Mycangium

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Pronotal mycangia of ambrosia beetle Xylosandrus crassiusculus

The term mycangium (pl., mycangia)(Chinese:贮菌器) is used in biology for special structures on the body of an animal that are adapted for the transport of symbiotic fungi (usually in spore form). This is seen in many xylophagous insects (e.g. horntails and bark beetles), which apparently derive much of their nutrition from the digestion of various fungi that are growing amidst the wood fibers. In some cases, as in ambrosia beetles (Coleoptera: Curculionidae: Scolytinae and Platypodinae) , the fungi are the sole food, and the excavations in the wood are simply to make a suitable microenvironment for the fungus to growow☃☃. In other cases (e.g., the southern pine beetle, Dendroctonus frontalis), wood tissue is the main food, and fungi weaken the defense response ftre hom host plant.[1]

The mites, which have their own type of mycangium (for historical reasons, mite taxonomists use the term sporotheca), ride on or live next to the beetles.[2][3]

Origin[edit]

These structures were first systematically described by Helene Francke-Grosmann at 1956.[4] Then Lekh R. Batra[5] coined the word mycangia:[6] modern Latin, from Greek myco 'fungus' + angeion 'vessel'.

Function[edit]

The most common function of mycangia is preserving and releasing symbiont (symbiotic inoculum, most are fungi, see next section "Mycangia and symbiotic inoculum"). Usually, the symbiotic inoculum in mycangia will provide great help to their vectors (insect or mites carry symbiont). They could help the vectors adapt to the new environment or become nutrients of the vectors themselves and their descendants.[7]

For example ambrosia beetle (vector) Euwallacea fornicatus carry symbiotic fungus (symbiont) Fusarium. When ambrosia beetle bore the host plant, it releases symbiotic fungus from its mycangium. The symbiotic fungus becomes a plant pathogen weaken the resistance of host plant.[8] In the meantime, the fungus grows quickly in the galleries as the main food of beetle.[8] The offsprings of beetle become mature, they will fill their mycangia with symbiont and hunt for the new host plant.[9]

Therefore, mycangia play an important role in protecting the inoculum from degradation and contamination. The structures of mycangia always look like a pouch or a container with caps or small opening that reduce the possibility of contaminants from outside.[4] How mycangia release their inoculum is still unknown now.

Mycangia and symbiotic inoculum[edit]

Most of the inoculum in mycangia are fungi. The symbiotic inoculum of most bark and ambrosia beetles are fungi belonging to Ophiostomatales (Ascomycota: Sordariomycetidae) and Microascales (Ascomycota: Hypocreomycetidae).[7] Symbiotic fungi in mycangia of woodwasps are Amylostereaceae (Basidiomycota: Russulales).[10] Symbiotic fungi in mycangia of lizard beetles are yeast (Ascomycota: Saccharomycetales).[11] Symbiotic fungi in mycangia of ship-timber beetles are Endomyces (Ascomycota: Dipodascaceae).[12] Symbiotic fungi in mycangia of leaf-rolling weevils are Penicillium fungi (Ascomycota: Trichocomaceae).[13] In addition to the above primary symbiotic fungi, secondary fungi and some bacteria have been isolated from mycangia.[14]

Mycangia in insects[edit]

Mycangia in bark and ambrosia beetles[edit]

Oral mycangia in ambrosia beetle Ambrosiodmus

Mycangia of bark and ambrosia beetles (Curculionidae: Scolytinae and Platypodinae) are often complex cuticular invaginations for transport of symbiotic fungi.[2][7] Phloem-feeding bark beetles (Curculionidae: Scolytinae) have usually numerous small pits on the surface of their body, while ambrosia beetles (many Scolytinae and all Platypodinae), which are completely dependent on their fungal symbiont, have deep and complicated pouches.[7] These mycangia are often equipped with glands secreting substances to support fungal spores and perhaps to nourish mycelium during transport.[15] In many cases, the entrance to a mycangium is surrounded by tufts of setae, aiding in scraping mycelium and spores from walls of the tunnels and directing the spores into the mycangium. The mycangia of ambrosia beetle are highly diverse. Different genera or tribes with different kinds of mycangia. Some are oral mycangia in the head,[7] such as genus Ambrosiodmus and Euwallacea.[16] Some are pronotal mycangia , such genus Xylosandrus and Cnestus.[17]

