Ralstonia solanacearum: The Bacterial Wilt Causal Agent
Monther Mohumad Tahat
Ralstonia solanacearum (race 3 biovar 2) is a bacterial wilt causal agent of many plant species. Infects (potatoes Solanum tuberosum, eggplant Solanum melongena, peppers Capsicum annuum, tomatoes Lycopersicon esculentum, geraniums, Geranium carolinianum, ginger Zingiber officinale and a few weed species including bittersweet Celastrus orbiculatus, nightshade Solanum karsense and stinging nettle Urtica dioica. Ralstonia solanacearum can be infectious in the soil for years in the presence of a host. Race 3 biovar 2 is most commonly transmitted by contaminated soil, equipment, water and insect, or by transplantation of infected seeds or seedlings. Management requires use of resistance cultivars, clean and certified seed, good cultural practices, some chemicals fumigation, antagonistic microbes as a biological control like (Mycorrhizal fungi, Streptomyces sp. and Tricoderma sp.) transgenic resistant plant, cropping systems, soil amendments, integrated control, genetically engineered antagonistic and virulent mutants of R. solanacearum.
August 16, 2010; Accepted: September 09, 2010;
Published: August 29, 2012
Ralstonia solanaceraum is an important pathogen of many crops. Formerly
called Pseudomonas solanacearum. Historically, strains of R. solanacearum
were classified into five races based loosely on host range and into five
biovars based on differential ability to produce acid from a panel of carbohydrates
The five races of R. solanacearum have different host ranges and geographic
distributions. Race 1 is a poorly-defined group with a very wide host range
and is endemic to the southern United States as well as Africa, Asia and South
America. Race 2 principally attacks bananas and is found mainly in Central America
and Southeast Asia. Race 3 is distributed worldwide and has primarily been associated
with potato. Race 4 affects ginger in much of Asia and Hawaii and race 5 affects
mulberries in China (Kelman, 1997; Denny,
2006). The origin of R. solanacearum is not clear, but Hayward
(1991) suggests it predates the geological separation of the continents
as the bacterium has been found in virgin jungle in South America and Indonesia.
However, race 3 biovar 2 strains are believed to originate in the Andean highlands
and this near-clonal subgroup is widely distributed in tropical ones throughout
the world and some temperate regions such as Europe and northern Asia.
Cultural practices, crop rotation and host resistance may provide limited control
of R. solanacearum (Kucharek, 1998; Pradhanang
et al., 2003). Several plant essential oils and their components
showed that some essential oils have significant efficacy against R. solanacearum
in vitro and under glasshouse (Momol et al.,
1999) and against several soilborne fungi of tomato (Momol
et al., 2000). Disease incidences of 15 to 55% have been reported
in fresh market tomato in Taiwan, causing losses exceeding 12 million U.S. dollars
annually (Hartman et al., 1991). In Hawaii, ginger
production was reported to have suffered losses of over 50% during 1998 and
1999 (Yu et al., 2003). Rapid early detection
of bacterial wilt is not only in tubers or plant debris but also in soil or
soil-related habitats is essential for disease management in the field to prevent
losses and further pathogen spread (Janse et al., 1998).
Disease causal agent: Ralstonia solanacearum is a highly heterogeneous
bacterial pathogen that causes severe wilting of many important plants (Smith
et al., 1995). The disease is also called Southern bacterial blight,
R. solanaceous wilt, Southern bacterial wilt and many other common names
in countries where it occurs (Buddenhagen and Kelman, 1964).
