Citrus canker
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Citrus canker is a disease affecting ''
Citrus ''Citrus'' is a genus of flowering trees and shrubs in the rue family, Rutaceae. Plants in the genus produce citrus fruits, including important crops such as oranges, lemons, grapefruits, pomelos, and limes. The genus ''Citrus'' is native to ...
'' species caused by the bacterium '' Xanthomonas'' (''X.axonopodis; X. campestris''). Infection causes lesions on the leaves, stems, and
fruit In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering. Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in partic ...
of citrus trees, including lime, oranges, and grapefruit. While not harmful to humans, canker significantly affects the vitality of citrus trees, causing leaves and fruit to drop prematurely; a fruit infected with canker is safe to eat, but too unsightly to be sold. The disease, which is believed to have originated in
Southeast Asia Southeast Asia, also spelled South East Asia and South-East Asia, and also known as Southeastern Asia, South-eastern Asia or SEA, is the geographical south-eastern region of Asia, consisting of the regions that are situated south of mainland ...
, is extremely persistent when it becomes established in an area. Citrus groves have been destroyed in attempts to eradicate the disease. Countries like Brazil and the United States also suffer from canker outbreaks.


Biology

''Xanthomonas axonopodis'' is a rod-shaped
Gram-negative Gram-negative bacteria are bacteria that do not retain the crystal violet stain used in the Gram staining method of bacterial differentiation. They are characterized by their cell envelopes, which are composed of a thin peptidoglycan cell wa ...
bacterium with polar flagella. The bacterium has a
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ...
length around 5 mega
base pair A base pair (bp) is a fundamental unit of double-stranded nucleic acids consisting of two nucleobases bound to each other by hydrogen bonds. They form the building blocks of the DNA double helix and contribute to the folded structure of both D ...
s. A number of types of citrus canker diseases are caused by different pathovars and variants of the bacterium:Gottwald, T.R, Graham, J.H. and Schubert, J.S. (2002)
Citrus canker: The pathogen and its impact
Online. ''Plant Health Progress''
*The Asiatic type of canker (canker A), ''X. axonopodis'' pv. ''citri'', caused by a group of strains originally found in Asia, is the most widespread and severe form of the disease. *Cancrosis B, caused by a group of ''X. axonopodis'' pv. ''aurantifolii'' strains originally found in South America is a disease of lemons, key lime,
bitter orange Bitter orange, Seville orange, bigarade orange, or marmalade orange is the citrus tree ''Citrus'' × ''aurantium'' and its fruit. It is native to Southeast Asia and has been spread by humans to many parts of the world. It is probably a cross be ...
, and
pomelo The pomelo ( ), ''Citrus maxima'', is the largest citrus fruit from the family Rutaceae and the principal ancestor of the grapefruit. It is a natural, non-hybrid, citrus fruit, native to Southeast Asia. Similar in taste to a sweet grapefr ...
. *Cancrosis C, also caused by strains within ''X. axonopodis'' pv. ''aurantifolii'', only infects key lime and bitter orange. *A* strains, discovered in
Oman Oman ( ; ar, عُمَان ' ), officially the Sultanate of Oman ( ar, سلْطنةُ عُمان ), is an Arabian country located in southwestern Asia. It is situated on the southeastern coast of the Arabian Peninsula, and spans the mouth of ...
,
Saudi Arabia Saudi Arabia, officially the Kingdom of Saudi Arabia (KSA), is a country in Western Asia. It covers the bulk of the Arabian Peninsula, and has a land area of about , making it the fifth-largest country in Asia, the second-largest in the Ara ...
,
Iran Iran, officially the Islamic Republic of Iran, and also called Persia, is a country located in Western Asia. It is bordered by Iraq and Turkey to the west, by Azerbaijan and Armenia to the northwest, by the Caspian Sea and Turkmeni ...
, and
India India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area, the List of countries and dependencies by population, second-most populous ...
, only infect key lime.