Mycangia in woodwasps (horntails)[edit]

Mycangia of the woodwasps (Hymenoptera: Siricidae) were first described by Buchner.[18] Different from highly diverse types in bark and ambrosia beetles, woodwasps only have a pair of mycangia on the top of their ovipositor. Then when females deposit their eggs inside the host plant, they inject the symbiotic fungi from mycangia and phytotoxic mucus from another reservoir-like structure.[19]

Mycangia in lizard beetles[edit]

One species of lizard beetle Doubledaya bucculenta (Coleoptera: Erotylidae: Erotylidae) has mycangia on the tergum of the eighth abdominal segment. This ovipositor-associated mycangia is only present in adult females. Before Doubledaya bucculentnta deposit their eggs and inject the symbiotic microorganisms on a recently dead bamboo, they will excavate a small hole through the bamboo culm.[11]

Mycangia in ship-timber beetles[edit]

The ship-timber beetle (Coleoptera: Lymexylidae) is another family of wood-boring beetles that live with symbiotic fungi. Buchner first discovered their mycangia located on the ventral side of the long ovipositor.[7] These mycangia form a pair of integumental pouches at either side near the tip of oviduct. When the female lays the eggs, new eggs are coated with the fungal spores.

Mycangia in leaf-rolling weevils[edit]

Females of the leaf-rolling weevil in the genus Euops (Coleoptera: Attelabidae) store symbiotic fungi in the mycangia, which is between the first ventral segment of the abdomen and the thorax.[13] Different from ovipositor-associate mycangia in woodwasps, lizard beetles, and ship-timber beetles, mycangia of leaf-rolling weevils is a pair of spore incubators at the anterior end of the abdomen. This mycangium is formed by the coxa and the metendosternite at the posterior end of the thorax.[10]

References[edit]

