Editing 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 = [[Carl Fredrik Fallén|Fallén]], 1823
| type_species = ''[[Drosophila funebris|Musca funebris]]''<!-- Yes, this is correct. The original name is used.-->
| type_species_authority = [[Johan Christian Fabricius|Fabricius]], 1787
| synonyms = ''Oinopota'' <small>Kirby & Spence, 1815</small>
| subdivision_ranks = Subgenera
| subdivision = {{ulist|''[[Chusqueophila]]''|''[[Dorsilopha]]''|''[[Drosophila (subgenus)|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 (biology)|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<ref name=taxodros>{{cite web| vauthors = Bächli G |year=1999–2006|url=http://www.taxodros.uzh.ch/|title=TaxoDros: the database on taxonomy of Drosophilidae}}</ref> 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 language|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 [[larva]]l to adult stage. Developing brain structures make these flies a prime candidate for neuro-genetic research.<ref name=pmid29654189>{{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 }}</ref> According to a study published in [[Nature (journal)|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 [[neuron]]s and 50 million [[synapsis]] were identified. In this study, the most detailed analysis ever conducted on the brain of an adult animal is represented.<ref>{{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}}</ref><ref>{{Cite web |title=FlyWire |url=https://flywire.ai/ |access-date=2024-12-24 |website=flywire.ai}}</ref> Many species, including the noted Hawaiian picture-wings, have distinct black patterns on the wings. The plumose (feathery) [[arista (biology)|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 ''Drosophila''e to [[detoxification|detox]] [[DDT]],<ref name = "Low_2007">{{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]] ([[Oxford University Press|OUP]]) | doi-access = free }}</ref>{{rp|Abstract}}<ref name = "Low_2007" />{{rp|1365}}<ref name = "Low_2007" />{{rp|1369}} by shaping the [[glutathione S-transferase|glutathione ''S''-transferase]] mechanism<ref name = "Low_2007" />{{rp|1365}}<ref name = "Low_2007" />{{rp|1369}} that metabolizes both.<ref name = "Low_2007" />{{rp|Abstract}}<ref name="Tang-Tu-1994">{{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 }}</ref>

===Selection===
The ''Drosophila'' genome is subject to a high degree of selection, especially unusually widespread [[negative selection (natural selection)|negative selection]] compared to other [[taxon|taxa]]. A majority of the genome is under selection of some sort, and a supermajority of this is occurring in [[non-coding DNA]].<ref name="Hough-et-al-2013">{{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 (publisher)|Annual Reviews]] | volume=44 | issue=1 | date=2013-11-23 | issn=1543-592X | doi=10.1146/annurev-ecolsys-110512-135851 | pages=31–49}}</ref>

[[Effective population size]] has been credibly suggested to positively correlate with the effect size of both negative and [[positive selection]]. [[Homologous recombination|Recombination]] is likely to be a significant source of [[genetic diversity|diversity]]. There is evidence that [[chromosomal crossover|crossover]] is positively correlated with [[gene polymorphism|polymorphism]] in ''D.'' populations.<ref name="Hough-et-al-2013" />

==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 [[Adaptive radiation|radiated]] into over 800 species.<ref>{{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}}</ref> They can be found in [[deserts]], [[tropical rainforest]], [[cities]], [[swamp]]s, and [[alpine climate|alpine zones]]. Some northern species [[hibernation|hibernate]]. The northern species ''[[Drosophila montana|D. montana]]'' is the best cold-adapted,<ref>{{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 }}</ref> and is primarily found at high altitudes.<ref>{{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 }}</ref> Most species breed in various kinds of decaying plant and [[fungus|fungal]] material, including [[fruit]], [[Bark (botany)|bark]], [[slime flux]]es, [[flower]]s, and [[mushroom]]s. ''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 citrifolia|morinda]], apples, pears, plums, and apricots belong into this category.<ref>{{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 }}</ref>

