Ascosphaera aggregata

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Ascosphaera aggregata
Ascosphaera journal.pone.0073419.g004.png
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Fungi
Division: Ascomycota
Class: Eurotiomycetes
Order: Onygenales
Family: Ascosphaeraceae
Genus: Ascosphaera
Species:
A. aggregata
Binomial name
Ascosphaera aggregata
Skou, JP (1975) [1]

Ascosphaera aggregata is a species of fungus.

History and taxonomy

Ascosphaera aggregata, discovered in 1975 by Jens-Peder Skou [1] is a fungus that is related to Ascosphaera apis . [2]

Contents

Habitat and ecology

Ascosphaera aggregata is an obligate parasite [3] that causes chalkbrood in bees, [4] symptom manifestations differ depending on age of the larva. [5] It primarily infects alfalfa leafcutting bees, Megachile rotundata . [2] [6] [7] [3] [8] Megachile rotundata infected with A. aggregata have been detected in the United States, Canada, [9] and South America. [5] Other bee species that A. aggregata has been seen to infect include the red mason bee ( Osmia rufa [8] ), the patchwork leafcutter bee ( Megachile centuncularis [8] ), Megachile pugnata and Megachile relativa . [2] [10] [8]

Growth, morphology and pathobiology

Ascosphaera aggregata is an obligate parasite that can cause chalkbrood by the fifth instar. [5] 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, [3] and is not possible via spore inhalation nor contact with the fungal vegetative form. [6] [5] Spores develop in the larva and cause it to swell, bursting the larval integument (giving the dead larvae a ragged appearance) [1] 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. [7] Bees covered in spores then contaminate food provisions for other broods [7] and spread the infection.

Early vegetative growth utilizes gut lumen nutrients. [6] A. aggregata grows through the midgut wall to the hemocoele (event trigger is unknown, not because of lack of space nor food) [6] eventually replacing larval tissue. [3] Resulting larvae are filled with a mycelial mat comprising two layers: a dense inner layer and a less dense outer layer. [6]

Sexual development

Ascospore morphology consists of two layers: an inner chitinous and smooth layer, and an outer layer that is rough, spotted, [1] and not composed of chitin nor cellulose.; [6] ) Ascospore development in A. aggregata is very unique and the resulting structure is referred to as a "spore cyst", or "ascocyst" or "synascus". [8] Sexual development occurs on the outer mycelial mat in the subcuticular region, [3] [6] and is documented to proceed as follows:

  1. The vegetative hyphae tips swell and form a thallus [6]
  2. The middle of the thallus grows and forms a nutriocyte (previously referred to as an archicarp [8] )
  3. The apical portion differentiates into the trichogyne cell. [6]
  4. Compatible trichogyne fuse and initiate plasmogamy. [6]
  5. Resulting dikaryotic fungal protoplasm then enters the nutriocyte and causes enlargement of the nutriocyte. [6]
  6. Nutriocyte growth causes the integument to rupture and initiate development of a fragile spherical structure without a cell wall. [6]
  7. Individual spores then pack together into a seemingly membrane-less spore ball. [11]
  8. Multiple spore balls then join and form a spore cyst. [11]
  9. Cell wall deposition changes spore colour from opaque white to grey to dull black [3] [6]

Physiology

Ascosphaera aggregata has been found to be unable to break down chitin. [6] [3]

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. [12] Recent investigations by James and Skinner (2005) [12] have discovered that PCR of the ITS domain of ribosomal DNA with species specific primer sets allows the detection of fungal DNA (working, even, in asymptomatic individuals). [12] The PCR technique can also be used on hair and honey samples to avoid the difficulty of culturing spores, [12] as spore were shown before to only germinate well in lipids. [13] Storage of the fungus has also proven to be difficult as it collapses after 1–2 months during normal culture passaging. [14] However, Jensen et al. (2009) found that spores could be preserved via cryopreservation or freeze-drying whereas hyphae unfortunately could not be preserved. [14]

Economic importance

Megachile rotundata is the primary pollinator of the commercially grown alfalfa seed, [11] [7] accounting for 46,000 metric tonnes of North American alfafa seed (two-thirds the global production) in 2004. [7] 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. [7] A. aggregata has been killing this economic pollinator in the US since 1972 [10] and has been reported to be able to kill greater than 50% of a population. [8]