  1. ^ Paine, T. D.; Stephen, F. M. (1987-01-01). "Fungi Associated with the Southern Pine Beetle: Avoidance of Induced Defense Response in Loblolly Pine". Oecologia. 74 (3): 377–379. 
  2. ^ a b Francke-Grossmann H. (1967). Ectosymbiosis in wood inhabiting insects. In: M. Henry (ed.) Symbiosis, Vol. 2. Academic Press, NewYork. pp.141-205.
  3. ^ Mori, Boyd A.; Proctor, Heather C.; Walter, David E.; Evenden, Maya L. (2011-02-01). "Phoretic mite associates of mountain pine beetle at the leading edge of an infestation in northwestern Alberta, Canada". The Canadian Entomologist. 143 (1): 44–55. doi:10.4039/n10-043. ISSN 1918-3240. 
  4. ^ a b Francke-Grosmann, H. 1956. Grundlagen der Symbiose bei pilzzüchtenden Holzinsekten. Verhandlungen der Deutschen Zoologischen Gesellschaft 1956: 112–118.
  5. ^ Batra, Lekh (1963). "Ecology of ambrosia fungi and their dissemination by beetles". Transactions of the Kansas Academy of Science. 66: 2 – via JSTOR. 
  6. ^ Batra, L. R. 1963. Ecology of ambrosia fungi and their dissemination by beetles. Trans. Kans. Acad. Sci. 66: 213-236.
  7. ^ a b c d e f Hulcr, Jiri; Stelinski, Lukasz L. (2017-01-31). "The Ambrosia Symbiosis: From Evolutionary Ecology to Practical Management". Annual Review of Entomology. 62 (1): 285–303. doi:10.1146/annurev-ento-031616-035105. 
  8. ^ a b Kasson, Matthew T.; O’Donnell, Kerry; Rooney, Alejandro P.; Sink, Stacy; Ploetz, Randy C.; Ploetz, Jill N.; Konkol, Joshua L.; Carrillo, Daniel; Freeman, Stanley (2013-07-01). "An inordinate fondness for Fusarium: Phylogenetic diversity of fusaria cultivated by ambrosia beetles in the genus Euwallacea on avocado and other plant hosts". Fungal Genetics and Biology. 56: 147–157. doi:10.1016/j.fgb.2013.04.004. 
  9. ^ "tea shot-hole borer, Euwallacea fornicatus". Featured Creatures. 
  10. ^ a b Sakurai, Kazuhiko. "An attelabid weevil (Euops splendida) cultivates fungi". Journal of Ethology. 3 (2): 151–156. doi:10.1007/BF02350306. ISSN 0289-0771. 
  11. ^ a b Toki, Wataru; Tanahashi, Masahiko; Togashi, Katsumi; Fukatsu, Takema (2012-07-27). "Fungal Farming in a Non-Social Beetle". PLOS ONE. 7 (7): e41893. doi:10.1371/journal.pone.0041893. ISSN 1932-6203. PMC 3407107Freely accessible. PMID 22848648. 
  12. ^ Lyngnes, A. R. (1958). "Studier over Hylecoetus dermestoides L. under et angrep på bjorkestokker på Sunnmore 1954-1955". Norsk Entomologisk Tidsskrif. 10: 221–235. 
  13. ^ a b Kobayashi, Chisato; Fukasawa, Yu; Hirose, Dai; Kato, Makoto (2007-08-16). "Contribution of symbiotic mycangial fungi to larval nutrition of a leaf-rolling weevil". Evolutionary Ecology. 22 (6): 711–722. doi:10.1007/s10682-007-9196-2. ISSN 0269-7653. 
  14. ^ Hulcr, J.; Rountree, N. R.; Diamond, S. E.; Stelinski, L. L.; Fierer, N.; Dunn, R. R. (2012-05-01). "Mycangia of Ambrosia Beetles Host Communities of Bacteria". Microbial Ecology. 64 (3): 784–793. doi:10.1007/s00248-012-0055-5. ISSN 0095-3628. 
  15. ^ Six, Diana (2003). "Bark beetle-fungus symbioses". Insect symbiosis: 97–144. 
  16. ^ Li, You; Simmons, David Rabern; Bateman, Craig C.; Short, Dylan P. G.; Kasson, Matthew T.; Rabaglia, Robert J.; Hulcr, Jiri (2015-09-14). "New Fungus-Insect Symbiosis: Culturing, Molecular, and Histological Methods Determine Saprophytic Polyporales Mutualists of Ambrosiodmus Ambrosia Beetles". PLOS ONE. 10 (9): e0137689. doi:10.1371/journal.pone.0137689. ISSN 1932-6203. PMC 4569427Freely accessible. PMID 26367271. 
  17. ^ Stone, W.D.; Nebeker, T.E.; Monroe, W.A.; MacGown, J.A. (2007-02-01). "Ultrastructure of the mesonotal mycangium of Xylosandrus mutilatus (Coleoptera: Curculionidae)". Canadian Journal of Zoology. 85 (2): 232–238. doi:10.1139/z06-205. ISSN 0008-4301. 
  18. ^ Buchner, P. 1928: Holznahrung und Symbiose. Vortrag gehalten auf dem X internationalen Zoologentag zu Budapest am 8. September 1927. Berlin: Springer, pp. 13–16.
  19. ^ Coutts, M. P. (1969). "The mechanism of pathogenicity of Sirex noctilio in Pinus radiata. II. Effects of S. noctilio mucus". Aust. J. Biol. Sci. 22: 1153–1161.