Fungal Effectors
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Fungal effectors are proteins or non-proteinaceous molecules (such as RNAs or small molecules) secreted by pathogenic fungi into a host organism in order to modulate the host's immune response.


Plant pathogenic fungi

In the first stages of infection, conserved molecules from the fungal pathogen's cell wall, such as polysaccharides and
chitin Chitin ( C8 H13 O5 N)n ( ) is a long-chain polymer of ''N''-acetylglucosamine, an amide derivative of glucose. Chitin is probably the second most abundant polysaccharide in nature (behind only cellulose); an estimated 1 billion tons of chit ...
, are recognised by membrane-localised pattern recognition receptors (PRRs) on the plant host's side. Such conserved molecules are generally described as pathogen-associated molecular patterns (PAMPs) or microbe-associated molecular patterns (MAMPs) and the initial innate immune response that their recognition triggers is known as PAMP-triggered immunity (PTI). In order to counteract PTI, fungal pathogens secrete effector proteins into the host, some of which may directly inhibit components of the innate immune response cascade. One example is the conserved effector NIS1, present in fungal pathogens from the
Ascomycota 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 def ...
and
Basidiomycota Basidiomycota () is one of two large divisions that, together with the Ascomycota, constitute the subkingdom Dikarya (often referred to as the "higher fungi") within the kingdom Fungi. Members are known as basidiomycetes. More specifically, Basi ...
phyla. NIS1 blocks PAMP-triggered immune responses by interacting with the PRR-associated kinases BAK1 and BIK1 and preventing these kinases from interacting with their downstream partners. To protect themselves from the actions of effector proteins, plants have evolved resistance proteins (R proteins), which may in turn recognise an effector and trigger a second tier of immune responses, known as
effector-triggered immunity Effector-triggered immunity (ETI) is one of the pathways, along with the Pattern-Triggered Immunity (PTI) pathway, by which the innate immune system recognises pathogenic organisms and elicits a protective immune response. ETI is elicited when an ...
(ETI).


Site of action

Plant pathogenic fungi use two distinct effector secretion systems and each secretory pathway is specific to an effector family: * apoplastic effectors act in the apoplast, the extracellular space outside the host plant's cells. In the model pathogen ''
Magnaporthe oryzae ''Magnaporthe'' is a genus of ascomycete fungi. Several of the species are cereal pathogen In biology, a pathogen ( el, πάθος, "suffering", "passion" and , "producer of") in the oldest and broadest sense, is any organism or agent that ca ...
,'' apoplastic effectors are secreted into a distinct compartment enclosing the growing hypha named the EIHM (extra-invasive hyphal membrane). * cytoplasmic effectors enter the host cells' cytoplasm. Cytoplasmic effectors of the pathogen ''
Magnaporthe oryzae ''Magnaporthe'' is a genus of ascomycete fungi. Several of the species are cereal pathogen In biology, a pathogen ( el, πάθος, "suffering", "passion" and , "producer of") in the oldest and broadest sense, is any organism or agent that ca ...
'' are accumulated into a complex plant-derived structure named the biotrophic interfacial complex (BIC) and they are later translocated across the EIHM inside the plant cell. It has been shown that cytoplasmic effectors can move through a few layers of plant cells, probably a way to prepare them for hyphal invasion.


Fungal pathogens


References

{{reflist Fungal proteins