It is an aerobic obligate organism; strains of the pathogen have minimum, optimum
and maximum temperature of 10, 35 and 41°C respectively (Kelman,
1953). Ralstonia solanacearum is a gram-negative, non-spore forming
rod, about 0.5-0.7 μm x1.5-2.0 μm with a single polar flagellum (Sneath
et al., 1986). The bacterium is aerobic and its colonies on solid
media are small, irregularly round, white in reflected light and tan in transmitted
light (Hayward, 1991). Four races and five biovars were
classified based on oxidation of sugar and sugar alcohol (Strider
et al., 1981). Different races of R. solanacearum have different
host range. For example, race 1 can affect many flowering crops, race 2 can
affect banana plant and race 3 can affect potato, tomato and some other Solanaceae,
race 4 can affect ginger and some wide plant (Denny and Hayward,
2001). R. solanacearum have variation in metabolic activity into
5 or 6 different biovars. It is a complex species with considerable diversity,
although workers have variously divided the species into "group, strains, pathovars,
biotype and race (Strider et al., 1981).
Ralstonia solanacearum classification:
Other names for R. solanacearum
Bacillus solanacearum Smith 1896
Burkholderia solanacearum (Smith, 1896) Yabuuchi and
Pseudomonas solanacearum (Smith, 1896) Smith (1914)
Ralstonia solanacearum (Smith, 1896) Yabuuchi et
al. (1995) (http://www.ncipmc.org/ralstonia)
Recently amore phylogenetically meaningful system has classified R. solanacearum
into four major genetic groups called phylotypes that reflect the geographical
origin and ancestral relationships between strains (Fegan and
Prior, 2005). Ralstonia solanacearum race 3 biovar 2 is a soil-borne
pathogen that persists in wet soils, depth soil layers (>75 cm) and reservoir
plants. The tetrazolium medium (TZC), described by Kelman
and Person (1954) is the best for culturing R. solanacearum. The
organism produces two easily distinguishable type of colonies; one is small,
flat, red and butyrous (Chen and Echandi, 1982) (Fig.
1a) while the other colony is large, elevated, mostly white with light pink
centers and full of fluid using Casamino Acid Peptone Glucose (CPG) (Fig.
1b) (Cuppels et al., 1978).
The most important race of R. solanacearum is race 3 biovar 2 (Table 1), which has valuable agricultural hosts.
||The most important races, host range, geographical distribution
and biovars of Ralstonia solanacearum
|Adapted from (Denny and Hayward, 2001)
||(a) Casamino Acid Peptone Glucose (CPG), (b) Tetrazolium Media
Occurrence and host range: Bacterial wilt caused by R. solanacearum
has been described on a wide range of hosts in many tropical and subtropical
regions (Agrios, 2005). In the absence of susceptible crops,
alternative weed hosts and non-host plants play important roles for the survival
of R. solanacearum strains (Granada and Sequeria,
1983). Ralstonia solanacearum causes lethal wilting disease in more
than 200 plant species (Denny, 2000), while more than
450 plant species were listed as host plants for R. solanacearum including
many important and economic crops (Hayward, 1991). Host
range of R. solanacearum includes several hundred species representing
44 families of plants and many newly recognized hosts. R. solanacearum
biovar 3 has been described on some woody perennial hosts including cashew (Anacardium
occideutale) and custard apple (Annona spp.) (Mayers
and Hutton, 1987).
The reason for these highly different and heterogeneous bacterial pathogen
is not obvious; nevertheless it is assumed that specific pathogenic strains
for certain hosts may have evolved only in certain parts of the world and are
not found elsewhere or these hosts may only be susceptible where a number of
environmental factors such as temperature, rainfall, soil type, inoculums and
other soil biological factors are conductive to disease expression coincide
(Hayward, 1991). There is an extensive literature on
the disease, particularly in terms of host range, geographic distribution, occurrence
of various biovars, strains and the ability to survive in soil and in association
with plants debris and weed hosts (Persley, 1992).
