Drosophila

{{Short description|Genus of flies}}

{{About|the entire genus|the most commonly used laboratory species, often just called Drosophila|Drosophila melanogaster|the type-subgenus|Drosophila (subgenus)|a mushroom that once shared this name|Psathyrella candolleana}}

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{{Automatic taxobox

| image = Drosophila_pseudoobscura-Male.png

| image_caption = Drosophila pseudoobscura

| taxon = Drosophila

| authority = Fallén, 1823

| type_species = Musca funebris

| type_species_authority = Fabricius, 1787

| synonyms = Oinopota Kirby & Spence, 1815

| subdivision_ranks = Subgenera

| subdivision = {{ulist|Chusqueophila|Dorsilopha|Drosophila|Dudaica|Phloridosa|Psilodorha|Siphlodora|Sophophora}}

}}

Drosophila ({{IPAc-en|d|r|ə|ˈ|s|ɒ|f|ᵻ|l|ə|,_|d|r|ɒ|-|,_|d|r|oʊ|-}}{{refn|{{cite book | vauthors = Jones D |author-link=Daniel Jones (phonetician) |title=English Pronouncing Dictionary| veditors = Roach P, Hartmann J, Setter J |place=Cambridge|publisher=Cambridge University Press|orig-year=1917|year=2003|isbn=978-3-12-539683-8 }}}}{{refn|{{MerriamWebsterDictionary|Drosophila}}}}), from Ancient Greek δρόσος (drósos), meaning "dew", and φίλος (phílos), meaning "loving", is a genus of fly, belonging to the family Drosophilidae, whose members are often called "small fruit flies" or pomace flies, vinegar flies, or wine flies, a reference to the characteristic of many species to linger around overripe or rotting fruit. They should not be confused with the Tephritidae, a related family, which are also called fruit flies (sometimes referred to as "true fruit flies"); tephritids feed primarily on unripe or ripe fruit, with many species being regarded as destructive agricultural pests, especially the Mediterranean fruit fly.

One species of Drosophila in particular, Drosophila melanogaster, has been heavily used in research in genetics and is a common model organism in developmental biology. The terms "fruit fly" and "Drosophila" are often used synonymously with D. melanogaster in modern biological literature. The entire genus, however, contains more than 1,500 species{{cite web| vauthors = Bächli G |year=1999–2006|url=http://www.taxodros.uzh.ch/|title=TaxoDros: the database on taxonomy of Drosophilidae}} and is very diverse in appearance, behavior, and breeding habitat.

Etymology

The term "Drosophila", meaning "dew-loving", is a modern scientific Latin adaptation from Greek words {{lang|el|δρόσος}}, {{lang|el|drósos}}, "dew", and {{lang|el|φίλος}}, {{lang|el|phílos}}, "loving".

Morphology

Drosophila species are small flies, typically pale yellow to reddish brown to black, with red eyes. When the eyes (essentially a film of lenses) are removed, the brain is revealed. Drosophila brain structure and function develop and age significantly from larval to adult stage. Developing brain structures make these flies a prime candidate for neuro-genetic research.{{cite journal | vauthors = Panikker P, Xu SJ, Zhang H, Sarthi J, Beaver M, Sheth A, Akhter S, Elefant F | display-authors = 6 | title = Restoring Tip60 HAT/HDAC2 Balance in the Neurodegenerative Brain Relieves Epigenetic Transcriptional Repression and Reinstates Cognition | journal = The Journal of Neuroscience | volume = 38 | issue = 19 | pages = 4569–4583 | date = May 2018 | pmid = 29654189 | pmc = 5943982 | doi = 10.1523/JNEUROSCI.2840-17.2018 }} According to a study published in Nature in October 2024, by the scientists examining the brain of an adult female Drosophila, the shape and location of each of its 130,000 neurons and 50 million synapsis were identified. In this study, the most detailed analysis ever conducted on the brain of an adult animal is represented.{{Cite web |date=2024-10-04 |title=Meyve sineğinin beyni, insanlardaki düşünme sürecine ışık tutuyor |url=https://www.bbc.com/turkce/articles/cq5e06gwz2jo |access-date=2024-12-24 |website=BBC News Türkçe |language=tr}}{{Cite web |title=FlyWire |url=https://flywire.ai/ |access-date=2024-12-24 |website=flywire.ai}} Many species, including the noted Hawaiian picture-wings, have distinct black patterns on the wings. The plumose (feathery) arista, bristling of the head and thorax, and wing venation are characters used to diagnose the family. Most are small, about {{Convert|2-4|mm|in}} long, but some, especially many of the Hawaiian species, are larger than a house fly.

Evolution

{{expand section|date=January 2021}}

=Detoxification mechanisms=

Environmental challenge by natural toxins helped to prepare Drosophilae to detox DDT,{{cite journal | vauthors = Low WY, Ng HL, Morton CJ, Parker MW, Batterham P, Robin C | title = Molecular evolution of glutathione S-transferases in the genus Drosophila | journal = Genetics | volume = 177 | issue = 3 | pages = 1363–1375 | date = November 2007 | pmid = 18039872 | pmc = 2147980 | doi = 10.1534/genetics.107.075838 | publisher = Genetics Society of America (OUP) | doi-access = free }}{{rp|Abstract}}{{rp|1365}}{{rp|1369}} by shaping the glutathione S-transferase mechanism{{rp|1365}}{{rp|1369}} that metabolizes both.{{rp|Abstract}}{{cite journal | vauthors = Tang AH, Tu CP | title = Biochemical characterization of Drosophila glutathione S-transferases D1 and D21 | journal = The Journal of Biological Chemistry | volume = 269 | issue = 45 | pages = 27876–27884 | date = November 1994 | pmid = 7961718 | doi = 10.1016/S0021-9258(18)46868-8 | doi-access = free }}

=Selection=

The Drosophila genome is subject to a high degree of selection, especially unusually widespread negative selection compared to other taxa. A majority of the genome is under selection of some sort, and a supermajority of this is occurring in non-coding DNA.{{cite journal | vauthors =Hough J, Williamson RJ, Wright SI | title=Patterns of Selection in Plant Genomes | journal=Annual Review of Ecology, Evolution, and Systematics | publisher=Annual Reviews | volume=44 | issue=1 | date=2013-11-23 | issn=1543-592X | doi=10.1146/annurev-ecolsys-110512-135851 | pages=31–49}}

Effective population size has been credibly suggested to positively correlate with the effect size of both negative and positive selection. Recombination is likely to be a significant source of diversity. There is evidence that crossover is positively correlated with polymorphism in D. populations.