Effective management of the fungus has yet to be discovered, as the current registered treatment in Canada (paraformaldehyde fumigation of spores [7] ) involves a carcinogen and other treatment options (heat and chloride treatments) are expensive and labour-intensive. [7]

Related Research Articles

<span class="mw-page-title-main">Ascomycota</span> Division or phylum of fungi

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 defining feature of this fungal group is the "ascus", a microscopic sexual structure in which nonmotile spores, called ascospores, are formed. However, some species of Ascomycota are asexual and thus do not form asci or ascospores. Familiar examples of sac fungi include morels, truffles, brewers' and bakers' yeast, dead man's fingers, and cup fungi. The fungal symbionts in the majority of lichens such as Cladonia belong to the Ascomycota.

<span class="mw-page-title-main">Alfalfa</span> Perennial flowering plant

Alfalfa, also called lucerne, is a perennial flowering plant in the legume family Fabaceae. It is cultivated as an important forage crop in many countries around the world. It is used for grazing, hay, and silage, as well as a green manure and cover crop. The name alfalfa is used in North America. The name lucerne is more commonly used in the United Kingdom, South Africa, Australia, and New Zealand. The plant superficially resembles clover, especially while young, when trifoliate leaves comprising round leaflets predominate. Later in maturity, leaflets are elongated. It has clusters of small purple flowers followed by fruits spiralled in two to three turns containing 10–20 seeds. Alfalfa is native to warmer temperate climates. It has been cultivated as livestock fodder since at least the era of the ancient Greeks and Romans.

<i>Megachile rotundata</i> Species of bee

Megachile rotundata, the alfalfa leafcutting bee, is a European bee that has been introduced to various regions around the world. As a solitary bee species, it does not build colonies or store honey, but is a very efficient pollinator of alfalfa, carrots, other vegetables, and some fruits. Because of this, farmers often use M. rotundata as a pollination aid by distributing M. rotundata prepupae around their crops. Each female constructs and provisions her own nest, which is built in old trees or log tunnels. Being a leafcutter bee, these nests are lined with cut leaves. These bees feed on pollen and nectar and display sexual dimorphism. This species has been known to bite and sting, but it poses no overall danger unless it is threatened or harmed, and its sting has been described as half as painful as a honey bee's.

<span class="mw-page-title-main">Megachilidae</span> Cosmopolitan family of bees

Megachilidae is a cosmopolitan family of mostly solitary bees. Characteristic traits of this family are the restriction of their pollen-carrying structure to the ventral surface of the abdomen, and their typically elongated labrum. Megachilid genera are most commonly known as mason bees and leafcutter bees, reflecting the materials from which they build their nest cells ; a few collect plant or animal hairs and fibers, and are called carder bees, while others use plant resins in nest construction and are correspondingly called resin bees. All species feed on nectar and pollen, but a few are kleptoparasites, feeding on pollen collected by other megachilid bees. Parasitic species do not possess scopae. The motion of Megachilidae in the reproductive structures of flowers is energetic and swimming-like; this agitation releases large amounts of pollen.

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<i>Ophiocordyceps sinensis</i> Species of fungus

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<span class="mw-page-title-main">Bee pollen</span> Ball of pollen gathered by worker honeybees

Bee pollen, also known as bee bread and ambrosia, is a ball or pellet of field-gathered flower pollen packed by worker honeybees, and used as the primary food source for the hive. It consists of simple sugars, protein, minerals and vitamins, fatty acids, and a small percentage of other components. Bee pollen is stored in brood cells, mixed with saliva, and sealed with a drop of honey. Bee pollen is harvested as food for humans and marketed as having various, but yet unproven, health benefits.

<i>Megachile</i> Genus of bees

The genus Megachile is a cosmopolitan group of solitary bees, often called leafcutter bees or leafcutting bees; it also includes the called resin bees and mortar bees. While other genera within the family Megachilidae may chew leaves or petals into fragments to build their nests, certain species within Megachile neatly cut pieces of leaves or petals, hence their common name. This is one of the largest genera of bees, with more than 1500 species in over 50 subgenera. The alfalfa leafcutter bee is managed on a commercial scale for crop pollination, and has been introduced by humans to various regions around the world.