Disease distribution: The first record of bacterial wilt Ralstonia
solanacearum (Smith, 1896) Pseudomonas solanacearum E. F. Smith)
in the world was reported by Burrill in 1890 in Japan. That was found to be
on tuber rot of potato (Gota, 1992). Across the world
there are differences between R. solanacearum races and biovars depending
on the geographical distribution (Hayward, 1991; Buddenhagen
and Kelman, 1964). Biovar 1 is predominant in USA and biovar 3 in Asia,
whereas biovar 2 and 5 occur in Australia (Pitkethley, 1981)
and China (He et al., 1983). It is also found
that biovar 4 occurs in India and Indonesia. In Africa the bacterial wilt disease
was recorded in Egypt, Libya, South Africa, Zambia and Burundi (OEPP/EPPO,
1999). In the Philippines all of biovars 1-4 have been found and here as
elsewhere in Asia biovar 3 is a predominant biovar in lowland regions. Strawberry
is a host in Japan and Taiwan but not in the southeastern USA (Hayward,
1991). Around 43 plant species were found as a plant host of the bacterial
wilt disease in Malaysia especially biovar 3 race 2 (Abdullah,
1982). Bacterial wilt disease was also observed in Cameron Highland/Malaysia
at about 1545 M above the sea level. In addition, the disease has been observed
and the pathogen was isolated from the infected crop plants and weeds at the
farm of Universiti Putra Malaysia, Selangor (Abdullah, 1988,
Ralstonia solanacearum is listed as a quarantine organism in the European
Union (EU) (Anonymous, 1995). In EPPO region the bacterial
wilt disease was found in Belgium, Spain, Netherland, Germany, United Kingdom
and Hungary (OEPP/EPPO, 1999). Bacterial wilt causal
agent can spread between countries by water, latently infected planting materials
and soil. One plant species that is seriously affected by bacterial wilt is
tomato and the efforts to grow tomato widely in the tropics have generally been
hampered by this disease (Hayward, 2000). Two new hosts
for Ralstonia solanacearum: davana (Artemisia pallens) and coleus
(Coleus forskohlii) were recorded by Chandrashekara
and Prasannakumar (2010), both are important crops in medicinal and aromatic
industries in India. Coleus and davana plants showing typical wilt symptoms
Disease symptoms: Ralstonia solanacearum is characterized by
sudden wilting of foliage and the young plant is affected more. The symptoms
occur as discoloration of the vascular system from pale yellow to dark (Gota,
1992). The plant infected with R. solanacearum may express all or
none of these symptoms, even under typical environmental conditions that are
ideal for the pathogen and typically this is a commonly observed condition known
as latency. The pathogen enters roots through wounds caused by transplanting,
cultivation, nematode, insects and through natural wounds. Then it starts to
multiply rapidly in the vascular system, finally the xylem elements are filled
with bacterial cell and slime (Kelman and Sequeira, 1965).
Molecular aptamers are single-stranded oligonucleotides that can specifically
bind with high affinity to a variety of molecules ranging from macromolecules
to small compounds. Aptamers potentially could be produced and used in the future
as reproducible, fast and highly-specific diagnostic tools for R3bv2. (Champoiseau
et al., 2009a).
Successful and efficient colonization requires production of molecular mass
Extracellular Polysaccharide (EPS) in a high amount and multiple extracellular
protein (EXPs) (Denny, 2000). The incidence the disease
infection may range from a few scattered plants or loci of infection in fields
where low or erratic natural infestations occur to the rapid death of the plants
(Kelman and Sequeira, 1965). The bacterium rapidly spreads
upward in the vascular system from secondary roots to larger roots and then
to the stem. After that, the plant starts to suffer from wilting irreversibly.
Older plant leaves first show wilting before the youngest leaves or one sided
wilting and stunting and finally the plant wilts permanently and dies (Agrios,
Massive invasion of the cortex might have resulted in the appearance of water-soaked
lesions on the external surface of the stem; if an infected stem is cut crosswise
tiny drops of dirty white or yellowish viscous ooze exude from several vascular
bundles (Champoiseau et al., 2009b). The pathogen
ingresses a plant through the roots, penetrate the xylem, systemically colonize
the stem and causes wilt symptoms (Kelman, 1953). This
bacterium causes wilt by infecting plants through roots and colonizing stem
vascular tissue. Although diseased plants can be found scattered in the field.