Biology

=Habitat=

Drosophila species are found all around the world, with more species in the tropical regions. Drosophila made their way to the Hawaiian Islands and radiated into over 800 species.{{Cite web | vauthors = Magnacca K | date = 8 October 2015 |url= https://www.fws.gov/endangered/about/ep_45_2015.html | archive-url = https://web.archive.org/web/20181201132456/https://www.fws.gov/endangered/about/ep_45_2015.html | archive-date = 1 December 2018 |title= Endangered Species: Featured Species: Relict Leopard Frog | publisher = U.S. Fish and Wildlife Service | work = Ecological Services Program |language=en-US|access-date=2018-03-10}} They can be found in deserts, tropical rainforest, cities, swamps, and alpine zones. Some northern species hibernate. The northern species D. montana is the best cold-adapted,{{cite journal | vauthors = Parker DJ, Wiberg RA, Trivedi U, Tyukmaeva VI, Gharbi K, Butlin RK, Hoikkala A, Kankare M, Ritchie MG | display-authors = 6 | title = Inter and Intraspecific Genomic Divergence in Drosophila montana Shows Evidence for Cold Adaptation | journal = Genome Biology and Evolution | volume = 10 | issue = 8 | pages = 2086–2101 | date = August 2018 | pmid = 30010752 | pmc = 6107330 | doi = 10.1093/gbe/evy147 }} and is primarily found at high altitudes.{{cite book | vauthors = Routtu J |year=2007 |url=https://rmbl.org/modules/Downloads/Publications/Routtu_phd_2007.pdf |title=Genetic and Phenotypic Divergence in Drosophila virilis and D. montana |location=Jyväskylä |publisher=University of Jyväskylä |page=13 }} Most species breed in various kinds of decaying plant and fungal material, including fruit, bark, slime fluxes, flowers, and mushrooms. Drosophila species that are fruit-breeding are attracted to various products of fermentation, especially ethanol and methanol. Fruits exploited by Drosophila species include those with a high pectin concentration, which is an indicator of how much alcohol will be produced during fermentation. Citrus, morinda, apples, pears, plums, and apricots belong into this category.{{Cite journal |last1=Keesey |first1=Ian W. |last2=Hansson |first2=Bill S. |date=2022-01-07 |title=Neuroecology of Alcohol Preference in Drosophila |journal=Annual Review of Entomology |volume=67 |issue=1 |pages=261–279 |doi=10.1146/annurev-ento-070721-091828 |pmid=34995092 |issn=0066-4170|doi-access=free }}

The larvae of at least one species, D. suzukii, can also feed in fresh fruit and can sometimes be a pest.{{cite web|url=http://cisr.ucr.edu/spotted_wing_drosophila_cherry_vinegar_fly.html| vauthors = Hoddle M |access-date=July 29, 2010|title=Spotted Wing Drosophila (Cherry Vinegar Fly) Drosophila suzukii|publisher=Center for Invasive Species Research}} A few species have switched to being parasites or predators. Many species can be attracted to baits of fermented bananas or mushrooms, but others are not attracted to any kind of baits. Males may congregate at patches of suitable breeding substrate to compete for the females, or form leks, conducting courtship in an area separate from breeding sites.{{citation needed|date=November 2019}}

Several Drosophila species, including Drosophila melanogaster, D. immigrans, and D. simulans, are closely associated with humans, and are often referred to as domestic species. These and other species (D. subobscura, and from a related genus Zaprionus indianus{{cite journal | vauthors = Vilela CR |title=Is Zaprionus indianus Gupta, 1970 (Diptera, Drosophilidae) currently colonizing the Neotropical Region |date=1 January 1999 |journal=Drosophila Information Service |volume=82 |pages=37–39 |url=https://www.scienceopen.com/document?vid=3dbd8296-d04c-41a5-9062-57e688fc25c0 }}{{cite journal | vauthors = van der Linde K, Steck GJ, Hibbard K, Birdsley JS, Alonso LM, Houle D |title=First records of Zaprionus indianus (Diptera, Drosophilidae), a pest species on commercial fruits, from Panama and the United States of America |journal=Florida Entomologist |date=September 2006 |volume=89 |issue=3 |pages=402–404 |doi=10.1653/0015-4040(2006)89[402:FROZID]2.0.CO;2 |issn=0015-4040 |doi-access=free }}{{cite journal | vauthors = Castrezana S |year=2007|title=New records of Zaprionus indianus Gupta, 1970 (Diptera, Drosophilidae) in North America and a key to identify some Zaprionus species deposited in the Drosophila Tucson Stock Center|journal=Drosophila Information Service|volume=90|pages=34–36 |url=http://www.ou.edu/journals/dis/DIS90/Research/Castrezana3.pdf }}) have been accidentally introduced around the world by human activities such as fruit transports.

File:Drosophila residua head.jpg

=Reproduction=

Males of this genus are known to have the longest sperm cells of any studied organism on Earth, including one species, Drosophila bifurca, that has sperm cells that are {{convert|58|mm|abbr=on}} long.{{cite journal | vauthors = Pitnick S, Spicer GS, Markow TA | title = How long is a giant sperm? | journal = Nature | volume = 375 | issue = 6527 | pages = 109 | date = May 1995 | pmid = 7753164 | doi = 10.1038/375109a0 | s2cid = 4368953 | bibcode = 1995Natur.375Q.109P | doi-access = free | author-link3 = Therese Ann Markow }} The cells mostly consist of a long, thread-like tail, and are delivered to the females in tangled coils. The other members of the genus Drosophila also make relatively few giant sperm cells, with that of D. bifurca being the longest.{{cite journal | vauthors = Joly D, Luck N, Dejonghe B | title = Adaptation to long sperm in Drosophila: correlated development of the sperm roller and sperm packaging | journal = Journal of Experimental Zoology. Part B, Molecular and Developmental Evolution | volume = 310 | issue = 2 | pages = 167–178 | date = March 2008 | pmid = 17377954 | doi = 10.1002/jez.b.21167 | bibcode = 2008JEZB..310..167J }} D. melanogaster sperm cells are a more modest 1.8 mm long, although this is still about 35 times longer than a human sperm. Several species in the D. melanogaster species group are known to mate by traumatic insemination.{{cite journal | vauthors = Kamimura Y | title = Twin intromittent organs of Drosophila for traumatic insemination | journal = Biology Letters | volume = 3 | issue = 4 | pages = 401–404 | date = August 2007 | pmid = 17519186 | pmc = 2391172 | doi = 10.1098/rsbl.2007.0192 }}

Drosophila species vary widely in their reproductive capacity. Those such as D. melanogaster that breed in large, relatively rare resources have ovaries that mature 10–20 eggs at a time, so that they can be laid together on one site. Others that breed in more-abundant but less nutritious substrates, such as leaves, may only lay one egg per day. The eggs have one or more respiratory filaments near the anterior end; the tips of these extend above the surface and allow oxygen to reach the embryo. Larvae feed not on the vegetable matter itself, but on the yeasts and microorganisms present on the decaying breeding substrate. Development time varies widely between species (between 7 and more than 60 days) and depends on the environmental factors such as temperature, breeding substrate, and crowding.

