Venturia inaequalis
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''Venturia inaequalis'' is an
ascomycete Ascomycota is a phylum of the kingdom Fungi that, together with the Basidiomycota, forms the subkingdom Dikarya. Its members are commonly known as the sac fungi or ascomycetes. It is the largest phylum of Fungi, with over 64,000 species. The defi ...
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 fr ...
that causes the
apple scab Apple scab is a common disease of plants in the rose family (Rosaceae) that is caused by the ascomycete fungus ''Venturia inaequalis''. While this disease affects several plant genera, including ''Sorbus, Cotoneaster,'' and ''Pyrus'', it is mos ...
disease.Evidence of Two Formae Speciales in Venturia inaequalis
/ref>


Systematics

''Venturia inaequalis''
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 a ...
s have been described under the names ''Fusicladium dendriticum'' and ''Spilocaea pomi''. Whether ''V. inaequalis'' is a single species or contains several
cryptic species In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each ot ...
has been a matter of debate for a long time. Recent genetic studies have revealed a considerable uniformity of the species. In addition, the fungus '' Spilocaea pyracanthae'', a parasite of ''
Pyracantha ''Pyracantha'' (from Greek "fire" and "thorn", hence firethorn) is a genus of large, thorny evergreen shrubs in the family Rosaceae, with common names firethorn or pyracantha. They are native to an area extending from Southwest Europe east to ...
'' appeared not to genetically differ from ''V. inaequalis'', being thus a special form of the latter.


Morphology

The
fruiting bodies The sporocarp (also known as fruiting body, fruit body or fruitbody) of fungi is a multicellular structure on which spore-producing structures, such as basidia or asci, are borne. The fruitbody is part of the sexual phase of a fungal life cyc ...
,
ascocarp 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 mos ...
s appear in the form of pseudothecia. They are solitary and embedded into the host plant tissue. A pseudothecium has small dark hairs around its opening, and contains
pseudoparaphyses {{Short pages monitor


Effectors

Effectors are proteins encoded by pathogens, which act to effect a response from a host cell - often modulating the host immune response. Where a host variety is able to recognise and mount a resistance response to the presence of an effector, the effector is referred to as an
Avirulence The gene-for-gene relationship was discovered by Harold Henry Flor who was working with rust (''Melampsora lini'') of flax ('' Linum usitatissimum''). Flor showed that the inheritance of both resistance in the host and parasite ability to cause di ...
protein. Presently, only one effector gene, ''AvrVg'', eliciting a resistance response in
Apple An apple is an edible fruit produced by an apple tree (''Malus domestica''). Apple trees are cultivated worldwide and are the most widely grown species in the genus '' Malus''. The tree originated in Central Asia, where its wild ancest ...
has been identified in ''V. inaequalis''


Hosts and symptoms

The ''Venturia inaequalis'' pathogen is a fungal organism that produces similar symptoms across a range of woody hosts. These include the common pear (''Pyrus spp.)'', firethorn (''Pyracantha spp.)'', mountain ash (''Sorbus spp.''), and most notably both commercial apples along with ornamental crabapples (''Malus spp.''). Symptoms of the infection occur on leaves, fruit, flowers, and young green shoots. Foliar symptoms begin to occur in the early spring around budbreak and mainly present as light green lesions that progress to an olive-brown color with a velvety texture due to conidia formation as time passes. These large scab-like lesions can warp the leaf's shape and can eventually lead to defoliation. Lesions formed by primary infection via ascospores tend to have more distinct borders when compared to lesions as a result of a secondary infection cycle via conidia. Young fruit, often infected by foliar conidia, can also display similar symptoms to infected leaves. In this case the lesions progress to bare, brown and corky spots. The apple skin and flesh can split open as the fruit enlarges, though young fruits often prematurely drop. Mature fruits are more resistant to infection and only form small, black ‘pin-head scabs’ which might not even be noticeable until after storage.


Importance

Economic loses due to apple scab over an extended period of time far outweigh the impact of any other apple pathogen. Historically one can find examples the symptoms of ''V.  inaequalis'' in paintings as far back as the fifteenth century. Fruit production can be limited due to defoliation while limiting fruit bud creation in the following year.  While the disease can cause total crop loss in optimal conditions without management, the main economic impact is due to the reduction in both size and marketable quality of the fruit. This issue is further compounded by the fact that apple cultivars with a high market share, Pink Lady for example, are susceptible to the pathogen while more resistant varieties are less well known or desired by consumers. At an industrial production level these threats are only reliably mitigated by expensive, labor intensive and repeated spraying of pesticides. In regions of apple production where the year's weather is conducive to infection up to 70% of the pesticides applied are used to control for the effects of apple scab. While the main host of economic interest are various apple cultivars the other host species are still affected by the pathogen. Considering most of these are commercially used as ornamental species, flowering crab apples for example, the importance of the pathogen shifts to be a more aesthetic nuisance. In the academic sense ''V. inaequalis'' has proved invaluable in the realm of genetic research of pathogenicity. It is one of the first ascomycete fungi to undergo genetic analysis and continues to be useful in that field. The fungi's mechanistic similarity to obligate parasites while still being able to be cultured in media has led to its repeated use in the study of the genes related to pathogenicity.


Management

Protection from initial inoculation, either via sexual ascospores or asexual conidia, with fungicide is the main form of pathogen management. Spray schedules should be created with plant and pathogen development timings in mind and thus should begin with an initial spray at budswell and repeated in 10 to 14 day intervals. Specifics of the spray intervals should also be determined by considering what spray is used, weather patterns (mainly rain and moisture), host species, host growth patterns, and the amount of fungal inoculum present. Cultural practices can also be used to limit or prevent apple scab infection. Orchard design and planting patterns focusing on increased aeration are important in ensuring that susceptible tissues dry prior to initial infection. Proper pruning can further aid this effect. Another form of cultural management is the implementation of proper sanitation. Fallen infected leaves in the fall should be collected and destroyed to reduce the total inoculum able to overwinter and infect the following year. Primary infection is mainly caused by the ascospores that overwinter in the fallen debris and the density of these spores in the spring is directly related to the speed and intensity of an apple scab outbreak. In addition to direct removal of fallen infected tissues, mulching of fallen leaves into the soil to destroy the ascospores is advisable. Application of nitrogenous fertilizer to speed up this process can be done immediately prior to leaf fall or as a ground application. More proactive methods of management exist in the form of genetically resistant cultivars and species. Apple cultivars such as Enterprise, Goldrush, Liberty, Jonafree, Macfree, Prima, Pristine, Redfree, and Sir Prize are resistant to apple scab infection along with multiple varieties of ornamental crab apples. Breeding of resistant lines has seen minimal acceptance in the Americas but more widespread success in Europe, though the durability of this resistance is always a concern.  More recently, biofungicidal methods of control have emerged in studies with some promise being shown by the organism ''Microsphaeropsis ochracea'' in reduction of initial ascospore inoculum of up to 70-80 percent. While still requiring a fungicide application, the use of the proposed mycoparasite allows for a delayed and lower volume use of chemicals.


References


External links

* Managing Resistance of ''Venturia inaequalis'' to the Strobilurin Fungicide

* Apple Scab, Plant Diagnostic Clinic, Cornell Universit

* Apple Scab, NYS IPM Tree Fruit Crops Fact Sheet Series (Revised), Wayne Wilco

* ''Venturia inaequalis'': Tom Volk's Fungus of the Month, September 200

* Apple and Pear Scab, Keepers Nurser

{{Taxonbar, from=Q10675708 Fungi described in 1866 Venturiaceae Fungal tree pathogens and diseases Apple tree diseases