Under natural conditions, the initial symptom in mature plants is wilting of
upper leaves during hot days followed by recovery throughout the evening and
early hours of the morning. The wilted leaves maintain their green color as
disease progresses. Under hot humid conditions disease complete wilting occurs
and the plant will die. The brown discoloration shown in the lower stem vascular
tissues (Kucharek, 1998).
It is difficult to control bacterial wilt disease in the soil (Jones,
1997). Various control strategies were developed to control and suppress
this disease including host-plant resistance and biological control (Dalal
et al., 1999). Potential biological agents were used to control bacterial
wilt of tomato (Lycopersicon esculentum) include vesicular-arbuscular
mycorrhizae (VAM) (Halos and Zorrilla, 1979) and some
naturally occurring antagonistic rhizobacteria such as Bacillus sp. (Silveira
et al., 1995), Pseudomonas sp. (Guo et
The most important and applicable methods are:
Resistance cultivars: The best strategy to control bacterial wilt caused
by R. solanacearum is breeding for resistance cultivar (Persley,
1992). A virulent mutant of R. solanacearum has been used as a bio-control
agent for the virulent pathogen (Trigalet and Trigalet-Demery,
1990). Abdalla and Abdulla (1998) found that the
degree of susceptibility to bacterial wilt is significantly different among
six tomato cultivars which were tested and this indicated that the additive
genes were more important than the non-additive genes. Thus, in breeding programs,
selection for disease resistant plants (genotype) after each generation is recommended.
In the United States, Southern tomato transplant growers sometimes prevent the
disease by avoiding infected fields (Hayward, 1964).
The infection by R. solanacearum can be controlled by the use of hot
pepper accessions (Capsicum annuum L.), sweet pepper (Capsicum annuum
L.) in Japan. Bacterial multiplication in stems of resistant tomato plants
was suppressed owing to the limitation of pathogen movement from the protoxylem
or the primary xylem to other xylem tissues. The limitation was most conspicuous
in Hawaii 7996. Grafting experiments indicated that the percentage of wilting
of Ponderosa scions was less on Hawaii 7996 rootstocks than that on the most
resistant rootstock (LS-89) used in Japan. Hawaii 7996 could be an alternative
genetic source for breeding for resistance to bacterial wilt (Nakaho
et al., 2004).
Cultural practices: Crop rotation, intercropping or incorporation, green
manure and planting a susceptible crop such as mungbean before the cultivation
have been practiced (Hartman et al., 1993). Crop
rotation with a non-susceptible crop provides some control, but this measure
is difficult to use because of the wide host range of the pathogen (Kelman,
1953). Islam and Toyota (2004) demonstrated that
the bacterial wilt of tomato was suppressed in the poultry and farmyard manure
(FYM) added soils and higher microbial activity was likely responsible. In Nepal,
the importance of crop rotation and resistance cultivars were studied by Adhikari
and Basyat (1998). They reported that the appearance of bacterial wilt symptoms
were delayed by 1-3 weeks and the wilt severity was reduced by 20-26% when a
susceptible tomato was grown after corn (Zea mays), ladys finger
(Abelomoschus esculentum), cowpea (Vigna unguiculata), or resistant
tomato. Finally it is found that crop rotation with corn, ladys finger,
cowpea and resistance cultivars appeared to be useful management strategies
to control tomato bacterial wilt. The infection of potato plant by Ralstonia
solanacearum can be significantly reduced by using non-susceptible crops
and crop rotation for 5-7 years (Smith et al., 1995).