Fruit flies lay eggs in response to environmental cycles. Eggs laid at a time (e.g., night) during which likelihood of survival is greater than in eggs laid at other times (e.g., day) yield more larvae than eggs that were laid at those times. Ceteris paribus, the habit of laying eggs at this 'advantageous' time would yield more surviving offspring, and more grandchildren, than the habit of laying eggs during other times. This differential reproductive success would cause D. melanogaster to adapt to environmental cycles, because this behavior has a major reproductive advantage.{{cite journal | vauthors = Howlader G, Sharma VK | title = Circadian regulation of egg-laying behavior in fruit flies Drosophila melanogaster | journal = Journal of Insect Physiology | volume = 52 | issue = 8 | pages = 779–785 | date = August 2006 | pmid = 16781727 | doi = 10.1016/j.jinsphys.2006.05.001 | bibcode = 2006JInsP..52..779H }}

Their median lifespan is 35–45 days.{{cite journal | vauthors = Broughton SJ, Piper MD, Ikeya T, Bass TM, Jacobson J, Driege Y, Martinez P, Hafen E, Withers DJ, Leevers SJ, Partridge L | display-authors = 6 | title = Longer lifespan, altered metabolism, and stress resistance in Drosophila from ablation of cells making insulin-like ligands | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 102 | issue = 8 | pages = 3105–3110 | date = February 2005 | pmid = 15708981 | pmc = 549445 | doi = 10.1073/pnas.0405775102 | doi-access = free | bibcode = 2005PNAS..102.3105B }}

{{multiple image|total_width=600|align=center

| title = Lifecycle of Drosophila

| width1=168|height1=140|image1 = Drosophila egg.png

| caption1 = Egg

| width2=598|height2=337|image2 = Fruit fly larva 01.jpg

| caption2 = Larva

| width3=688|height3=460|image3 = Fruit fly pupae 01.jpg

| caption3 = Pupae (brown specimens are older than the white ones)

| width4=546|height4=424|image4=Drosophila melanogaster - side (aka).jpg

| caption4=Adult D. melanogaster

}}

=Aging=

DNA damage accumulates in Drosophila intestinal stem cells with age.{{cite journal |vauthors=Park JS, Lee SH, Na HJ, Pyo JH, Kim YS, Yoo MA |title=Age- and oxidative stress-induced DNA damage in Drosophila intestinal stem cells as marked by Gamma-H2AX |journal=Exp Gerontol |volume=47 |issue=5 |pages=401–5 |date=May 2012 |pmid=22387531 |doi=10.1016/j.exger.2012.02.007 }} Deficiencies in the Drosophila DNA damage response, including deficiencies in expression of genes involved in DNA damage repair, accelerates intestinal stem cell (enterocyte) aging.{{cite journal |vauthors=Park JS, Jeon HJ, Pyo JH, Kim YS, Yoo MA |title=Deficiency in DNA damage response of enterocytes accelerates intestinal stem cell aging in Drosophila |journal=Aging (Albany NY) |volume=10 |issue=3 |pages=322–338 |date=March 2018 |pmid=29514136 |pmc=5892683 |doi=10.18632/aging.101390 }} Sharpless and Depinho{{cite journal | last1 = Sharpless | first1 = NE | last2 = DePinho | first2 = RA | date = Sep 2007 | title = How stem cells age and why this makes us grow old | journal = Nat Rev Mol Cell Biol | volume = 8 | issue = 9| pages = 703–13 | doi = 10.1038/nrm2241 | pmid = 17717515 | s2cid = 36305591 }} reviewed evidence that stem cells undergo intrinsic aging and speculated that stem cells grow old, in part, as a result of DNA damage.

= Mating systems =

== Courtship behavior ==

The following section is based on the following Drosophila species: Drosophila simulans and Drosophila melanogaster.

Courtship behavior of male Drosophila is an attractive behaviour.{{cite journal | vauthors = Pan Y, Robinett CC, Baker BS | title = Turning males on: activation of male courtship behavior in Drosophila melanogaster | journal = PLOS ONE | volume = 6 | issue = 6 | pages = e21144 | year = 2011 | pmid = 21731661 | pmc = 3120818 | doi = 10.1371/journal.pone.0021144 | doi-access = free | bibcode = 2011PLoSO...621144P }} Females respond via their perception of the behavior portrayed by the male.{{cite journal | vauthors = Cook RM | title = Courtship processing in Drosophila melanogaster. II. An adaptation to selection for receptivity to wingless males | journal = Animal Behaviour | volume = 21 | issue = 2 | pages = 349–358 | date = May 1973 | pmid = 4198506 | doi = 10.1016/S0003-3472(73)80077-6 }} Male and female Drosophila use a variety of sensory cues to initiate and assess courtship readiness of a potential mate.{{cite journal | vauthors = Crossley SA, Bennet-Clark HC, Evert HT |year=1995 |title=Courtship song components affect male and female Drosophila differently |journal=Animal Behaviour |volume=50 |issue=3 |pages=827–839 |doi=10.1016/0003-3472(95)80142-1|s2cid=53161217 }} The cues include the following behaviours: positioning, pheromone secretion, following females, making tapping sounds with legs, singing, wing spreading, creating wing vibrations, genitalia licking, bending the stomach, attempt to copulate, and the copulatory act itself.{{cite journal | vauthors = Ejima A, Griffith LC | title = Measurement of Courtship Behavior in Drosophila melanogaster | journal = Cold Spring Harbor Protocols | volume = 2007 | issue = 10 | pages = pdb.prot4847 | date = October 2007 | pmid = 21356948 | doi = 10.1101/pdb.prot4847 }} The songs of Drosophila melanogaster and Drosophila simulans have been studied extensively. These luring songs are sinusoidal in nature and varies within and between species.

The courtship behavior of Drosophila melanogaster has also been assessed for sex-related genes, which have been implicated in courtship behavior in both the male and female. Recent experiments explore the role of fruitless (fru) and doublesex (dsx), a group of sex-behaviour linked genes.{{cite journal | vauthors = Certel SJ, Savella MG, Schlegel DC, Kravitz EA | title = Modulation of Drosophila male behavioral choice | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 104 | issue = 11 | pages = 4706–4711 | date = March 2007 | pmid = 17360588 | pmc = 1810337 | doi = 10.1073/pnas.0700328104 | doi-access = free | bibcode = 2007PNAS..104.4706C }}

The fruitless (fru) gene in Drosophila helps regulate the network for male courtship behavior; when a mutation to this gene occurs altered same sex sexual behavior in males is observed.{{cite journal | vauthors = Demir E, Dickson BJ | title = fruitless splicing specifies male courtship behavior in Drosophila | journal = Cell | volume = 121 | issue = 5 | pages = 785–794 | date = June 2005 | pmid = 15935764 | doi = 10.1016/j.cell.2005.04.027 | doi-access = free }} Male Drosophila with the fru mutation direct their courtship towards other males as opposed to typical courtship, which would be directed towards females.{{cite journal | vauthors = Yamamoto D, Kohatsu S | title = What does the fruitless gene tell us about nature vs. nurture in the sex life of Drosophila? | journal = Fly | volume = 11 | issue = 2 | pages = 139–147 | date = April 2017 | pmid = 27880074 | pmc = 5406164 | doi = 10.1080/19336934.2016.1263778 }} Loss of the fru mutation leads back to the typical courtship behavior.