Pathology

Plants infected with citrus canker have characteristic lesions on leaves, stems, and fruit with raised, brown, water-soaked margins, usually with a yellow
halo Halo, halos or haloes usually refer to: * Halo (optical phenomenon) * Halo (religious iconography), a ring of light around the image of a head HALO, halo, halos or haloes may also refer to: Arts and entertainment Video games * ''Halo'' (franch ...
or ring effect around the lesion. Older lesions have a corky appearance, still in many cases retaining the halo effect. The bacterium propagates in lesions in leaves, stems, and fruit. The lesions ooze bacterial cells that, when dispersed by windblown rain, can spread to other plants in the area. Infection may spread further by
hurricane A tropical cyclone is a rapidly rotating storm system characterized by a low-pressure center, a closed low-level atmospheric circulation, strong winds, and a spiral arrangement of thunderstorms that produce heavy rain and squalls. Dep ...
s. The disease can also be spread by contaminated equipment, and by transport of infected or apparently healthy plants. Due to latency of the disease, a plant may appear to be healthy, but actually be infected. Citrus canker bacteria can enter through a plant's
stomata In botany, a stoma (from Greek ''στόμα'', "mouth", plural "stomata"), also called a stomate (plural "stomates"), is a pore found in the epidermis of leaves, stems, and other organs, that controls the rate of gas exchange. The pore is b ...
or through wounds on leaves or other green parts. In most cases, younger leaves are considered to be the most susceptible. Also, damage caused by citrus leaf miner larvae (''Phyllocnistis citrella'') can be sites for infection to occur. Within a controlled laboratory setting, symptoms can appear in 14 days following inoculation into a susceptible host. In the field environment, the time for symptoms to appear and be clearly discernible from other foliar diseases varies; it may be on the order of several months after infection. Lower temperatures increase the latency of the disease. Citrus canker bacteria can stay viable in old lesions and other plant surfaces for several months.


Pathogenicity

''Xanthomonas axonopodis'' has the capability to form a
biofilm A biofilm comprises any syntrophic consortium of microorganisms in which cells stick to each other and often also to a surface. These adherent cells become embedded within a slimy extracellular matrix that is composed of extracellular po ...
for attachment on the host. The biofilm is the result of the production of extracellular polysaccharides (
xanthan Xanthan gum () is a polysaccharide with many industrial uses, including as a common food additive. It is an effective thickening agent, emulsifier, and stabilizer that prevents ingredients from separating. It can be produced from simple suga ...
). The biofilm ensures the virulence and epiphytic survival of ''X. axonopodis'' pv. ''citri'' prior to the development of citrus canker. In addition, the bacteria secrete transcriptional activator-like (TAL) effectors through the type III secretion system. The effector interacts with host machinery to induce transcription for genes that regulate plant hormones such as
gibberellin Gibberellins (GAs) are plant hormones that regulate various developmental processes, including stem elongation, germination, dormancy, flowering, flower development, and leaf and fruit senescence. GAs are one of the longest-known classes of plan ...
and
auxin Auxins (plural of auxin ) are a class of plant hormones (or plant-growth regulators) with some morphogen-like characteristics. Auxins play a cardinal role in coordination of many growth and behavioral processes in plant life cycles and are essenti ...
.


Disease cycle

''Xanthomonas axonopodis'' pv. ''citri'' overseason in an infected area which appears as a canker lesion on leaf or stem. The bacteria ooze out of the lesions when there is free moisture. During rainy weather, wind-blown rain carries the inoculum to new susceptible hosts. The bacteria infect new plants through stomata and wounds. Pruning or hedging can cut open mesophyll tissues, creating wounds through which the plant may be directly infected. The rain can also cause water congestion on the leaf surface, form columns of water through the stomata and promote infection through natural openings. Infections can form on fruit, foliage and young stem. Leaves and stems are most susceptible to infection within the first six weeks of initial growth. Infection of fruit is most likely to occur during the 90 day period after petal fall during fruit formation. The varied size of lesions on citrus fruit is because of the multiple cycle of infections and can reflect different-aged lesions on the same fruit.


Favorable environmental conditions

Wind-driven rain plays a major role in the dispersal of ''X. axonopodis''. The bacteria are said to be readily dispersed by splashed rain and wind and the quantity of ''X. axonopodis'' declines after the first event of wind-blown rain dispersal. Apart from that, the bacteria also favor warm weather. The cases of citrus canker are more acute in areas that receive high rainfall and have high mean temperature, such as Florida. Often, cankers emerge briskly during fall, slowly during winter and most rapidly in mid to late spring.


Detection

The disease can be detected in groves and on fruit by the appearance of lesions. Early detection is critical in quarantine situations. Bacteria can be tested for pathogenicity by inoculating multiple citrus species with them. Additional diagnostic tests (antibody detection), fatty-acid profiling, and genetic procedures using
polymerase chain reaction The polymerase chain reaction (PCR) is a method widely used to rapidly make millions to billions of copies (complete or partial) of a specific DNA sample, allowing scientists to take a very small sample of DNA and amplify it (or a part of it) ...
can be conducted to confirm diagnosis and may help to identify the particular canker strain. Clara H. Hasse determined that citrus canker was not of fungoid origin but was caused by a
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were am ...
l parasite. Her research published in the 1915 ''Journal of Agricultural Research'' played a major part in saving citrus crops in multiple states.


Susceptibility

Not all species and varieties of citrus have been tested for citrus canker. Most of the common species and varieties of citrus are susceptible to it. Some species are more susceptible than others, while a few species are resistant to infection.