Rice husks, corn stalke, cow manure, oyster shell powder and mineral ash have
been used by improved mixture called Sun and Hung (SH) (Sun
and Hung, 1985). The application of the organic amendment (Wydra
et al., 2005)and compost released biologically active substances
from crop residues and soil microorganisms such as allelochemicals (Bailey
and Lazarovits, 2003). The Sun hemp mixture reduced the bacterial population
after 4 weeks of incubation at 2, 6 and 10% rates of incorporation of inoculated
plants wilted and died after the second week (Hartman et
al., 1993). The addition of household compost resulted in enhanced decline
rate of Ralstonia solanacearum population (Schonfeld
et al., 2003). Corn stalk, rice straw and tree bark were used for
the suppression of tomato bacterial wilt caused by R. solanacearum in
Malaysia and the results showed that all compost used significantly reduced
the disease severity index but the disease incidence was the lowest in bark
compost (Masyitah, 2004). Abdullah
et al. (1983) found that soil type and moisture levels individually
as well as in combination had a significant effect on the severity of the bacterial
wilt of groundnut. Studying the distribution of disease in Malaysia related
to the soil types illustrated that the tobacco plants were infected more when
grown on organic soils and light, heavy and intermediate types of mineral soils.
Also disease incidence in organic soils (more than 65% organic matter) is normally
low (Abdullah, 1988). Cow manure fertilization suppressed
brown rot caused by R. solanacearum in most soils with a clear shift
in rhizospere bacterial community. Stenotrophomonas maltophilia, isolated
from the rhizosphere of eggplant in the Egyptian Delta, was antagonistic to
R. solanacearum in vitro; its antagonistic activity was not Fe-siderophore
dependent. A selected S. maltophilia strain survived longer and reduced
R. solanacearum survival more in Egyptian than in Dutch clay soils and
suppressed potato brown rot in Egyptian soil (by at least 36%) but not in Dutch
soil (Messiha, 2006). The populations of native Ralstonia
spp. were reduced from 2.4-7x108 colony forming units (CFU) g-1
to 0-115 CFU g-1. Heat treatment reduced bacterial wilt incidence
by 50-75% (Kongkiattikajorn et al., 2006).
Chemical methods: Disease control using chemicals has been difficult
because of the localization of the pathogen inside the xylem and its ability
for survival at depth in the soil. In addition, this kind of control is not
economically feasible in the field. Some scientists reported that there are
no bactericides available for chemical control of the bacterial wilt disease
(Hartman et al., 1993), while others, reported
that it is difficult to control bacterial with chemicals (Grimault
et al., 1992). The bactericides Terlai have been tested in Taiwan
under both green house and field conditions (Hartman et
al., 1993) and it was found that chemical control through soil fumigation
and antibiotics (Penicillin, Ampicillin, Tetracycline and Streptomycin) has
shown little suppression of Ralstonia solanacearum (Murakoshi
and Takahashi, 1984). Some fumigation bactericides increased the extent
of wilt caused by Ralostonia solanacearum such as chloroform (Shiomi
et al., 1999). Antagonistic, avirulent bacteriocin-producing Ralstonia
solanacearum strains were investigated for potential biological control
of tomato (Lycopersicon esculentum) bacterial wilt in Brazil, it was
demonstrated that none of the bacteriocin-producing strains was inhibited by
its own bacteriocin (Araujo et al., 2004).
During 1992-1995 crop season, the field trials on the management of bacterial
wilt Ralostonia solanacearum of tomato revealed that Streptomycin proved
to be most effective as it gave highest disease control (79.5%) and lead to
maximum yield production (274.60 ha-1) compared to asafetida (Ferula
foetida) (70.30%) and turmeric powder (Curcuma longa Linn.) (69%)
(Mazumder, 1998). Many plant species produce volatile
essential oil compounds. These oils are considered to play a role in host defense
mechanisms against plant pathogens (Mihaliak et al.,
1991). Essential oils and their components, usually from medicinal plants,
have been recognized as having fungicidal effects (Wilson
et al., 1997), but their efficacy as a biofumigant on R. solanacearum
has not been studied prior to 1999.