The larvae of at least one species, ''[[Drosophila suzukii|D. suzukii]]'', can also feed in fresh fruit and can sometimes be a pest.<ref>{{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]]}}</ref> A few species have switched to being [[parasitism|parasites]] or [[predator]]s. Many species can be attracted to baits of fermented [[banana]]s 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 [[Lek (animal behaviour)|leks]], conducting courtship in an area separate from breeding sites.{{citation needed|date=November 2019}}

Several ''Drosophila'' species, including ''Drosophila melanogaster'', ''[[Drosophila immigrans|D. immigrans]]'', and ''[[Drosophila simulans|D. simulans]]'', are closely associated with humans, and are often referred to as [[Domestication|domestic]] species. These and other species (''[[Drosophila subobscura|D. subobscura]]'', and from a related genus ''[[Zaprionus indianus]]''<ref>{{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 }}</ref><ref>{{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 }}</ref><ref>{{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 }}</ref>) have been accidentally introduced around the world by human activities such as fruit transports.
[[File:Drosophila residua head.jpg|thumb|Side view of head showing characteristic bristles above the eye]]

===Reproduction===
Males of this genus are known to have the longest [[spermatozoon|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.<ref name="Pitnick">{{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 }}</ref> 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.<ref name="Joly-2007">{{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 }}</ref> ''D. melanogaster'' sperm cells are a more modest 1.8&nbsp;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]].<ref name="kamimura">{{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 }}</ref>

''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 [[yeast]]s and [[microorganism]]s 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.<ref>{{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 }}</ref>

Their median lifespan is 35–45 days.<ref>{{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 }}</ref>

{{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 (naturally occurring)|DNA damage]] accumulates in ''Drosophila'' intestinal [[stem cell]]s with age.<ref>{{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 }}</ref> Deficiencies in the ''Drosophila'' DNA damage response, including deficiencies in expression of genes involved in [[DNA repair|DNA damage repair]], accelerates intestinal stem cell ([[enterocyte]]) aging.<ref>{{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 }}</ref>  Sharpless and Depinho<ref>{{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 }}</ref> 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.<ref name="Pan">{{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 }}</ref> Females respond via their perception of the behavior portrayed by the male.<ref name="Cook">{{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 }}</ref> Male and female ''Drosophila'' use a variety of sensory cues to initiate and assess courtship readiness of a potential mate.<ref name="Pan" /><ref name="Cook" /><ref name="Crossly">{{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 }}</ref> 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.<ref name="Ejimal and Griffith">{{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 }}</ref><ref name="Pan" /><ref name="Cook" /><ref name="Crossly" /> 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.<ref name="Crossly" />

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.<ref name="Pan" /> Recent experiments explore the role of fruitless (''fru'') and doublesex (''dsx''), a group of sex-behaviour linked genes.<ref name="Certel">{{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 }}</ref><ref name="Pan" />

The [[Fruitless (gene)|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.<ref>{{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 }}</ref> Male ''Drosophila'' with the ''fru'' mutation direct their courtship towards other males as opposed to typical courtship, which would be directed towards females.<ref name=":0">{{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 }}</ref> Loss of the ''fru'' mutation leads back to the typical courtship behavior.<ref name=":0" />

====Pheromones====
A novel class of [[pheromone]]s was found to be conserved across the subgenus ''Drosophila'' in 11 desert dwelling species.<ref name="pmid24618898">{{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 }}</ref>  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 in animals|Polyandry]] is a prominent mating system among ''Drosophila''.<ref name=" Frentiu and Chenoweth">{{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 }}</ref><ref name=" Puurtinen and Fromhage ">{{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 }}</ref><ref name=" Herrera et al. ">{{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 }}</ref><ref name=" Pinzone and Dyer.">{{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 }}</ref> Females mating with multiple sex partners has been a beneficial mating strategy for ''Drosophila''.<ref name=" Frentiu and Chenoweth " /><ref name=" Puurtinen and Fromhage " /><ref name=" Herrera et al." /><ref name=" Pinzone and Dyer." /> 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.<ref name=" Frentiu and Chenoweth " /><ref name=" Puurtinen and Fromhage " /> Post copulatory strategies include sperm competition, mating frequency, and sex-ratio meiotic drive.<ref name=" Frentiu and Chenoweth " /><ref name=" Puurtinen and Fromhage " /><ref name=" Herrera et al." /><ref name=" Pinzone and Dyer." />