Management

Quarantine measures are implemented in areas where citrus canker is not endemic or has been obliterated to prevent the introduction of ''X. axonopodis.'' On the other hand, in regions where citrus canker occurs,
Integrated Pest Management Integrated pest management (IPM), also known as integrated pest control (IPC) is a broad-based approach that integrates both chemical and non-chemical practices for economic control of pests. IPM aims to suppress pest populations below the eco ...
(IPM) is utilized. The most notable feature of this management program is the transposition of susceptible citrus plants to field resistant citrus cultivars. Apart from using resistant cultivars in fields, there are several measures that are taken to control citrus canker from causing failed crop. The measures can be divided into three major categories: exclusion, eradication and sanitation.


Exclusion

Citrus trees or fruits from outside of the country are inspected to ensure they are bacteria-free trees. Under the management program, the production of ''Xac'' (''X. axonopodis'' pv. ''citri'')-free nursery trees for exclusion of canker from orchard is also mandatory. Because the bacteria can be introduced from countries with endemic canker or canker outbreaks, strict restrictions on citrus importation are implemented in citrus-growing countries. Citrus trees will only be grown on canker-free fields at least one year after effective eradication. Planting sites are also chosen to minimize favorable environmental conditions for the spread of ''X. axonopodis''. For example, areas with strong wind are avoided to decrease the dispersal of bacterial inoculum to the susceptible citrus trees.


Eradication

Once citrus canker is introduced into a field, removal of the infected trees is enacted to halt further spread of the bacteria. For instance, in Florida between 2000 and 2006, all citrus trees within of infected trees were required to be eradicated. In the process, the infected trees are uprooted and burned. In urban areas, the trees are cut down and chipped, then disposed of in landfills.


Sanitation

''X. axonopodis'' pv. ''citri'' can be transmitted by mechanical means such as humans and machinery. As a sanitation measure, the workers in citrus orchards are required to do thorough decontamination of personnel and equipment to prevent the spread of bacteria from the infected areas. Aerosol inoculum is able to cause infection in wetted foliage in the zone of bacterial dispersal. Vehicles can also become contaminated by contacting the wet foliage. Contaminated equipment and machines can be disinfected by spraying bactericide.


Distribution and economic impact

Citrus canker is thought to have originated in the area of Southeast Asia-India. It is now also present in
Japan Japan ( ja, 日本, or , and formally , ''Nihonkoku'') is an island country in East Asia. It is situated in the northwest Pacific Ocean, and is bordered on the west by the Sea of Japan, while extending from the Sea of Okhotsk in the n ...
,
South South is one of the cardinal directions or compass points. The direction is the opposite of north and is perpendicular to both east and west. Etymology The word ''south'' comes from Old English ''sūþ'', from earlier Proto-Germanic ''*sunþa ...
and
Central Africa Central Africa is a subregion of the African continent comprising various countries according to different definitions. Angola, Burundi, the Central African Republic, Chad, the Democratic Republic of the Congo, the Republic of the Co ...
, the
Middle East The Middle East ( ar, الشرق الأوسط, ISO 233: ) is a geopolitical region commonly encompassing Arabian Peninsula, Arabia (including the Arabian Peninsula and Bahrain), Anatolia, Asia Minor (Asian part of Turkey except Hatay Pro ...
,
Bangladesh Bangladesh (}, ), officially the People's Republic of Bangladesh, is a country in South Asia. It is the eighth-most populous country in the world, with a population exceeding 165 million people in an area of . Bangladesh is among the mo ...
, the
Pacific Islands Collectively called the Pacific Islands, the islands in the Pacific Ocean are further categorized into three major island groups: Melanesia, Micronesia, and Polynesia. Depending on the context, the term ''Pacific Islands'' may refer to one of se ...
, some countries in South America, and
Florida Florida is a state located in the Southeastern region of the United States. Florida is bordered to the west by the Gulf of Mexico, to the northwest by Alabama, to the north by Georgia, to the east by the Bahamas and Atlantic Ocean, and ...
. Some areas of the world have eradicated citrus canker and others have ongoing eradication programs, but the disease remains endemic in most areas where it has appeared. Because of its rapid spread, high potential for damage, and impact on export sales and domestic trade, citrus canker is a significant threat to all citrus-growing regions.