==Pheromones==

A novel class of pheromones was found to be conserved across the subgenus Drosophila in 11 desert dwelling species.{{cite journal | vauthors = Chin JS, Ellis SR, Pham HT, Blanksby SJ, Mori K, Koh QL, Etges WJ, Yew JY | display-authors = 6 | title = Sex-specific triacylglycerides are widely conserved in Drosophila and mediate mating behavior | journal = eLife | volume = 3 | issue = | pages = e01751 | date = March 2014 | pmid = 24618898 | pmc = 3948109 | doi = 10.7554/eLife.01751 | doi-access = free }} These pheromones are triacylglycerides that are secreted exclusively by males from their ejaculatory bulb and transferred to females during mating. The function of the pheromones is to make the females unattractive to subsequent suitors and thus inhibit courtship by other males.

==Polyandry==

The following section is based on the following Drosophila species: Drosophila serrata, Drosophila pseudoobscura, Drosophila melanogaster, and Drosophila neotestacea. Polyandry is a prominent mating system among Drosophila.{{cite journal | vauthors = Frentiu FD, Chenoweth SF | title = Polyandry and paternity skew in natural and experimental populations of Drosophila serrata | journal = Molecular Ecology | volume = 17 | issue = 6 | pages = 1589–1596 | date = March 2008 | pmid = 18266626 | doi = 10.1111/j.1365-294X.2008.03693.x | bibcode = 2008MolEc..17.1589F | s2cid = 23286566 }}{{cite journal | vauthors = Puurtinen M, Fromhage L | title = Evolution of male and female choice in polyandrous systems | journal = Proceedings. Biological Sciences | volume = 284 | issue = 1851 | page = 20162174 | date = March 2017 | pmid = 28330914 | pmc = 5378073 | doi = 10.1098/rspb.2016.2174 }}{{cite journal | vauthors = Herrera P, Taylor ML, Skeats A, Price TA, Wedell N | title = Can patterns of chromosome inversions in Drosophila pseudoobscura predict polyandry across a geographical cline? | journal = Ecology and Evolution | volume = 4 | issue = 15 | pages = 3072–3081 | date = August 2014 | pmid = 25247064 | pmc = 4161180 | doi = 10.1002/ece3.1165 | bibcode = 2014EcoEv...4.3072H }}{{cite journal | vauthors = Pinzone CA, Dyer KA | title = Association of polyandry and sex-ratio drive prevalence in natural populations of Drosophila neotestacea | journal = Proceedings. Biological Sciences | volume = 280 | issue = 1769 | pages = 20131397 | date = October 2013 | pmid = 24004936 | pmc = 3768301 | doi = 10.1098/rspb.2013.1397 }} Females mating with multiple sex partners has been a beneficial mating strategy for Drosophila. The benefits include both pre and post copulatory mating. Pre-copulatory strategies are the behaviours associated with mate choice and the genetic contributions, such as production of gametes, that are exhibited by both male and female Drosophila regarding mate choice. Post copulatory strategies include sperm competition, mating frequency, and sex-ratio meiotic drive.

These lists are not inclusive. Polyandry among the Drosophila pseudoobscura in North America vary in their number of mating partners. There is a connection between the number of time females choose to mate and chromosomal variants of the third chromosome. It is believed that the presence of the inverted polymorphism is why re-mating by females occurs. The stability of these polymorphisms may be related to the sex-ratio meiotic drive.

However, for Drosophila subobscura, the main mating system is monandry, not normally seen in Drosophila.{{cite journal | vauthors = Holman L, Freckleton RP, Snook RR | title = What use is an infertile sperm? A comparative study of sperm-heteromorphic Drosophila | journal = Evolution; International Journal of Organic Evolution | volume = 62 | issue = 2 | pages = 374–385 | date = February 2008 | pmid = 18053077 | doi = 10.1111/j.1558-5646.2007.00280.x | s2cid = 12804737 | doi-access = free }}