<i>Didymella bryoniae</i> Species of fungus

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<i>Ascosphaera</i> Genus of fungi

Ascosphaera is a genus of fungi in the family Ascosphaeraceae. It was described in 1955 by mycologists Charles F. Spiltoir and Lindsay S. Olive. Members of the genus are insect pathogens. The type species, A. apis, causes chalkbrood disease in honey bees. The reproductive ascospores of the fungus are produced within a unique structure, the spore cyst, or sporocyst.

<span class="mw-page-title-main">California carpenter bee</span> Species of bee

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<span class="mw-page-title-main">Alkali bee</span> Species of insect

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<i>Taphrina caerulescens</i> Species of fungus

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Megachile zombae is a species of bee in the Megachilidae family. Natively endemic to Malawi and identified in 1977, these are solitary bees. The name derives from Greek mega 'large' + cheil- (χειλ) 'lip' and a district Zomba, in Malawi.

References

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  2. 1 2 3 Goulson, Dave (2010). Bumblebees : behaviour, ecology, and conservation (2nd ed.). New York: Oxford University Press. p.  230. ISBN   978-0-19-955306-8.
  3. 1 2 3 4 5 6 7 Capinera, John L. (2008). Encyclopedia of entomology (2nd ed.). Springer. p.  304. ISBN   978-1-4020-6242-1.
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  6. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 McManus, William R.; Youssef, Nabil N. (September 1984). "Life Cycle of the Chalk Brood Fungus, Ascosphaera aggregata, in the Alfalfa Leafcutting Bee, Megachile rotundata, and Its Associated Symptomatology". Mycologia. 76 (5): 830–842. doi:10.2307/3793139. JSTOR   3793139.
  7. 1 2 3 4 5 6 7 8 Pitts-Singer, Theresa L.; Cane, James H. (7 January 2011). "The Alfalfa Leafcutting Bee, Megachile rotundata: The World's Most Intensively Managed Solitary Bee". Annual Review of Entomology. 56 (1): 221–237. doi:10.1146/annurev-ento-120709-144836. PMID   20809804.
  8. 1 2 3 4 5 6 7 Bissett, John (December 1988). "Contribution toward a monograph of the genus". Canadian Journal of Botany. 66 (12): 2541–2560. doi:10.1139/b88-346.
  9. Rank, GH; Goerzen, DW (1981). "Native leafcutter bee species and associated parasites in commercial hives in Saskatchewan, Canada". Apidologie. 12 (3): 211–220. doi: 10.1051/apido:19810301 .
  10. 1 2 Goerzen, D.W.; Erlandson, M.A.; Bissett, J. (31 May 2012). "Occurrence of Chalkbrood Caused by Ascosphaera Aggregata Skou in a Native Leafcutting Bee, Megachile Relativa Cresson (Hymenoptera: Megachilidae), in Saskatchewan". The Canadian Entomologist. 122 (6): 1269–1270. doi:10.4039/Ent1221269-11. S2CID   85760024.
  11. 1 2 3 Richards, K.W. (31 May 2012). "Detection of a Chalkbrood Fungus, Ascosphaera Aggregata, in Larvae of the Alfalfa Leafcutter Bee (Hymenoptera: Megachilidae) from Western Canada". The Canadian Entomologist. 117 (9): 1143–1145. doi:10.4039/Ent1171143-9. S2CID   84474941.
  12. 1 2 3 4 James, R. R.; Skinner, J. S. (1 October 2005). "PCR diagnostic methods for Ascosphaera infections in bees". Journal of Invertebrate Pathology. 90 (2): 98–103. doi:10.1016/j.jip.2005.08.004. ISSN   0022-2011. PMID   16214164.
  13. James, R. R.; Buckner, J. S. (1 October 2004). "Lipids stimulate spore germination in the entomopathogenic ascomycete Ascosphaera aggregata". Mycopathologia. 158 (3): 293–302. doi:10.1007/s11046-004-2910-5. ISSN   1573-0832. PMID   15645171. S2CID   21838968.
  14. 1 2 Jensen, A. B.; James, R. R.; Eilenberg, J. (1 June 2009). "Long-term storage of Ascosphaera aggregata and Ascosphaera apis, pathogens of the leafcutting bee (Megachile rotundata) and the honey bee (Apis mellifera)". Journal of Invertebrate Pathology. 101 (2): 157–160. doi:10.1016/j.jip.2009.03.004. ISSN   0022-2011. PMID   19332075.