Ascosphaera Aggregata
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''Ascosphaera aggregata'' is a species of fungus.


History and taxonomy

''Ascosphaera aggregata'' (''A. aggregata''), discovered in 1975 by Jens-Peder Skou is a fungus that is related to ''
Ascosphaera apis ''Ascosphaera apis'' is a species of fungus belonging to the family Ascosphaeraceae. It was one of the first entomopathogen genomes to be sequenced. It has a cosmopolitan distribution In biogeography, cosmopolitan distribution is the term fo ...
''.


Habitat and ecology

''A. aggregata'' is an
obligate parasite An obligate parasite or holoparasite is a parasitic organism that cannot complete its life-cycle without exploiting a suitable host. If an obligate parasite cannot obtain a host it will fail to reproduce. This is opposed to a facultative parasite, ...
that causes chalkbrood in bees, symptom manifestations differ depending on age of the larva. It primarily infects alfalfa leafcutting bees, '' Megachile rotundata''. ''Megachile rotundata'' infected with ''A. aggregata'' have been detected in the United States, Canada, and South America. Other bee species that ''A. aggregata'' has been seen to infect include the red mason bee ('' Osmia rufa''), the patchwork leafcutter bee ('' Megachile centuncularis''), '' Megachile pugnata'' and ''
Megachile relativa ''Megachile relativa'' is a species of bee in the family Megachilidae Megachilidae is a cosmopolitan family of mostly solitary bees. Both that their pollen-carrying structure (called a '' scopa'') is restricted to the ventral surface of the ...
''.


Growth, morphology and pathobiology

''A. aggregata'' is an obligate parasite that can cause chalkbrood by the fifth
instar An instar (, from the Latin '' īnstar'', "form", "likeness") is a developmental stage of arthropods, such as insects, between each moult (''ecdysis''), until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or ass ...
. The majority of the life cycle and growth of ''A. aggregata'' occurs in ''M. rotundata'' larvae. Infection of bee larvae occurs only via ingestion of resting spores, and is not possible via spore inhalation nor contact with the fungal vegetative form. Spores develop in the larva and cause it to swell, bursting the larval integument (giving the dead larvae a ragged appearance) and furthering the spread of the fungus. Buildup of larval cadavers traps the unaffected emerging bees, forcing them to chew through the cadavers and be covered in spores. Bees covered in spores then contaminate food provisions for other broods and spread the infection. Early vegetative growth utilizes gut lumen nutrients. ''A. aggregata'' grows through the midgut wall to the hemocoele (event trigger is unknown, not because of lack of space nor food) eventually replacing larval tissue. Resulting larva are filled with a
mycelial 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 ...
mat comprising two layers: a dense inner layer and a less dense outer layer.


Sexual development

Ascospore morphology consists of two layers: an inner chitinous and smooth layer, and an outer layer that is rough, spotted, and not composed of chitin nor cellulose.;) Ascospore development in ''A. aggregata'' is very unique and the resulting structure is referred to as a "spore cyst", or "ascocyst" or "synascus". Sexual development occurs on the outer mycelial mat in the subcuticular region, and is documented to proceed as follows: # The vegetative hyphae tips swell and form a
thallus Thallus (plural: thalli), from Latinized Greek (), meaning "a green shoot" or "twig", is the vegetative tissue of some organisms in diverse groups such as algae, fungi, some liverworts, lichens, and the Myxogastria. Many of these organisms wer ...
# The middle of the thallus grows and forms a nutriocyte (previously referred to as an archicarp) # The apical portion differentiates into th
trichogyne
cell. # Compatible trichogyne fuse and initiate plasmogamy. # Resulting
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 ...
fungal protoplasm then enters the nutriocyte and causes enlargement of the nutriocyte. # Nutriocyte growth causes the integument to rupture and initiate development of a fragile spherical structure without a cell wall. # Individual spores then pack together into a seemingly membrane-less spore ball. # Multiple spore balls then join and form a spore cyst. # Cell wall deposition changes spore colour from opaque white to grey to dull black


Physiology

''A. aggregata'' has been found to be unable to break down
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 ...
.


Diagnostic Considerations

Although ascospore development is very unique, it is very hard to identify ''A. aggregata'' because the spore balls and conidia tend to resemble other species. Recent investigations by James and Skinner (2005) have discovered that
PCR PCR or pcr may refer to: Science * Phosphocreatine, a phosphorylated creatine molecule * Principal component regression, a statistical technique Medicine * Polymerase chain reaction ** COVID-19 testing, often performed using the polymerase chain r ...
of the ITS domain of ribosomal DNA with species specific
primer Primer may refer to: Arts, entertainment, and media Films * ''Primer'' (film), a 2004 feature film written and directed by Shane Carruth * ''Primer'' (video), a documentary about the funk band Living Colour Literature * Primer (textbook), a t ...
sets allows the detection of fungal DNA (working, even, in asymptomatic individuals). The PCR technique can also be used on hair and honey samples to avoid the difficulty of culturing spores, as spore were shown before to only germinate well in lipids. Storage of the fungus has also proven to be difficult as it collapses after 1–2 months during normal culture passaging. However, Jensen et al. (2009) found that spores could be preserved via cryopreservation or freeze-drying whereas hyphae unfortunately could not be preserved.


Economic importance

''M. rotundata'' is the primary pollinator of the commercially grown alfalfa seed, accounting for 46,000 metric tonnes of North American alfafa seed (two-thirds the global production) in 2004. ''M. rotundata'' is also the second most valuable field crop pollinator, behind the honey bee, because of the value of alfalfa in animal feed and hay. ''A. aggregata'' has been killing this economic pollinator in the US since 1972 and has been reported to be able to kill greater than 50% of a population. Effective management of the fungus has yet to be discovered, as the current registered treatment in Canada (paraformaldehyde fumigation of spores) involves a carcinogen and other treatment options (heat and chloride treatments) are expensive and labour-intensive.


References

{{Taxonbar , from=Q85743263 Onygenales