These lists are not inclusive. Polyandry among the ''Drosophila pseudoobscura'' in North America vary in their number of mating partners.<ref name=" Herrera et al." />  There is a connection between the number of time females choose to mate and chromosomal variants of the third chromosome.<ref name=" Herrera et al." /> It is believed that the presence of the inverted [[Chromosomal polymorphism|polymorphism]] is why re-mating by females occurs.<ref name=" Herrera et al." /> The stability of these polymorphisms may be related to the sex-ratio meiotic drive.<ref name=" Pinzone and Dyer."/>

However, for ''[[Drosophila subobscura]],'' the main mating system is monandry, not normally seen in ''Drosophila.''<ref>{{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 }}</ref>

====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.<ref name="Manier et al">{{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 }}</ref><ref name="Clark, Begun and Prout">{{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 }}</ref><ref name="Mack et al">{{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 }}</ref><ref name=" Manier et al. 2013">{{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 }}</ref><ref name="Ala-Honkola and Manier">{{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 }}</ref> 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.<ref name="Ala-Honkola and Manier"/> However, some species of ''Drosophila'' have evolved to only use one or the other.<ref>{{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 }}</ref> Females have little control when it comes to [[cryptic female choice]].<ref name=" Manier et al. 2013"/><ref name="Clark, Begun and Prout"/> Female ''Drosophila'' through cryptic choice, one of several post-copulatory mechanisms, which allows for the detection and expelling of sperm that reduces inbreeding possibilities.<ref name="Mack et al"/><ref name="Clark, Begun and Prout"/> 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.<ref name="Manier et al. 2013"/> Among the preceding species there are variations at each stage that play a role in the natural selection process.<ref name="Manier et al. 2013"/> This sperm competition has been found to be a driving force in the establishment of reproductive isolation during speciation.<ref>{{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 }}</ref><ref>{{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 }}</ref>

==== 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''.<ref>{{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}}</ref><ref>{{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}}</ref>

===Laboratory-cultured animals===
''[[Drosophila melanogaster|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 (mutation)|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 [[chromosome]]s as the vector of inheritance for genes. This and other ''Drosophila'' species are widely used in studies of [[genetics]], [[Drosophila embryogenesis|embryogenesis]], [[Drosophila circadian rhythm|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.<ref>{{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}}</ref>

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

=== Use in genetic research ===
''Drosophila'' is considered one of the most valuable genetic model organisms; both adults and embryos are used in experiments.<ref name="Perez-Perri-et-al-2020">{{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])}}.</ref> ''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''.<ref>{{Cite web | url=https://www.yourgenome.org/theme/fruit-flies-in-the-laboratory/ |title = Fruit flies in the laboratory | work = yourgenome.org }}</ref> The fly has approximately 15,500 genes on its four chromosomes, whereas humans have about 22,000 genes among their 23 chromosomes.<ref>{{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 }}</ref> 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.<ref>{{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 }}</ref><ref name=pmid29654189/> 

''Drosophila'' is a useful in vivo tool to analyze Alzheimer's disease.<ref name="Prüßing_2013">{{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 }}</ref> [[Rhomboid protease]]s were first detected in ''Drosophila'' but then found to be [[highly conserved]] across [[eukaryote]]s, [[mitochondrion|mitochondria]], and [[bacteria]].<ref name="Freeman-2008">{{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 (publisher)|Annual Reviews]] }}</ref><ref name="Freeman-2014">{{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 (publisher)|Annual Reviews]] | s2cid = 31705365 | url = https://ora.ox.ac.uk/objects/uuid:eb7bd447-12a5-421d-b643-b22a82491652 | doi-access = free }}</ref> 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.<ref name="Mosse-et-al-2001">{{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 }}</ref>

