Penicillium Expansum
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''Penicillium expansum'' is a
psychrophilic Psychrophiles or cryophiles (adj. ''psychrophilic'' or ''cryophilic'') are extremophilic organisms that are capable of growth and reproduction in low temperatures, ranging from to . They have an optimal growth temperature at . They are found in ...
blue mold that is common throughout the world in soil. It causes Blue Mold of apples, one of the most prevalent and economically damaging post-harvest diseases of apples. Though primarily known as a disease of apples, this
plant pathogen Plant pathology (also phytopathology) is the scientific study of diseases in plants caused by pathogens (infectious organisms) and environmental conditions (physiological factors). Organisms that cause infectious disease include fungi, oomyc ...
can infect a wide range of hosts, including pears, strawberries, tomatoes, corn, and rice. ''Penicillium expansum'' produces the carcinogenic metabolite
patulin Patulin is an organic compound classified as a polyketide. It is a white powder soluble in acidic water and in organic solvents. It is a lactone that is heat-stable, so it is not destroyed by pasteurization or thermal denaturation.http://www.s ...
, a
neurotoxin Neurotoxins are toxins that are destructive to nerve tissue (causing neurotoxicity). Neurotoxins are an extensive class of exogenous chemical neurological insultsSpencer 2000 that can adversely affect function in both developing and mature ner ...
that is harmful when consumed. Patulin is produced by the fungus as a
virulence factor Virulence factors (preferably known as pathogenicity factors or effectors in plant science) are cellular structures, molecules and regulatory systems that enable microbial pathogens (bacteria, viruses, fungi, and protozoa) to achieve the following ...
as it infects the host. Patulin levels in foods are regulated by the governments of many developed countries. Patulin is a particular health concern for young children, who are often heavy consumers of apple products. The fungus can also produce the mycotoxin
citrinin Citrinin is a mycotoxin which is often found in food. It is a secondary metabolite produced by fungi that contaminates long-stored food and it causes different toxic effects, like nephrotoxic, hepatotoxic and cytotoxic effects. Citrinin is mainly ...
.


Hosts and disease development

''Penicillium expansum'' has a wide host range, causing similar symptoms on fruits which include apples, pears, cherries, and citrus . Initial infection most often occurs at sites of fruit injury, such as bruises or puncture wounds. Although infections may start in the field, infected spots often become evident post-harvest, and expand while fruit is in storage. Infected areas are clearly delineated and light brown, and soft decaying tissue can be easily "scooped" out of the surrounding healthy tissue., Spore masses later appear on the surfaces of infected fruit, initially appearing as white mycelium, then turning blue to blue-green in color as the asexual spores mature. Fruit affected by ''P. expansum'' typically has an earthy, musty odor. Lesions measure 1–1.25 inches in diameter eight to ten weeks after infection if kept under cold storage conditions. Age factors into ''P. expansum'' infection, in that overripe or mature fruits are most susceptible to infection, while those picked underripe are less likely to become infected. In apples, the colors of the lesions may vary with variety, from lighter-brown on green and yellow apple varieties to dark-brown on the deeper-red and other darker-color varieties. Varieties particularly susceptible to ''P. expansum'' infection include McIntosh, Golden Supreme, and Golden Delicious. Both sweet and sour cherries are affected by ''P. expansum''. Cherry varieties found to be particularly susceptible to ''P. expansum'' infection were mainly early varieties, including Navalinda and Burlat.


Diagnosis

''Penicillium expansum'' can be identified by its morphological characteristics and secondary metabolites in fruit or in axenic culture. The presence of the secondary metabolite patulin can suggest ''P. expansum'' infection, but this method is not species-specific as a number of different ''Penicillium'' species and their allies produce patulin. Patulin presence can be assayed using high-performance liquid chromatography with ultraviolet detection. Molecular methods based on species-specific genes can speed identification.


Environment

''Penicillium expansum'' grows best in wet, cool (<25C) conditions. ''P. expansum'' was found to grow most efficiently in a temperature range of 15-27 degrees Celsius, with slower growth at lower and higher temperatures. ''P. expansum'' grows best in wet conditions; growth rate was fastest at a relative humidity of 90%. ''P. expansum'' infection acidifies host tissues via the secretion of organic acids, and that acidification enhances fungal development, indicating a link between environmental acidity and ''P. expansum'' virulence.


Disease cycle

''P. expansum'' infects a fruit via wounds through which the
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 ...
are able to enter. Usually, puncturing, bruising, and limb rubs occur during harvesting, packaging, and processing of the fruit, all of which provide sites through which spores can enter the fruit. Conidia can be found in soil, decaying debris, and tree bark, and can survive cold temperatures. Conidia may be isolated from the air of the orchard and packaging house, on the walls of the packaging houses, and from the water and fungicide solution into which harvested fruits are dunked before packaging or storage. Exposure to conidia at any step of growth, harvesting, processing, shipping, and storage can lead to inoculation and disease. Conidia that have gained access via a wound can germinate to form a germ tube. This germ tube will continue to grow as hyphae which colonize the fruit, killing fruit cells in an expanding infection. If the fungus has colonized the fruit with mycelium, the formation of conidiophores occurs on the surface or subsurface of the hyphae. The conidiophores are mostly smooth-walled terverticillate penicilli. A terverticillate pencilii has multiple branch points below the phialides, the cells that the conidia are attached to. However, at times, the penicilli may be rough or biverticillate (only two levels of branching). The phialides are packed close together with nearly a cylindrical shape. The conidia are dry, smooth, elliptical, and "dull-green" in color and are often disseminated by wind currents. Sexual reproduction has not been observed in nature for ''P. expansum''.


Management

Due to the susceptibility to infection of mature and overripe fruit, post-harvest treatment of fruit with fungicides is the most common method of combating ''P. expansum''. Proper sanitation and careful handling of the fruit are two non-chemical methods that can help control the disease. Good sanitation reduces contact with orchard soil either on the fruit or in transportation containers. And since the fungus needs a wound to infect, careful handling can reduce infection even when the fungus is present. Chemical treatment with a chlorine bath can be effective in killing spores. Biofungicides using active ingredients such as bacteria and yeast have been successful in preventing infection but are ineffective against existing infections.


Importance

''Penicillium expansum'' produces the mycotoxin
patulin Patulin is an organic compound classified as a polyketide. It is a white powder soluble in acidic water and in organic solvents. It is a lactone that is heat-stable, so it is not destroyed by pasteurization or thermal denaturation.http://www.s ...
, a neurotoxin that can enter the food supply via apples and apple products such as juice and cider. China and the United States are the leading producers of apples in the world. In all, roughly 69 million tons of apples were grown worldwide. In the United States, approximately 60% of all apples are grown in Washington state. In Washington, 10-12 billion apples are hand-picked each year and the industry is the largest in the state. Considering the size of the apple product industry and the large number of people that may come into contact with infected fruits, control of ''P. expansum'' is vitally important.


References


External links


USDA ARS Fungal Database
{{Taxonbar, from=Q147358 Fungal plant pathogens and diseases Apple tree diseases expansum Taxa named by Johann Heinrich Friedrich Link Fungi described in 1809