==Sperm competition==

The following section is based on the following Drosophila species: Drosophila melanogaster, Drosophila simulans, and Drosophila mauritiana. Sperm competition is a process that polyandrous Drosophila females use to increase the fitness of their offspring.{{cite journal | vauthors = Manier MK, Belote JM, Berben KS, Lüpold S, Ala-Honkola O, Collins WF, Pitnick S | title = Rapid diversification of sperm precedence traits and processes among three sibling Drosophila species | journal = Evolution; International Journal of Organic Evolution | volume = 67 | issue = 8 | pages = 2348–2362 | date = August 2013 | pmid = 23888856 | doi = 10.1111/evo.12117 | s2cid = 24845539 | doi-access = free }}{{cite journal | vauthors = Clark AG, Begun DJ, Prout T | title = Female x male interactions in Drosophila sperm competition | journal = Science | volume = 283 | issue = 5399 | pages = 217–220 | date = January 1999 | pmid = 9880253 | doi = 10.1126/science.283.5399.217 | s2cid = 43031475 | jstor = 2897403 }}{{cite journal | vauthors = Mack PD, Hammock BA, Promislow DE | title = Sperm competitive ability and genetic relatedness in Drosophila melanogaster: similarity breeds contempt | journal = Evolution; International Journal of Organic Evolution | volume = 56 | issue = 9 | pages = 1789–1795 | date = September 2002 | pmid = 12389723 | doi = 10.1111/j.0014-3820.2002.tb00192.x | s2cid = 2140754 | doi-access = free }}{{cite journal | vauthors = Manier MK, Lüpold S, Pitnick S, Starmer WT | title = An analytical framework for estimating fertilization bias and the fertilization set from multiple sperm-storage organs | journal = The American Naturalist | volume = 182 | issue = 4 | pages = 552–561 | date = October 2013 | pmid = 24021407 | doi = 10.1086/671782 | bibcode = 2013ANat..182..552M | s2cid = 10040656 | url = https://www.zora.uzh.ch/id/eprint/113525/1/Manier%20etal%20AmNat%202013.pdf | author-link4 = William T. Starmer }}{{cite journal | vauthors = Ala-Honkola O, Manier MK |year=2016 |title=Multiple mechanisms of cryptic female choice act on intraspecific male variation in Drosophila simulans |journal=Behavioral Ecology and Sociobiology |volume=70 |issue=4 |pages=519–532 |doi=10.1007/s00265-016-2069-3 |bibcode=2016BEcoS..70..519A |s2cid=17465840 |url=http://urn.fi/URN:NBN:fi:jyu-201603211902 }} The female Drosophila has two sperm storage organs, the spermathecae and seminal receptacle, that allows her to choose the sperm that will be used to inseminate her eggs. However, some species of Drosophila have evolved to only use one or the other.{{cite journal | vauthors = Pitnick S, Marrow T, Spicer GS | title = Evolution of Multiple Kinds of Female Sperm-Storage Organs in Drosophila | journal = Evolution; International Journal of Organic Evolution | volume = 53 | issue = 6 | pages = 1804–1822 | date = December 1999 | pmid = 28565462 | doi = 10.2307/2640442 | jstor = 2640442 }} Females have little control when it comes to cryptic female choice. Female Drosophila through cryptic choice, one of several post-copulatory mechanisms, which allows for the detection and expelling of sperm that reduces inbreeding possibilities. Manier et al. 2013 has categorized the post copulatory sexual selection of Drosophila melanogaster, Drosophila simulans, and Drosophila mauritiana into the following three stages: insemination, sperm storage, and fertilizable sperm. Among the preceding species there are variations at each stage that play a role in the natural selection process. This sperm competition has been found to be a driving force in the establishment of reproductive isolation during speciation.{{cite journal | vauthors = Lüpold S, Manier MK, Puniamoorthy N, Schoff C, Starmer WT, Luepold SH, Belote JM, Pitnick S | display-authors = 6 | title = How sexual selection can drive the evolution of costly sperm ornamentation | journal = Nature | volume = 533 | issue = 7604 | pages = 535–538 | date = May 2016 | pmid = 27225128 | doi = 10.1038/nature18005 | s2cid = 4407752 | bibcode = 2016Natur.533..535L | url = https://zenodo.org/record/1000843 }}{{cite journal | vauthors = Zajitschek S, Zajitschek F, Josway S, Al Shabeeb R, Weiner H, Manier MK | title = Costs and benefits of giant sperm and sperm storage organs in Drosophila melanogaster | journal = Journal of Evolutionary Biology | volume = 32 | issue = 11 | pages = 1300–1309 | date = November 2019 | pmid = 31465604 | doi = 10.1111/jeb.13529 | s2cid = 191162620 | biorxiv = 10.1101/652248 | doi-access = free }}

== Parthenogenesis and gynogenesis ==

Parthenogenesis does not occur in D. melanogaster, but in the gyn-f9 mutant, gynogenesis occurs at low frequency. The natural populations of D. mangebeirai are entirely female, making it the only obligate parthenogenetic species of Drosophila. Parthenogenesis is facultative in parthenogenetica and mercatorum.{{Cite journal |last=Markow |first=Therese Ann |date=2013-04-01 |title=Parents Without Partners: Drosophila as a Model for Understanding the Mechanisms and Evolution of Parthenogenesis |journal=G3: Genes, Genomes, Genetics |language=en |volume=3 |issue=4 |pages=757–762 |doi=10.1534/g3.112.005421 |issn=2160-1836 |pmc=3618362 |pmid=23550124}}{{Cite journal |last1=Loppin |first1=Benjamin |last2=Dubruille |first2=Raphaëlle |last3=Horard |first3=Béatrice |date=August 2015 |title=The intimate genetics of Drosophila fertilization |journal=Open Biology |language=en |volume=5 |issue=8 |pages=150076 |doi=10.1098/rsob.150076 |issn=2046-2441 |pmc=4554920 |pmid=26246493}}

=Laboratory-cultured animals=

D. melanogaster is a popular experimental animal because it is easily cultured en masse out of the wild, has a short generation time, and mutant animals are readily obtainable. In 1906, Thomas Hunt Morgan began his work on D. melanogaster and reported his first finding of a white eyed mutant in 1910 to the academic community. He was in search of a model organism to study genetic heredity and required a species that could randomly acquire genetic mutation that would visibly manifest as morphological changes in the adult animal. His work on Drosophila earned him the 1933 Nobel Prize in Medicine for identifying chromosomes as the vector of inheritance for genes. This and other Drosophila species are widely used in studies of genetics, embryogenesis, chronobiology, speciation, neurobiology, and other areas.{{citation needed|date=November 2019}}

However, some species of Drosophila are difficult to culture in the laboratory, often because they breed on a single specific host in the wild. For some, it can be done with particular recipes for rearing media, or by introducing chemicals such as sterols that are found in the natural host; for others, it is (so far) impossible. In some cases, the larvae can develop on normal Drosophila lab medium, but the female will not lay eggs; for these it is often simply a matter of putting in a small piece of the natural host to receive the eggs.{{Cite web |title=Drosophila Fallén, 1823 |url=https://www.gbif.org/species/112842957 |access-date=2022-06-24 | work = Global Biodiversity Information Facility (GBIF) |language=en}}

The Drosophila Species Stock Center located at Cornell University in Ithaca, New York, maintains cultures of hundreds of species for researchers.{{cite web |title=The National Drosophila Species Stock Center |url=http://blogs.cornell.edu/drosophila/ |publisher=College of Agriculture and Life Science, Cornell University }}

= Use in genetic research =

Drosophila is considered one of the most valuable genetic model organisms; both adults and embryos are used in experiments.{{cite journal | vauthors = Perez-Perri JI, Noerenberg M, Kamel W, Lenz CE, Mohammed S, Hentze MW, Castello A | title = Global analysis of RNA-binding protein dynamics by comparative and enhanced RNA interactome capture | journal = Nature Protocols | volume = 16 | issue = 1 | pages = 27–60 | date = January 2021 | pmid = 33208978 | pmc = 7116560 | doi = 10.1038/s41596-020-00404-1 | publisher = Nature Portfolio }} {{small|1=(JIPP ORCID: [http://orcid.org/0000-0001-5395-6549 0000-0001-5395-6549])}}. {{small|1=(MN ORCID: [http://orcid.org/0000-0002-4834-8888 0000-0002-4834-8888])}}. {{small|1=(SM ORCID: [http://orcid.org/0000-0003-2640-9560 0000-0003-2640-9560])}}. Drosophila is a prime candidate for genetic research because the relationship between human and fruit fly genes is very close; disease-producing genes in humans can be linked to those in Drosophila.{{Cite web | url=https://www.yourgenome.org/theme/fruit-flies-in-the-laboratory/ |title = Fruit flies in the laboratory | work = yourgenome.org }} The fly has approximately 15,500 genes on its four chromosomes, whereas humans have about 22,000 genes among their 23 chromosomes.{{Cite encyclopedia | url=http://modencode.sciencemag.org/drosophila/introduction | encyclopedia = Model Organism Encyclopedia of DNA Elements (modENCODE) | title = Drosophila as a model organism | access-date=2019-11-19 | archive-date=2020-02-05 | archive-url=https://web.archive.org/web/20200205072908/http://modencode.sciencemag.org/drosophila/introduction | url-status=dead }} The low number of chromosomes make Drosophila easier to study. Genetic traits can be studied through different Drosophila lineages, and the findings can be applied to deduce genetic trends in humans. Research conducted on Drosophila has helped to determine the ground rules for genetic inheritance in many organisms.{{cite journal | vauthors = Jennings BH |title=Drosophila – a versatile model in biology & medicine |journal=Materials Today |date=May 2011 |volume=14 |issue=5 |pages=190–195 |doi=10.1016/S1369-7021(11)70113-4 |doi-access=free }}