===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 [[Microbiome in the Drosophila gut|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.<ref>{{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 }}<br />
* {{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 }}</ref> 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.<ref>{{cite web |title=Wolbachia |url=http://www.eliminatedengue.com/our-research/wolbachia | work = World Mosquito Program (WMP) | location =  Melbourne, Australia | publisher = Monash University }}</ref> 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.<ref>{{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 }}</ref><ref>{{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 }}</ref><ref>{{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 }}</ref>

Since the ''Drosophila'' species is one of the most used model organisms, it was vastly used in genetics. However, the effect [[Abiotic component|abiotic factors]],<ref>{{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  }}</ref> 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&nbsp;°C) lead to an increase of ''Acetobacter'' populations in the [[Gut microbiota|gut microbiome]] of ''[[Drosophila melanogaster]]'' as compared to lower temperatures (13&nbsp;°C). In low temperatures (13&nbsp;°C), the flies were more cold resistant and also had the highest concentration of ''Wolbachia.''<ref>{{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 }}</ref>

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.<ref>{{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 }}</ref>

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 display|courtship]] and [[mating]].<ref>{{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 }}</ref>

===Predators===
''Drosophila'' species are prey for many generalist predators, such as [[Asilidae|robber flies]]. In [[Hawaii]], the introduction of [[Vespula|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, [[Staphylinidae|staphylinid]] [[beetle]]s, and [[ant]]s.<ref>{{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}}</ref>

===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 [[SNARE (protein)|SNARE]]s, 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 [[synapse]]s can substitute for [[synaptobrevin]].<ref name="Ungar-Hughson-2003">{{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 (publisher)|Annual Reviews]] }}</ref>

===Immunity===
The [[Spätzle (gene)|Spätzle]] protein is a [[ligand]] of [[Toll signaling|Toll]].<ref name="Belmonte-et-al-2020">{{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 }}</ref><ref name="Cerenius-Soderhall-2021">{{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 }}</ref> 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.<ref name="Belmonte-et-al-2020" /><ref name="Cerenius-Soderhall-2021" /> 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 (biology)|crosstalk]] between these pathways.<ref name="Belmonte-et-al-2020" /><ref name="Cerenius-Soderhall-2021" />

