Drosophila Neotestacea

''Drosophila neotestacea'' is a member of the ''testacea'' species group of '' Drosophila''. Testacea species are specialist fruit flies that breed on the fruiting bodies of mushrooms. These flies will choose to breed on psychoactive mushrooms such as the Fly Agaric ''Amanita muscaria''. ''Drosophila neotestacea'' can be found in temperate regions of North America, ranging from the north eastern United States to western Canada.

Immunity

''Drosophila neotestacea'' and other mushroom-breeding Drosophila have been studied extensively for their interactions with '' Howardula'' nematode parasites, particularly '' Howardula aoronymphium''. Unlike related species, ''D. neotestacea'' is sterilized by ''H. aoronymphium'' infection. The genetic basis of this susceptibility is unknown, and is nematode-dependent. For instance, a related ''Howardula'' species from Japan does not sterilize ''D. neotestacea'', even though the European and North American ''Howardula'' species do. Moreover, the related '' Drosophila orientacea'' is resistant to infection by the European ''Howardula'' nematodes, but susceptible to the Japanese ''Howardula'' nematodes. Accordingly, nematode infection strongly suppresses genes involved in egg development. Comparisons between ''D. neotestacea'' and nematode-resistant members of the Testacea species group can help tease apart interactions of fly immunity genetics and nematode parasitism genetics.

Initially discovered in ''D. neotestacea'', mushroom-feeding flies are commonly infected with the trypanosomatid parasite '' Jaenimonas drosophilae''.

The major innate immunity pathways of ''Drosophila'' are found in ''D. neotestacea'', however the antimicrobial peptide Diptericin B has been lost. This loss of Diptericin B is also common to the related ''Drosophila testacea'' and ''Drosophila guttifera'', but not the also-related ''Drosophila innubila''. As such, these loss events appear to have been independent, suggesting that Diptericin B is actively selected against in these species; indeed, ''Diptericin B'' is conserved in all other Drosophila species. It also seems that unrelated Tephritid fruit flies have independently derived a ''Diptericin'' gene strikingly similar to the ''Drosophila'' ''Diptericin B'' gene. Like mushroom-feeding flies, these Tephritids also have a non- frugivorous sub-lineage that has similarly lost the Tephritid Diptericin B gene. These evolutionary patterns in mushroom-breeding ''Drosophila'' and other fruit flies suggests that the immune system's effectors (like antimicrobial peptides) are directly shaped by host ecology.

Symbiosis

''Drosophila neotestacea'' can harbour bacterial symbionts including '' Wolbachia'' and notably '' Spiroplasma poulsonii''. The ''S. poulsonii'' strain of ''D. neotestacea'' has spread westward across North America due to the selective pressure imposed by the sterilizing nematode parasite '' Howardula aoronymphium''. While ''S. poulsonii'' can be found in other ''Drosophila'' species, the ''D. neotestacea'' strain is unique in defending its host against nematode infestation. Like other ''S. poulsonii'' strains, the ''D. neotestacea'' strain also protects its host from parasitic wasp infestation.

The mechanism through which ''S. poulsonii'' protects flies from nematodes and parasitic wasps relies on the presence of toxins called ribosome-inactivating proteins (RIPs), similar to Sarcin or Ricin. These toxins cut a conserved structure in ribosomal RNA, ultimately changing the nucleotide sequence at a specific site. This leaves a signature of RIP attack in nematode and wasp RNA. ''Spiroplasma poulsonii'' likely avoids damaging its host fly by carrying parasite-specific complements of RIP toxins encoded on bacterial plasmids. This allows genes for RIP toxins to readily move between species by horizontal gene transfer, as ''D. neotestacea'' ''Spiroplasma'' RIPs are shared by ''Spiroplasma'' of other mushroom-feeding flies, such as '' Megaselia nigra''.

Selfish genetic elements

The Testacea species group is used in population genetics to study sex-ratio distorting 'selfish' or 'driving' X chromosomes. Selfish X chromosomes bias the offspring of males such that fathers only produce daughters. This increases the spread of the selfish X chromosome, as Y chromosome-bearing sperm are never transmitted. In wild populations, up to 30% of ''D. neotestacea'' individuals can harbour a selfish X chromosome. The spread of the ''D. neotestacea'' selfish X is limited by climatic factors, predicted by the harshness of winter. Thus, its frequency in the wild may be affected by ongoing climate change. The mechanism of X chromosome drive may be related to a duplication of an importin gene, a type of nuclear transport protein.

Often, selfish X chromosomes suppress genetic recombination during meiosis. This process maintains the gene clusters that promote X chromosome drive, but also can lead to an accumulation of deleterious mutations via a process known as Muller's ratchet. The ''D. neotestacea'' selfish X suppresses recombination in lab settings, but occasional recombination occurs in the wild evidenced by recombinant genetic regions in wild-caught flies. Other Testacea species harbour selfish X chromosomes, raising the question of whether X chromosome drive played a role in speciation of the Testacea group. At least one selfish X in Testacea group flies is old enough to have been present in the last-common ancestor of '' Drosophila testacea'' and '' Drosophila orientacea''.

See also

* Drosophila testacea species group
* '' Spiroplasma''
* Meiotic drive

References

{{Taxonbar, from=Q14591958

neotestacea
Insects described in 1992