Australia

The citrus industry is the largest fresh-fruit exporting industry in Australia. Australia has had three outbreaks of citrus canker, all of which have been successfully eradicated. The disease was found twice during the 1900s in the
Northern Territory The Northern Territory (commonly abbreviated as NT; formally the Northern Territory of Australia) is an Australian territory in the central and central northern regions of Australia. The Northern Territory shares its borders with Western Aust ...
and was eradicated each time. In 2004, an unexplained outbreak occurred in central Queensland. The state and federal governments ordered all commercial groves, all noncommercial citrus trees, and all native lime trees ('' C. glauca'') in the vicinity of
Emerald Emerald is a gemstone and a variety of the mineral beryl (Be3Al2(SiO3)6) colored green by trace amounts of chromium or sometimes vanadium.Hurlbut, Cornelius S. Jr. and Kammerling, Robert C. (1991) ''Gemology'', John Wiley & Sons, New York, p ...
to be destroyed rather than trying to isolate infected trees. Eradication was successful, with permission to replant being granted to farmers by the biosecurity unit of the
Queensland Department of Primary Industries The Department of Agriculture and Fisheries is a department of the Queensland Government which aims to maximise the economic potential for Queensland's primary industries on a sustainable basis through strategic industrial development. The secti ...
in early 2009. Citrus canker was detected again in April 2018 and confirmed in May 2018 in the
Northern Territory The Northern Territory (commonly abbreviated as NT; formally the Northern Territory of Australia) is an Australian territory in the central and central northern regions of Australia. The Northern Territory shares its borders with Western Aust ...
and
Western Australia Western Australia (commonly abbreviated as WA) is a state of Australia occupying the western percent of the land area of Australia excluding external territories. It is bounded by the Indian Ocean to the north and west, the Southern Ocean to t ...
.


Brazil

Citrus is an important domestic and export crop for Brazil. Citrus agriculture is the second-most important agricultural activity in the state of São Paulo, the largest sweet orange production area in the world.Xanthomonas axonopodis pv citri GENOME PROJECT
Over 100,000 groves are in São Paulo, and the area planted with citrus is increasing. Of the estimated 2 million trees, greater than 80% are a single variety of orange, and the remainder is made up of tangerine and lemon trees. Because of the uniformity in citrus variety, the state has been adversely affected by canker, causing crop and monetary losses. In Brazil, rather than destroying entire groves to eradicate the disease, contaminated trees and trees within a 30-m radius are destroyed; by 1998, over half a million trees had been destroyed.


United States

Citrus canker was first found in the United States in 1910 not far from the
Georgia Georgia most commonly refers to: * Georgia (country), a country in the Caucasus region of Eurasia * Georgia (U.S. state), a state in the Southeast United States Georgia may also refer to: Places Historical states and entities * Related to the ...
– Florida border. Subsequently, canker was discovered in 1912 in Dade County, more than away. Beyond Florida, the disease was discovered in the Gulf states and reached as far north as
South Carolina )''Animis opibusque parati'' ( for, , Latin, Prepared in mind and resources, links=no) , anthem = " Carolina";" South Carolina On My Mind" , Former = Province of South Carolina , seat = Columbia , LargestCity = Charleston , LargestMetro = ...
. It took more than 20 years to eradicate that outbreak of citrus canker, from 1913 through 1931, $2.5 million in state and private funds were spent to control it—a sum equivalent to $28 million in 2000 dollars.Citrus Canker Animal and Plant Health Inspection Service
/ref> In 26 counties, some 257,745 grove trees and 3,093,110 nursery trees were destroyed by burning. Citrus canker was detected again on the Gulf Coast of Florida in 1986 and declared eradicated in 1994. The most recent outbreak of citrus canker was discovered in
Miami-Dade County, Florida Miami-Dade County is a county located in the southeastern part of the U.S. state of Florida. The county had a population of 2,701,767 as of the 2020 census, making it the most populous county in Florida and the seventh-most populous county in ...
, on September 28, 1995, by Louis Willio Francillon, a Florida Department of Agriculture agronomist. Despite eradication attempts, by late 2005, the disease had been detected in many places distant from the original discovery, for example, in Orange Park, 315 miles (500 km) away. In January 2000, the Florida Department of Agriculture adopted a policy of removing all infected trees and all citrus trees within a 1900-ft radius of an infected tree in both residential areas and commercial groves. Previous to this eradication policy, the department eradicated all citrus trees within 125 ft of an infected one. The program ended in January 2006 following a statement from the USDA that eradication was not feasible.United States Department of Agricultur
USDA Secretary Deputy Chuck Conner's Memo to Commissioner Bronson
Florida Department of Agriculture and Consumer Services Press Releas

/ref>


See also

* Bacterial blight of cassava * Clara H. Hasse


References


External links


Species Profile- Citrus Canker (''Xanthomonas axonopodis'')
National Invasive Species Information Center,
United States National Agricultural Library The United States National Agricultural Library (NAL) is one of the world's largest agricultural research libraries, and serves as a national library of the United States and as the library of the United States Department of Agriculture. Located ...
. Lists general information and resources for Citrus Canker.
Electronic Data Information Source - Citrus Canker
University of Florida IFAS Extension
Type strain of ''Xanthomonas axonopodis'' at Bac''Dive'' - the Bacterial Diversity Metadatabase
{{Taxonbar, from=Q2955986 Bacterial citrus diseases Xanthomonadales