Drosophila is a useful in vivo tool to analyze Alzheimer's disease.{{cite journal | vauthors = Prüßing K, Voigt A, Schulz JB | title = Drosophila melanogaster as a model organism for Alzheimer's disease | journal = Molecular Neurodegeneration | volume = 8 | issue = | pages = 35 | date = November 2013 | pmid = 24267573 | pmc = 4222597 | doi = 10.1186/1750-1326-8-35 | doi-access = free }} Rhomboid proteases were first detected in Drosophila but then found to be highly conserved across eukaryotes, mitochondria, and bacteria.{{cite journal | vauthors = Freeman M | title = Rhomboid proteases and their biological functions | journal = Annual Review of Genetics | volume = 42 | issue = 1 | pages = 191–210 | year = 2008 | pmid = 18605900 | doi = 10.1146/annurev.genet.42.110807.091628 | publisher = Annual Reviews }}{{cite journal | vauthors = Freeman M | title = The rhomboid-like superfamily: molecular mechanisms and biological roles | journal = Annual Review of Cell and Developmental Biology | volume = 30 | issue = 1 | pages = 235–254 | date = 2014-10-11 | pmid = 25062361 | doi = 10.1146/annurev-cellbio-100913-012944 | publisher = Annual Reviews | s2cid = 31705365 | url = https://ora.ox.ac.uk/objects/uuid:eb7bd447-12a5-421d-b643-b22a82491652 | doi-access = free }} Melanin's ability to protect DNA against ionizing radiation has been most extensively demonstrated in Drosophila, including in the formative study by Hopwood et al. in 1985.{{cite conference | vauthors = Mosse I, Plotnikova S, Kostrova L, Molophei V, Dubovic B | title=Melanin is Effective Radioprotector against Chronic Irradiation and Low Radiation Doses | website=INIS | veditors = Obelic B, Ranogajev-Komor M, Miljanic S, Krajcar Bronic I | publisher=Croatian Radiation Protection Association | page=35 (of 268) | conference=IRPA Regional Congress on Radiation Protection in Central Europe: Radiation Protection and Health | location=Dubrovnik (Croatia) | date = May 2001 }}

=Microbiome=

Like other animals, Drosophila is associated with various bacteria in its gut. The fly gut microbiota or microbiome seems to have a central influence on Drosophila fitness and life history characteristics. The microbiota in the gut of Drosophila represents an active current research field.

Drosophila species also harbour vertically transmitted endosymbionts, such as Wolbachia and Spiroplasma. These endosymbionts can act as reproductive manipulators, such as cytoplasmic incompatibility induced by Wolbachia or male-killing induced by the D. melanogaster Spiroplasma poulsonii (named MSRO). The male-killing factor of the D. melanogaster MSRO strain was discovered in 2018, solving a decades-old mystery of the cause of male-killing. This represents the first bacterial factor that affects eukaryotic cells in a sex-specific fashion, and is the first mechanism identified for male-killing phenotypes.{{cite journal | vauthors = Harumoto T, Lemaitre B | title = Male-killing toxin in a bacterial symbiont of Drosophila | journal = Nature | volume = 557 | issue = 7704 | pages = 252–255 | date = May 2018 | pmid = 29720654 | pmc = 5969570 | doi = 10.1038/s41586-018-0086-2 | bibcode = 2018Natur.557..252H }}

  • {{lay source |template=cite web | vauthors = Papageorgiou N |title=Mystery solved: The bacterial protein that kills male fruit flies |url=https://actu.epfl.ch/news/mystery-solved-the-bacterial-protein-that-kills--3/ |date=5 July 2018 | work = École polytechnique fédérale de Lausanne (EPFL) | trans-work = Swiss Federal Institute of Technology Lausanne | language = French }} Alternatively, they may protect theirs hosts from infection. Drosophila Wolbachia can reduce viral loads upon infection, and is explored as a mechanism of controlling viral diseases (e.g. Dengue fever) by transferring these Wolbachia to disease-vector mosquitoes.{{cite web |title=Wolbachia |url=http://www.eliminatedengue.com/our-research/wolbachia | work = World Mosquito Program (WMP) | location = Melbourne, Australia | publisher = Monash University }} The S. poulsonii strain of Drosophila neotestacea protects its host from parasitic wasps and nematodes using toxins that preferentially attack the parasites instead of the host.{{cite journal | vauthors = Haselkorn TS, Jaenike J | title = Macroevolutionary persistence of heritable endosymbionts: acquisition, retention and expression of adaptive phenotypes in Spiroplasma | journal = Molecular Ecology | volume = 24 | issue = 14 | pages = 3752–3765 | date = July 2015 | pmid = 26053523 | doi = 10.1111/mec.13261 | bibcode = 2015MolEc..24.3752H | s2cid = 206182327 }}{{cite journal | vauthors = Hamilton PT, Peng F, Boulanger MJ, Perlman SJ | title = A ribosome-inactivating protein in a Drosophila defensive symbiont | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 113 | issue = 2 | pages = 350–355 | date = January 2016 | pmid = 26712000 | pmc = 4720295 | doi = 10.1073/pnas.1518648113 | doi-access = free | bibcode = 2016PNAS..113..350H }}{{cite journal | vauthors = Ballinger MJ, Perlman SJ | title = Generality of toxins in defensive symbiosis: Ribosome-inactivating proteins and defense against parasitic wasps in Drosophila | journal = PLOS Pathogens | volume = 13 | issue = 7 | pages = e1006431 | date = July 2017 | pmid = 28683136 | pmc = 5500355 | doi = 10.1371/journal.ppat.1006431 | doi-access = free }}