==Systematics==
{{Cladogram
|clades={{Clade
| style=font-size:85%;line-height:85%
|2={{Clade
   |1=&nbsp;Old&nbsp;World&nbsp;''[[Sophophora]]''
   |2={{Clade
      |1=&nbsp;New&nbsp;World&nbsp;''Sophophora''
      |2=&nbsp;''[[Lordiphosa]]''
      |3=&nbsp;''[[Hirtodrosophila duncani]]''
      }}
   }}
|1={{Clade
   |1={{Clade
      |1={{Clade
         |1=&nbsp;[[immigrans-tripunctata radiation|''immigrans-tripunctata''&nbsp;radiation]]
         |2={{Clade
            |1={{Clade
               |1=&nbsp;[[Drosophila quadrilineata species group|''D.&nbsp;quadrilineata''&nbsp;species&nbsp;group]]               |2=&nbsp;''[[Samoaia]]''
               }}
            |2={{Clade
               |1=&nbsp;''[[Zaprionus]]''
               |2={{Clade
                  |1=&nbsp;[[Drosophila tumiditarsus species group|''D.&nbsp;tumiditarsus''&nbsp;species&nbsp;group]]
                  |2=&nbsp;''[[Liodrosophila]]''
                  }}
               }}
            }}
         }}
      |2={{Clade
         |1=&nbsp;''[[Dichaetophora (flies)|Dichaetophora]]''
         |2=&nbsp;''[[Hirtodrosophila]]''
         |3={{Clade
            |1=&nbsp;''[[Mycodrosophila]]''
            |2=&nbsp;''[[Paramycodrosophila]]''
            }}
         }}
      |3={{Clade
         |1={{Clade
            |1=&nbsp;{{nowrap|[[virilis-repleta radiation|''virilis-repleta'' radiation (in part)]]}}
            |2={{Clade
               |1=&nbsp;[[Siphlodora|subgenus&nbsp;''Siphlodora'']]
               |2=&nbsp;{{nowrap|[[virilis-repleta radiation|''virilis-repleta'' radiation (in part)]]}}
               }}
            }}
         |2={{Clade
            |1=&nbsp;[[Hawaiian Drosophila|Hawaiian ''Drosophila'']]
            |2=&nbsp;''[[Scaptomyza]]''
            }}
         |3=&nbsp;[[Drosophila polychaeta species group|''D.&nbsp;polychaeta''&nbsp;species&nbsp;group]]
         }}
      }}
   |2=&nbsp;''[[Dorsilopha]]''
   }}
}}
}}
[[File:Drosophila setosimentum.jpg|thumb|upright|''[[Drosophila setosimentum|D. setosimentum]]'', a species of Hawaiian picture-wing fly]]
The [[genus]] ''Drosophila'' as currently defined is [[paraphyletic]] (see below) and contains 1,450 described species,<ref name=taxodros/><ref>{{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}}</ref> while the total number of species is estimated at thousands.<ref>{{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}}</ref> The majority of the [[species]] are members of two subgenera: ''Drosophila'' (about 1,100 species) and ''[[Sophophora]]'' (including ''[[Drosophila melanogaster|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'',<ref name=taxodros/><ref>{{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}}</ref> 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'':<ref>{{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 }}</ref><ref>{{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 }}</ref>
*''[[Liodrosophila]]'' <small>Duda, 1922</small>
*''[[Mycodrosophila]]'' <small>Oldenburg, 1914</small>
*''[[Samoaia]]'' <small>Malloch, 1934</small>
*''[[Scaptomyza]]'' <small>Hardy, 1849</small>
*''[[Zaprionus]]'' <small>Coquillett, 1901</small>
*''[[Zygothrica]]'' <small>Wiedemann, 1830</small>
*''[[Hirtodrosophila]]'' <small>Duda, 1923</small> (position uncertain)

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:<ref name="LBL">{{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}}</ref>
*''[[Drosophila melanogaster|Drosophila (Sophophora) melanogaster]]''
*''[[Drosophila simulans|Drosophila (Sophophora) simulans]]''
*''[[Drosophila sechellia|Drosophila (Sophophora) sechellia]]''
*''[[Drosophila yakuba|Drosophila (Sophophora) yakuba]]''
*''[[Drosophila erecta|Drosophila (Sophophora) erecta]]''
*''[[Drosophila ananassae|Drosophila (Sophophora) ananassae]]''
*''[[Drosophila pseudoobscura|Drosophila (Sophophora) pseudoobscura]]''
*''[[Drosophila persimilis|Drosophila (Sophophora) persimilis]]''
*''[[Drosophila willistoni|Drosophila (Sophophora) willistoni]]''
*''[[Drosophila mojavensis|Drosophila (Drosophila) mojavensis]]''
*''[[Drosophila virilis|Drosophila (Drosophila) virilis]]''
*''[[Drosophila grimshawi|Drosophila (Drosophila) grimshawi]]''

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 (biology)|hybrid]] offspring. The ''Drosophila'' genome is often compared with the genomes of more distantly related species such as the honeybee ''[[Western 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.<ref>{{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 }}</ref>

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

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 element]]s (TEs) and illumination of their evolutionary history.<ref name="Gilbert-et-al-2021">{{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 (publisher)|Annual Reviews]] | s2cid = 221747772 | url = https://hal.archives-ouvertes.fr/hal-03376520/file/Gilbert_et_al_30Jan20_text%2BFig.pdf }}</ref> 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''.<ref name="Gilbert-et-al-2021" />

