Banana Freckle
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Banana freckle is a disease caused by the
fungus A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from th ...
'' Guignardia musae'' (
teleomorph In mycology, the terms teleomorph, anamorph, and holomorph apply to portions of the life cycles of fungi in the phyla Ascomycota and Basidiomycota: *Teleomorph: the sexual reproductive stage (morph), typically a fruiting body. *Anamorph: an ase ...
) or '' Phyllosticta musarum'' (
anamorph In mycology, the terms teleomorph, anamorph, and holomorph apply to portions of the life cycles of fungi in the phyla Ascomycota and Basidiomycota: *Teleomorph: the sexual reproductive stage (morph), typically a fruiting body. *Anamorph: an ase ...
). Generally, the causal agent of disease is referred to as ''Guignardia-Phyllosticta'' sp. There are several different strains of the fungus that exist to infect different banana varieties around the globe. Symptoms include yellowing of the tissue and formation of small dark brown spots on the leaves and fruit. Within the spots,
conidia A conidium ( ; ), sometimes termed an asexual chlamydospore or chlamydoconidium (), is an asexual, non-motile spore of a fungus. The word ''conidium'' comes from the Ancient Greek word for dust, ('). They are also called mitospores due to the ...
or
pycnidia A pycnidium (plural pycnidia) is an asexual fruiting body produced by mitosporic fungi, for instance in the order Sphaeropsidales ( Deuteromycota, Coelomycetes) or order Pleosporales (Ascomycota, Dothideomycetes). It is often spherical or inve ...
can be found. Banana freckle is easily propagated and spread from plant to plant by rain splash and movement of infected tissue or fruit. Management of the disease consists of cutting out infected leaves, using the paper bag method,
fungicide Fungicides are biocidal chemical compounds or biological organisms used to kill parasitic fungi or their spores. A fungistatic inhibits their growth. Fungi can cause serious damage in agriculture, resulting in critical losses of yield, quality, ...
application, and proper sanitation techniques. This devastating disease is extremely relevant for the major banana exporting countries of the world. In the absence of chemical control, there is about a 78% yield loss. Banana freckle disease needs to be carefully monitored in order to prevent further spread of the disease.


Hosts and symptoms

Several strains of this pathogen exist, and some are species-specific. For example, the fungal strain that infects Australian bananas,
Lady Finger Lady finger(s), ladyfinger(s), lady's finger, or ladies' fingers may refer to: Food * Ladyfinger (biscuit), a type of sponge cake * Ladyfinger, a kirsch cocktail Botany * Okra, or ladies' fingers, a pod vegetable plant * Ladyfinger cactus, a ...
and
Bluggoe Bluggoe, Orinoco, ''Musa'' Orinoco, or burro is a cultivar of banana. Genome Bluggoe is a triploid ABB cultivar. Cultivation Bluggoe is a cold hardy banana, growing in USDA zones 810 or 710. Tree to tall. Width of leaves same dimensions. Flo ...
, does not infect
Cavendish banana Cavendish bananas are the fruits of one of a number of banana cultivars belonging to the Cavendish subgroup of the AAA banana cultivar group. The same term is also used to describe the plants on which the bananas grow. They include commercially ...
s. Individual strains are found in various locations across the globe (see “Environment” section for more information on pathogen strains). Banana freckle are not known to infect any other types of fruits. Symptoms typically appear after 2–4 weeks after the banana cluster has opened, and seem to be primarily localized on older leaves. Infected tissues may also yellow with age, and eventually
senesce Senescence () or biological aging is the gradual deterioration of functional characteristics in living organisms. The word ''senescence'' can refer to either cellular senescence or to senescence of the whole organism. Organismal senescence invol ...
in the most severe case of symptoms. Even in the cases of minor infections, the fungus renders the banana fruit unmarketable at the loss of the grower. Major macroscopic diagnostic signs include small brown to dark brown specks on leaves and fruits, which indicate the presence of fungal
spores In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, f ...
protruding through the infected tissues. Depending on which stage (sexual or asexual) the fungus completes its life cycle, the spots will be either pycnidia or conidia. Microscopically, these dark spots, or freckles, can range from in diameter to . As is the case for most fungal diseases, banana freckle is diagnosed based on the presence of its spores. Fruiting bodies and spores obtained from lesions and fungal cultures are key components when identifying the
pathogen In biology, a pathogen ( el, πάθος, "suffering", "passion" and , "producer of") in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a germ ...
. The causal strain of the fungus can then be determined by the type of banana it has infected.