Since the Drosophila species is one of the most used model organisms, it was vastly used in genetics. However, the effect abiotic factors,{{cite encyclopedia |title=Abiotic Factors |encyclopedia=Encyclopedia of Entomology |year=2004 |pages=7 |place=Dordrecht |publisher=Kluwer Academic Publishers |doi=10.1007/0-306-48380-7_8 |isbn=0-7923-8670-1 }} such as temperature, has on the microbiome on Drosophila species has recently been of great interest. Certain variations in temperature have an impact on the microbiome. It was observed that higher temperatures (31 °C) lead to an increase of Acetobacter populations in the gut microbiome of Drosophila melanogaster as compared to lower temperatures (13 °C). In low temperatures (13 °C), the flies were more cold resistant and also had the highest concentration of Wolbachia.{{cite journal | vauthors = Moghadam NN, Thorshauge PM, Kristensen TN, de Jonge N, Bahrndorff S, Kjeldal H, Nielsen JL | title = Strong responses of Drosophila melanogaster microbiota to developmental temperature | journal = Fly | volume = 12 | issue = 1 | pages = 1–12 | date = January 2018 | pmid = 29095113 | pmc = 5927714 | doi = 10.1080/19336934.2017.1394558 }}

The microbiome in the gut can also be transplanted among organisms. It was found that Drosophila melanogaster became more cold-tolerant when the gut microbiota from Drosophila melanogaster that were reared at low temperatures. This depicted that the gut microbiome is correlated to physiological processes.{{Cite journal | vauthors = Ferguson LV, Dhakal P, Lebenzon JE, Heinrichs DE, Bucking C, Sinclair BJ |date= October 2018 | veditors = Barribeau S |title=Seasonal shifts in the insect gut microbiome are concurrent with changes in cold tolerance and immunity |journal=Functional Ecology |language=en |volume=32 |issue=10 |pages=2357–2368 |doi=10.1111/1365-2435.13153 |bibcode= 2018FuEco..32.2357F |s2cid= 91035207 |issn=0269-8463|url= https://ir.lib.uwo.ca/cgi/viewcontent.cgi?article=1103&context=biologypub }}

Moreover, the microbiome plays a role in aggression, immunity, egg-laying preferences, locomotion and metabolism. As for aggression, it plays a role to a certain degree during courtship. It was observed that germ-free flies were not as competitive compared to the wild-type males. Microbiome of the Drosophila species is also known to promote aggression by octopamine OA signalling. The microbiome has been shown to impact these fruit flies' social interactions, specifically aggressive behaviour that is seen during courtship and mating.{{cite journal | vauthors = Rohrscheib CE, Bondy E, Josh P, Riegler M, Eyles D, van Swinderen B, Weible MW, Brownlie JC | display-authors = 6 | title = Wolbachia Influences the Production of Octopamine and Affects Drosophila Male Aggression | journal = Applied and Environmental Microbiology | volume = 81 | issue = 14 | pages = 4573–4580 | date = July 2015 | pmid = 25934616 | doi = 10.1128/AEM.00573-15 | pmc = 4551182 | bibcode = 2015ApEnM..81.4573R | veditors = Goodrich-Blair H }}

=Predators=

Drosophila species are prey for many generalist predators, such as robber flies. In Hawaii, the introduction of yellowjackets from mainland United States has led to the decline of many of the larger species. The larvae are preyed on by other fly larvae, staphylinid beetles, and ants.{{Cite web |title=Ecology and Genetics of Social Arthropods |url=https://purcelllab.ucr.edu/blog2.html |access-date=2024-06-23 |website=purcelllab.ucr.edu}}

=Neurochemistry=

Fruit flies use several fast-acting neurotransmitters, similar to those found in humans, which allow neurons to communicate and coordinate behavior. Acetylcholine, glutamate, gamma-aminobutyric acid (GABA), dopamine, serotonin, and histamine are all neurotransmitters that can be found in humans, but Drosophila also have another neurotransmitter, octopamine, the analog of norepinephrine. Acetylcholine is the primary excitatory neurotransmitter and GABA is the primary inhibitory neurotransmitter utilized in the drosophila central nervous system. In Drosophila, the effects of many neurotransmitters can vary depending on the receptors and signaling pathways involved, allowing them to act as excitatory or inhibitory signals under different contexts. This versatility enables complex neural processing and behavioral flexibility.

Glutamate can serve as an excitatory neurotransmitter, specifically at the neuromuscular junction in fruit flies. This differs from vertebrates, where acetylcholine is used at these junctions.

In Drosophila, histamine primarily functions as a neurotransmitter in the visual system. It is released by photoreceptor cells to transmit visual information from the eye to the brain, making it essential for vision.

As with many Eukaryotes, this genus is known to express SNAREs, and as with several others the components of the SNARE complex are known to be somewhat substitutable: Although the loss of SNAP-25 - a component of neuronal SNAREs - is lethal, SNAP-24 can fully replace it. For another example, an R-SNARE not normally found in synapses can substitute for synaptobrevin.{{cite journal | vauthors = Ungar D, Hughson FM | title = SNARE protein structure and function | journal = Annual Review of Cell and Developmental Biology | volume = 19 | issue = 1 | pages = 493–517 | year = 2003 | pmid = 14570579 | doi = 10.1146/annurev.cellbio.19.110701.155609 | publisher = Annual Reviews }}

=Immunity=

The Spätzle protein is a ligand of Toll.{{cite journal | vauthors = Belmonte RL, Corbally MK, Duneau DF, Regan JC | title = Sexual Dimorphisms in Innate Immunity and Responses to Infection in Drosophila melanogaster | journal = Frontiers in Immunology | volume = 10 | pages = 3075 | date = 2020-01-31 | pmid = 32076419 | pmc = 7006818 | doi = 10.3389/fimmu.2019.03075 | publisher = Frontiers Media | doi-access = free }}{{cite journal | vauthors = Cerenius L, Söderhäll K | title = Immune properties of invertebrate phenoloxidases | journal = Developmental and Comparative Immunology | volume = 122 | pages = 104098 | date = September 2021 | pmid = 33857469 | doi = 10.1016/j.dci.2021.104098 | publisher = Elsevier | doi-access = free }} In addition to melanin's more commonly known roles in the endoskeleton and in neurochemistry, melanization is one step in the immune responses to some pathogens. Dudzic et al. 2019 additionally find a large number of shared serine protease messengers between Spätzle/Toll and melanization and a large amount of crosstalk between these pathways.