Different sugars and amino acids were added to a conducive soil to study their
effects on bacterial wilt of tomato caused by Ralstonia solanacearum
YU1Rif43. Most of the seeds failed to germinate in soils to which serine, glycine
and alanine were applied at the same rate. At 2.5 mg g-1, the inhibitory
effect on tomato germination disappeared except for methionine. The compounds
that showed the most suppressive effect were glucose, proline, glutamine, serine,
arginine and lysine. The pathogen utilized glucose, proline and glutamine, but
not serine, arginine and lysine (Posas et al., 2007).
Biological control: Biological control can be defined as the direct
and accurate management of common components of ecosystem to protect plants
against pathogens. It is acceptable as a key practice in sustainable agriculture
(Azcon-Augiler and Barea, 1996). Biological control
preserves environmental quality by reducing the dependency on chemical input
and maintaining sustainable management practices (Barea and
Jeffries, 1995). Wall and Sanchez (1992) reported
that when bacteriophages, which are capable of attacking the bacterial wilt
pathogen, used as a bio-control agent showed that bacteriophages play an important
role in suppressing the population dynamics of R. solanacearum. Plant
growth promoting bacteria (PGPR) strains were reported to be a promising bio-control
agent to control R. solanacearum. It was found that they were able to
reduce the disease in different levels and increased the yield of tomato plant
(Guo et al., 2004; Kenichi,
118 strains of rhizobacteria, were screened against an Ethiopian R. solanacearum
strain. On the basis of in vitro screening, six strains (RP87, B2G, APF1,
APF2, APF3 and APF4) shown good inhibitory effect were selected for in planta
testing in a greenhouse. The study showed that APF1 and B2G strains reduced
significantly disease incidence and increased weight of tomato plants. Area
under Disease Progress Curves (AUDPC) was reduced by 60 and 56% in plants inoculated
with APF1 and B2G strains, respectively. APF1 was found to be the most beneficial
strain in disease suppression and also growth promotion resulting in 63% dry
weight increase compared to untreated control (Lemessa and
The effect of Biological Soil Disinfestation (BSD) was tested in glass vessels,
microplots and in an accidentally infested commercial field. BSD is based on
production of toxic organic acids through anaerobic digestion of fresh organic
matter. BSD was accomplished by incorporating grass or potato haulms in soil
and covering the soil with airtight plastic survival of R. solanacearum in
soil and potato tubers was significantly reduced in the BSD treatment (>93%)
and not in separate grass amendment -or plastic cover treatments (Messiha,
2006). There have been a few studies of the potential role of AMF for the
protection of plants from bacterial wilt (Hayward, 1991).
Glomus mosseae was able to suppress the infection of tomato plant by
R. solanacearum in the glasshouse conditions (Tahat
et al., 2010). Zhu and Yao (2004), found
that the Localized and systemic increase of phenol in tomato roots induced by
Glomus versiforme inhibits Ralstonia solanacearum.
Bacterial wilt, caused by Ralstonia solanacearum, is responsible for severe losses to many important crops, mainly Solanaceous plants and bananas, This microorganism is the causal agent of bacterial wilt, moko disease in banana southern wilt of geranium and potato brown rot. Develop disease management to control bacterial wilt on tomato, potato and geraniums including: 1-Screen additional chemical and biological control products. 2-Exclude the pathogen from tomato transplants, potato seeds and geranium cuttings and develop vegetative plant material certification schemes. 3-Study the effects of crop rotation, cover crops and mulches on pathogen dynamics and disease incidence 4- Develop additional DNA and immunological detection tools that can be used reliably to distinguish subgroups of R. solanacearum, especially Race 3 biovare 2, from other endemic strains of Ralstonia. 5-Educate county extension, growers and crop advisors in sampling, monitoring and management of related diseases.
The authors would like to thank the University Putra Malaysia for supporting this work.
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