== See also ==
*[[Drosophila hybrid sterility]]
*[[Laboratory experiments of speciation]]
*[[List of Drosophila species|List of ''Drosophila'' species]]
*''Caenorhabditis'' [[Drosophilae supergroup|'Drosophilae' species supergroup]], a group of species generally found on rotten fruits and transported by ''Drosophila'' flies

== References ==
{{Reflist}}

== External links ==
{{Commons category|Drosophila}}
{{Wikispecies}}
* {{cite web | url = http://flybase.bio.indiana.edu/ | publisher = Indiana University | title = Fly Base | archive-url = https://web.archive.org/web/20090815020557/http://flybase.bio.indiana.edu/ | archive-date = 2009-08-15 }} FlyBase is a comprehensive database for information on the genetics and molecular biology of ''Drosophila''. It includes data from the ''Drosophila'' Genome Projects and data curated from the literature.
* {{cite web | url = http://www.flymine.org | title = FlyMine | work = Department of Genetics | publisher = University of Cambridge, United Kingdom| archive-url = https://web.archive.org/web/20191004144454/http://www.flymine.org/ | archive-date=2019-10-04 }} is an integrated database of genomic, expression and protein data for ''Drosophila''
* University of California, Santa Cruz
** {{cite web | url = http://genome.ucsc.edu/cgi-bin/hgTracks?db=dm6 | title = D. melanogaster | work = UCSC Genome browser | publisher = University of California, Santa Cruz (UCSC) }}
** {{cite web | url = http://stockcenter.ucsd.edu/ | archive-url = https://archive.today/20121212132427/http://stockcenter.ucsd.edu/ | archive-date = 12 December 2012 | publisher = University of California, Santa Cruz (UCSC) | title = ''Drosophila'' Stock Center }} breeds hundreds of species and supplies them to researchers
* Lawrence Berkeley National Laboratory
** {{cite web | url = http://www.fruitfly.org/ | title = Berkeley ''Drosophila'' Genome Project (BDGP) | publisher = Lawrence Berkeley National Laboratory (LBNL) | location = Berkeley, CA }}
** {{cite web | url = http://rana.lbl.gov/drosophila/ | publisher = Lawrence Berkeley National Laboratory (LBNL) | location = Berkeley, CA | archive-url = https://web.archive.org/web/20051214205852/http://rana.lbl.gov/drosophila/ | archive-date = 14 December 2005 | title = Assembly, Alignment and Annotation of 12 ''Drosophila'' species }}
* {{cite web | vauthors = Anderson N | date = 27 September 2022  | url = http://www.thebugsquad.com/fruit-flies/drosophila-melanogaster | title = ''Drosophila Melanogaster'' | work = The Bug Squad }}
* {{cite web | vauthors = Bächli G | url = http://taxodros.uzh.ch/ | title = TaxoDros: The database on Taxonomy of Drosophilidae | publisher = University of Zürich }}
* {{cite web | vauthors = Manning G | date = 5 May 2010 | url = http://ceolas.org/fly/index.html | title = The ''Drosophila'' Virtual library }} is library of ''Drosophila'' on the web
* {{cite web | url = https://genetics-gsa.org/drosophila/ | title = Annual ''Drosophila'' Research Conference | work = Genetics Society of America (GSA) }}
* {{cite web | url = http://ccamp.res.in/fly-facility.html | title = Fly (Drosophila) Facility  | work = Centre for Cellular and Molecular Platforms Fly facility (C-CAMP) | location = Bangalore, India | archive-url = https://web.archive.org/web/20130121085710/http://ccamp.res.in/fly-facility.html | archive-date = 2013-01-21 }} – In India microinjection service for the generation of transgenic lines, Screening Platforms, ''Drosophila'' strain development

{{Model Organisms}}
{{Portal bar|Biology|Insects}}
{{Taxonbar|from=Q312154}}
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[[Category:Drosophila| ]]
[[Category:Drosophilidae genera]]
[[Category:Taxa named by Carl Fredrik Fallén]]
[[Category:Animal models]]