Disease cycle


Contact

Spores land on
banana A banana is an elongated, edible fruit – botanically a berry – produced by several kinds of large herbaceous flowering plants in the genus ''Musa''. In some countries, bananas used for cooking may be called "plantains", distinguis ...
plants or fruits by rain splash or movement of infected tissue or fruit.


Germination and Infection

The spores that landed on the plant
germinate Germination is the process by which an organism grows from a seed or spore. The term is applied to the sprouting of a seedling from a seed of an angiosperm or gymnosperm, the growth of a sporeling from a spore, such as the spores of fungi, fer ...
and as
hyphae A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one or ...
grow on the outside of the host into masses known as
mycelium Mycelium (plural mycelia) is a root-like structure of a fungus consisting of a mass of branching, thread-like hyphae. Fungal colonies composed of mycelium are found in and on soil and many other substrate (biology), substrates. A typical single ...
, they also form
appressoria An appressorium is a specialized cell typical of many fungal plant pathogens that is used to infect host plants. It is a flattened, hyphal "pressing" organ, from which a minute infection peg grows and enters the host, using turgor pressure capable ...
in order to penetrate the host cells. After entering, the hyphae form a
haustorium In botany and mycology, a haustorium (plural haustoria) is a rootlike structure that grows into or around another structure to absorb water or nutrients. For example, in mistletoe or members of the broomrape family, the structure penetrates t ...
which allows them to absorb nutrients through the
plasma membrane The cell membrane (also known as the plasma membrane (PM) or cytoplasmic membrane, and historically referred to as the plasmalemma) is a biological membrane that separates and protects the interior of all cells from the outside environment (t ...
of the host cell.


Reproduction

* Asexual – ''Phyllosticta musarum'' – Conidia in Pycnidium * Sexual – ''Guignardia musae'' – Hyphae of opposite mating types undergo
plasmogamy Plasmogamy is a stage in the sexual reproduction of fungi, in which the protoplasm of two parent cells (usually from the mycelia) fuse without the fusion of nuclei, effectively bringing two haploid nuclei close together in the same cell. This sta ...
and then
karyogamy Karyogamy is the final step in the process of fusing together two haploid eukaryotic cells, and refers specifically to the fusion of the two nuclei. Before karyogamy, each haploid cell has one complete copy of the organism's genome. In order for ...
to form
ascocarps An ascocarp, or ascoma (), is the fruiting body ( sporocarp) of an ascomycete phylum fungus. It consists of very tightly interwoven hyphae and millions of embedded asci, each of which typically contains four to eight ascospores. Ascocarps are mo ...
with
asci ASCI or Asci may refer to: * Advertising Standards Council of India * Asci, the plural of ascus, in fungal anatomy * Accelerated Strategic Computing Initiative * American Society for Clinical Investigation * Argus Sour Crude Index * Association of ...
.


Development

Dikaryotic The dikaryon is a cell nucleus, nuclear feature which is unique to certain fungi. (The green alga ''Derbesia'' had been long considered an exception, until the heterokaryotic hypothesis was challenged by later studies.) Compatible cell-types can fus ...
hyphae grow and develop ascocarps, where asci develop diploid
ascospores An ascus (; ) is the sexual spore-bearing cell produced in ascomycete fungi. Each ascus usually contains eight ascospores (or octad), produced by meiosis followed, in most species, by a mitotic cell division. However, asci in some genera or ...
which undergo
meiosis Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately resu ...
to become
haploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
.


Dispersal

Asci survive in tissue and can be spread to new host by water or movement of infected tissue.


Environment

Banana freckle has been identified in 27 countries. Warm, humid environments like that of Southeast Asia, Australia, and Oceania are ideal growing conditions for the causal genera guignardium and phyllosticta. The high annual precipitation of these tropical areas is also very conducive to the spread of the spores by splash.