Systematics

{{Cladogram

|clades={{Clade

| style=font-size:85%;line-height:85%

|2={{Clade

|1= Old World Sophophora

|2={{Clade

|1= New World Sophophora

|2= Lordiphosa

|3= Hirtodrosophila duncani

}}

}}

|1={{Clade

|1={{Clade

|1={{Clade

|1= immigrans-tripunctata radiation

|2={{Clade

|1={{Clade

|1= D. quadrilineata species group |2= Samoaia

}}

|2={{Clade

|1= Zaprionus

|2={{Clade

|1= D. tumiditarsus species group

|2= Liodrosophila

}}

}}

}}

}}

|2={{Clade

|1= Dichaetophora

|2= Hirtodrosophila

|3={{Clade

|1= Mycodrosophila

|2= Paramycodrosophila

}}

}}

|3={{Clade

|1={{Clade

|1= {{nowrap|virilis-repleta radiation (in part)}}

|2={{Clade

|1= subgenus Siphlodora

|2= {{nowrap|virilis-repleta radiation (in part)}}

}}

}}

|2={{Clade

|1= Hawaiian Drosophila

|2= Scaptomyza

}}

|3= D. polychaeta species group

}}

}}

|2= Dorsilopha

}}

}}

}}

File:Drosophila setosimentum.jpg, a species of Hawaiian picture-wing fly]]

The genus Drosophila as currently defined is paraphyletic (see below) and contains 1,450 described species,{{cite book| vauthors = Markow TA, O'Grady PM |year=2005|title=Drosophila: A guide to species identification and use|location=London|publisher=Elsevier|isbn=978-0-12-473052-6}} while the total number of species is estimated at thousands.{{cite book | vauthors = Patterson C |title=Evolution|publisher=Cornell University Press|year=1999|isbn=978-0-8014-8594-7|author-link=Colin Patterson (biologist) }}{{page needed|date=November 2019}} The majority of the species are members of two subgenera: Drosophila (about 1,100 species) and Sophophora (including D. (S.) melanogaster; around 330 species).

The Hawaiian species of Drosophila (estimated to be more than 500, with roughly 380 species described) are sometimes recognized as a separate genus or subgenus, Idiomyia,{{cite book | vauthors = Brake I, Bächli G |year=2008 |title=Drosophilidae (Diptera) |series=World Catalogue of Insects |isbn=978-87-88757-88-0 }}{{page needed|date=November 2019}} but this is not widely accepted. About 250 species are part of the genus Scaptomyza, which arose from the Hawaiian Drosophila and later recolonized continental areas.

Evidence from phylogenetic studies suggests these genera arose from within the genus Drosophila:{{cite journal | vauthors = O'Grady P, Desalle R | title = Out of Hawaii: the origin and biogeography of the genus Scaptomyza (Diptera: Drosophilidae) | journal = Biology Letters | volume = 4 | issue = 2 | pages = 195–199 | date = April 2008 | pmid = 18296276 | pmc = 2429922 | doi = 10.1098/rsbl.2007.0575 }}{{cite journal | vauthors = Remsen J, O'Grady P | title = Phylogeny of Drosophilinae (Diptera: Drosophilidae), with comments on combined analysis and character support | journal = Molecular Phylogenetics and Evolution | volume = 24 | issue = 2 | pages = 249–264 | date = August 2002 | pmid = 12144760 | doi = 10.1016/s1055-7903(02)00226-9 | bibcode = 2002MolPE..24..249R }}

Several of the subgeneric and generic names are based on anagrams of Drosophila, including Dorsilopha, Lordiphosa, Siphlodora, Phloridosa, and Psilodorha.

Genetics

Drosophila species are extensively used as model organisms in genetics (including population genetics), cell biology, biochemistry, and especially developmental biology. Therefore, extensive efforts are made to sequence drosophilid genomes. The genomes of these species have been fully sequenced:{{cite web|url=http://rana.lbl.gov/drosophila/index.html|title=12 Drosophila Genomes Project|publisher=Lawrence Berkeley National Laboratory|access-date=July 29, 2010|archive-url=https://web.archive.org/web/20100527141042/http://rana.lbl.gov/drosophila/index.html|archive-date=May 27, 2010|url-status=dead}}

The data have been used for many purposes, including evolutionary genome comparisons. D. simulans and D. sechellia are sister species, and provide viable offspring when crossed, while D. melanogaster and D. simulans produce infertile hybrid offspring. The Drosophila genome is often compared with the genomes of more distantly related species such as the honeybee Apis mellifera or the mosquito Anopheles gambiae.

The Drosophila modEncode project conducted extensive work to annotate Drosophila genomes, profile transcripts, histone modifications, transcription factors, regulatory networks, and other aspects of Drosophila genetics, and make predictions about gene expression among others.{{Cite journal |last1=The modENCODE Consortium |last2=Roy |first2=Sushmita |last3=Ernst |first3=Jason |last4=Kharchenko |first4=Peter V. |last5=Kheradpour |first5=Pouya |last6=Negre |first6=Nicolas |last7=Eaton |first7=Matthew L. |last8=Landolin |first8=Jane M. |last9=Bristow |first9=Christopher A. |last10=Ma |first10=Lijia |last11=Lin |first11=Michael F. |last12=Washietl |first12=Stefan |last13=Arshinoff |first13=Bradley I. |last14=Ay |first14=Ferhat |last15=Meyer |first15=Patrick E. |date=2010-12-24 |title=Identification of Functional Elements and Regulatory Circuits by Drosophila modENCODE |journal=Science |language=en |volume=330 |issue=6012 |pages=1787–1797 |doi=10.1126/science.1198374 |issn=0036-8075 |pmc=3192495 |pmid=21177974|bibcode=2010Sci...330.1787R }}

FlyBase serves as a centralized database of curated genomic data on Drosophila.{{Cite web |title=FlyBase Homepage |url=https://flybase.org/ |access-date=2024-10-28 |website=flybase.org}}

The {{visible anchor|Drosophila 12 Genomes Consortium}} has presented ten new genomes and combines those with previously released genomes for D. melanogaster and D. pseudoobscura to analyse the evolutionary history and common genomic structure of the genus. This includes the discovery of transposable elements (TEs) and illumination of their evolutionary history.{{cite journal | vauthors = Gilbert C, Peccoud J, Cordaux R | title = Transposable Elements and the Evolution of Insects | journal = Annual Review of Entomology | volume = 66 | issue = 1 | pages = 355–372 | date = January 2021 | pmid = 32931312 | doi = 10.1146/annurev-ento-070720-074650 | publisher = Annual Reviews | s2cid = 221747772 | url = https://hal.archives-ouvertes.fr/hal-03376520/file/Gilbert_et_al_30Jan20_text%2BFig.pdf }} Bartolomé et al. 2009 find at least {{frac|3}} of the TEs in D. melanogaster, D. simulans and D. yakuba have been acquired by horizontal transfer. They find an average rate of 0.035 horizontal transfer events per TE family per million years. Bartolomé also finds horizontal transfer of TEs follows other relatedness metrics, with transfer events between D. melanogaster and D. simulans being twice as common as either of them with D. yakuba.

See also

References

{{Reflist}}