because
they might have carried with them pathogen propagules and brought to the non-infested
area or vice-versa. So, before using in the new area, tires and its parts should be properly
cleaned and disinfested. That is IN and OUT protocol.
Well, the unavoidable spread of the disease was during the great flood of the area
mostly in ComVal province after the hit of typhoon Pablo. The previously non-infested
area the plants became infected after recovery of the typhoon and later on abandoned
because of the total damage brought by Fusarium wilt. Meaning, water is a successful
mean of disease spread.
The pathogen forms chlamydospores as a mean of survival. This activates with
the presence of the hosts exudates or from any alternate hosts. Therefore, alternate hosts
should be removed as they can serve as reservoir of inoculums.
Though banana growers and experts/scientists have not yet perfected the weapons
for this dreadful disease, proper identification (diagnosis) is essential for everyone
concerned in the banana industry and be reported at once for any sightings or early

detection to right personnel. Alternative procedure for eradication should be strictly

implemented and followed at once. Inoculum source and dead infected mats should be
totally removed in the area through proper burning. And lastly, area isolation and use of
disinfectant and strong quarantine implementation should never be handled as a joke.
The efforts of the Department of Agriculture and the Department of Science and
Technology as to mapping the disease spread are very relevant this time. We mostly
knew that the disease all started in Davao region and even identified through PCR the
identity that it was the dreadful TR4. The survey and mapping did not only looked into
Cavendish but including our Lakatan, Latundan and Cardaba. There are still areas
where there are no sightings yet of the disease, this is where our quarantine protocols be
strictly imposed and provincial legislations be made.

On-farm disease management: A critical review

Provided the strain of Foc involved in an outbreak of Fusarium wilt does not pose
a quarantine risk, or the property concerned is not beyond a defined local quarantine
zone, banana production is allowed to continue. Banana growers are advised to follow
simple yet practical measures to minimize the spread of the disease and to extend the
productive life of the plantation.
It is essential to prevent the spread of the pathogen to other farms (e.g. no
movement of planting material or infested soil and, if possible, containing or diverting
associated drainage water), and to minimize the spread of Foc within the affected farm.
The common procedure in by farmers and growers for isolated cases of Fusarium wilt in
an otherwise healthy block is to immediately inject the affected plants with herbicide
(e.g. Roundup/glyphosate), and let them die in situ without further disturbance of the
plants. Often a single or double ring of healthy plants around the affected ones may also
be injected or otherwise treated with herbicide and allowed to die in place. This has the
two-fold effect of preventing mat-to-mat spread of the pathogen through living banana
roots, and also reducing the population of the fungus at that site by killing its preferred
host plant. It is important that the affected plants and the soil around them be disturbed as
little as possible in this process as any disturbance and movement of soil will increase the

chances of moving the pathogen. Fencing-off the area also signals to workers or visitors
to keep out, reducing the risk of movement of contaminated soil on boots, machinery or
in suckers.
Banana growers must not be tempted to use planting material from known
infected plantations, even if some rhizomes appear clean. If Foc has been identified at a
plantation, any movement of soil or plants from the affected block must be considered as
an unacceptable risk.
If disease-free areas are available for banana production it is vitally important not
to contaminate them through the lack of basic farm hygiene. The key to keeping diseasefree areas free from Foc is to minimize, if not remove, all opportunities for infested soil,
plant material or irrigation water (that may be carrying spores of the fungus attached to
soil particles) from entering the area. Risk minimization can be achieved in several ways:

Using only certified disease-free vegetative planting material or tissue-cultured

plants to initiate new plantations;

Cleaning equipment, footwear and vehicles to avoid moving infested soil or plant
matter from an affected block to disease-free blocks;

Where practicable, isolating irrigation water that drains from diseased parts of a
plantation. Such drainage water may be carrying spores of Foc and should not be
used to irrigate new or disease free areas if possible. At this time no commercial
treatments are known for controlling Foc in irrigation water or water storages.

Educating farm workers about how the fungus is spread and providing posters or
other visual material to enable symptoms of Fusarium wilt to be recognized to aid
in early detection of new outbreaks.

Use of disinfectant applied to vehicles (particularly tyres and inside wheel arches)
as a foaming spray, which is then left for about 10 minutes to allow it to soak into
any soil that may still be attached before being rinsed off. It is also used in
footbaths for cleaning boots when moving between infected and disease-free areas
within and between properties.

Recommendations for Management and Future Research

The following are my recommendations for continued searched for best options
against Foc TR4 in our banana plantations:

Updated mapping on the distribution of Foc TR4 and distribution of results to

concerned agencies and discussing its relevance;

Continued developing/validating disease management measure: varietal resistance

trial, IPM approaches, biological control, annual cropping, soil health
management, eradication, fungicide treatment, and quarantine

Understanding mechanisms of soil suppression studies

Microbial population interaction studies and development and identification of

indicators

Reproduction of diagnostic tools

Establishing collaborative studies

Remote sensing and image analysis

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CAREFOOT, G.L. and E.R. SPROTT. (1969). Famine on the wind. Angus and
Robertson: London.
GENERALAO, L.C. (2013). Panama Program Report. SMARRDEC-DOST-PCAARRD.
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A.E. BROWNE. (1995). Banana diseases. In: Bananas and Plantains. (ed. S.
Gowen). Chapman and Hall: London. pp. 337-343.

JONES, D.R. (2000). Diseases of banana, abaca and enset. CAB International:
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PLOETZ, R.C. (2005). Panama disease, an old enemy rears its ugly head: parts 1 and 2.
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STOVER, R.H. (1962). Fusarium wilt (Panama disease) of bananas and other Musa
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WALDUCK, G. (2002). New varieties and quarantine help solve Panama riddle.
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