Management

There are a number of control methods to prevent and reduce the banana freckle disease. The paper bag method seems to be the most effective way to gain physical control of the pathogen. The infected leaves are the primary source of spores, and placing a bag over the bananas, once harvested, creates a barrier to prevent
inoculum In biology, inoculum refers to the source material used for inoculation. ''Inoculum'' may refer to: * In medicine, material that is the source of the inoculation in a vaccine * In microbiology, propagules: cells, tissue, or viruses that are used t ...
from spreading to the fruit. Some cultural controls include pruning out infectious plant material, planting in pathogen-free fields, and practicing proper sanitation techniques. In the Philippines, pruning and cutting out patches of infected tissue have prevented the spread of the pathogen in the plant during disease outbreaks. General sanitation practices have also reduced the spread of inoculum. When planters failed to maintain sanitary equipment, seeds, and soil, they witnessed severe fruit infections. The more freckles seen on the leaves of the plant, the more the fruit develops symptoms of the disease. Inversely, less freckles corresponded to less disease. In addition, multiple fungicides have been seen to reduce banana freckle disease. In Hawaii, spraying the leaves and fruit with
maneb Maneb is a fungicide and a polymeric complex of manganese with the ethylene bis(dithiocarbamate) anionic ligand.Arno H. Reidies "Manganese Compounds" Ullmann's Encyclopedia of Chemical Technology 2007; Wiley-VCH, Weinheim. Applications It can be ...
( water plus of sticker-spreader) every 2 weeks or once a month throughout the year has remarkably reduced the spread of inoculum. In Taiwan, spraying fungicides, such as
phaltan Phaltan () is a town, a tehsil, and a municipal council in the Satara district in the Indian state of Maharashtra. The town is about northeast of the city of Satara and about 110 km from Pune. History Phaltan was one of the non-salu ...
, orthocide,
chlorothalonil Chlorothalonil (2,4,5,6-tetrachloroisophthalonitrile) is an organic compound mainly used as a broad spectrum, nonsystemic fungicide, with other uses as a wood protectant, pesticide, acaricide, and to control mold, mildew, bacteria, algae. Chlor ...
,
dithiocarbamates In organic chemistry, a dithiocarbamate is a functional group with the general formula and structure . It is the analog of a carbamate in which both oxygen atoms are replaced by sulfur atoms (when only 1 oxygen is replaced the result is thiocar ...
, and
propiconazole Propiconazole is a triazole fungicide, also known as a DMI, or demethylation inhibiting fungicide due to its binding with and inhibiting the 14-alpha demethylase enzyme from demethylating a precursor to ergosterol. Without this demethylation st ...
, biweekly have produced effective results against the disease. In the Philippines, chemical controls used against
Black Black is a color which results from the absence or complete absorption of visible light. It is an achromatic color, without hue, like white and grey. It is often used symbolically or figuratively to represent darkness. Black and white have o ...
or Yellow Sigatoka disease have been helpful. These consist of
mancozeb Mancozeb is a dithiocarbamate non-systemic agricultural fungicide with multi-site, protective action on contact. It is a combination of two other dithiocarbamates: maneb and zineb. The mixture controls many fungal diseases in a wide range of field ...
,
triazoles A triazole is a heterocyclic compound featuring a five-membered ring of two carbon atoms and three nitrogen atoms with molecular formula C2H3N3. Triazoles exhibit substantial isomerism, depending on the positioning of the nitrogen atoms within the ...
, tridemorph, and
strobilurin Strobilurins are a group of natural products and their synthetic analogs. A number of strobilurins are used in agriculture as fungicides. They are part of the larger group of QIs (Quinone outside Inhibitors), which act to inhibit the respiratory ch ...
. Mancozeb seems to be the most effective fungicide against banana freckle disease in Hawaii and the Philippines.Corcolon, Benny M.; Raymundo, Avelino D. (2008)
"Estimating Yield Loss in Banana Due to Freckle Disease Caused by ''Phyllosticta musarum'' (Cke.) Van Der Aa"
''Philippine Journal of Crop Science'' 33(2): 75–85.
These fungicides do not eliminate the pathogen completely, but they reduce the inoculum levels and eventually reduce yield loss. Lastly, eradication of infected plants can prevent further infection of other fruit around the area.


Importance

Banana freckle disease has become a major problem for many countries involved with banana export and production. In the absence of disease management, there is an estimated 78% yield loss. However, the expenses that go toward methods for disease control is 25% of the selling price. Although the fruit is still edible when affected by the disease, market value is reduced.
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 north ...
does not tolerate any symptoms and will not purchase any bananas with suspicious blemishes. This causes problems for the exporting business in Taiwan and the Philippines. Banana freckle was first identified in Hawaii in 1917 and then soon after in the Philippines. The disease is most common in Hawaii, Indonesia, Thailand, Taiwan, and New Guinea to name a few. This devastating fungal disease can infect the leaves during any stage of the plant’s life cycle. In July 2013, ''Phyllosticta cavendishii'' (banana freckle) was detected in the Northern Territory on the Cavendish variety of bananas on two rural properties. In January 2019, after a successful eradication program, absence of banana freckle has been declared for Australia (exclusive of external territories).


References

{{reflist Fungal fruit diseases Banana diseases