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Drosophila melanogaster

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Drosophila melanogaster
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{{short description|Species of fruit fly}}{{Italic title}}{{Taxobox| name = Drosophila melanogaster | image = Drosophila melanogaster Proboscis.jpg| image_caption = | regnum = Animalia| phylum = Arthropoda| classis = Insecta
Fly>Diptera| familia = Drosophilidae| genus = Drosophila| subgenus = SophophoraDrosophila melanogaster species group>Drosophila melanogaster groupDrosophila melanogaster species subgroup>Drosophila melanogaster subgroup| species_complex = Drosophila melanogaster complex| species = D. melanogaster| binomial = Drosophila melanogasterJohann Wilhelm Meigen>Meigen, 1830MEIGEN JW > TITLE = SYSTEMATISCHE BESCHREIBUNG DER BEKANNTEN EUROPäISCHEN ZWEIFLüGELIGEN INSEKTEN. (VOLUME 6) YEAR = 1830 URL = HTTPS://DLIB.STANFORD.EDU:6521/TEXT1/DD-ILL/INSEKTEN6.PDF ARCHIVE-URL = HTTPS://WWW.WEBCITATION.ORG/5NDBHU1QX?URL=HTTPS://DLIB.STANFORD.EDU:6521/TEXT1/DD-ILL/INSEKTEN6.PDF, 2010-02-01, }}Drosophila melanogaster is a species of fly (the taxonomic order Diptera) in the family Drosophilidae. The species is known generally as the common fruit fly (though inaccuratelyWEB,weblink Vinegar Flies, 19 July 2018, ) or vinegar fly. Starting with Charles W. Woodworth's proposal of the use of this species as a model organism, D. melanogaster continues to be widely used for biological research in genetics, physiology, microbial pathogenesis, and life history evolution. As of 2017, eight Nobel prizes had been awarded for research using Drosophila.WEB,weblink Nobel Prizes, D. melanogaster is typically used in research because it can be readily reared in the laboratory, has only four pairs of chromosomes, breeds quickly, and lays many eggs.ENCYCLOPEDIA, James H., Sang, Eric C. R., Reeve, vanc, Encyclopedia of genetics, Drosophila melanogaster: The Fruit Fly,weblink 2009-07-01, 2001-06-23, Fitzroy Dearborn Publishers, I, USA, 157, 978-1-884964-34-3, Its geographic range includes all continents, including islands.JOURNAL, Markow TA, The secret lives of Drosophila flies, English, eLife, 4, June 2015, 26041333, 4454838, 10.7554/eLife.06793, D. melanogaster is a common pest in homes, restaurants, and other places where food is served.WEB,weblink Vinegar Flies, Drosophila species, Family: Drosophilidae, Department of Entomology, College of Agricultural Sciences, Pennsylvania State University, 2017, 20 July 2017, Flies belonging to the family Tephritidae are also called "fruit flies". This can cause confusion, especially in the Mediterranean, Australia, and South Africa, where the Mediterranean fruit fly Ceratitis capitata is an economic pest.

Physical appearance

(File:Biology Illustration Animals Insects Drosophila melanogaster.svg|thumb|left|Female (left) and male (right) D. melanogaster){{multiple image
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}}Wildtype fruit flies are yellow-brown, with brick-red eyes and transverse black rings across the abdomen. They exhibit sexual dimorphism; females are about {{convert|2.5|mm|abbr=on}} long; males are slightly smaller with darker backs. Males are easily distinguished from females based on colour differences, with a distinct black patch at the abdomen, less noticeable in recently emerged flies, and the sexcombs (a row of dark bristles on the tarsus of the first leg). Furthermore, males have a cluster of spiky hairs (claspers) surrounding the reproducing parts used to attach to the female during mating. Extensive images are found at FlyBase.WEB, FlyBase: A database of Drosophila genes and genomes, Genetics Society of America, 2009, August 11, 2009,weblink yes,weblink" title="web.archive.org/web/20090815020557weblink">weblink August 15, 2009,

Lifecycle and reproduction

(File:Drosophila egg.png|thumb|left|Egg of D. melanogaster)Under optimal growth conditions at {{convert|25|C|F}}, the D. melanogaster lifespan is about 50 days from egg to death.JOURNAL, Linford NJ, Bilgir C, Ro J, Pletcher SD, Measurement of lifespan in Drosophila melanogaster, Journal of Visualized Experiments, 71, January 2013, 23328955, 3582515, 10.3791/50068, The developmental period for D. melanogaster varies with temperature, as with many ectothermic species. The shortest development time (egg to adult), 7 days, is achieved at {{convert|28|C|F}}.BOOK, Ashburner M, Thompson JN, Michael Ashburner, The laboratory culture of Drosophila, The genetics and biology of Drosophila, Ashburner M, Wright TRF, 2A, 1–81, Academic Press, 1978, true, BOOK, Ashburner M, Golic KG, Hawley RS, Michael Ashburner, Drosophila: A Laboratory Handbook., 162–4, 2nd, Cold Spring Harbor Laboratory Press, 2005, 978-0-87969-706-8, Development times increase at higher temperatures (11 days at {{convert|30|C|F|disp=or}}) due to heat stress. Under ideal conditions, the development time at {{convert|25|C|F}} is 8.5 days,Bloomington Drosophila Stock Center at Indiana University: Basic Methods of Culturing Drosophila at {{convert|18|C|F}} it takes 19 days and at {{convert|12|C|F}} it takes over 50 days. Under crowded conditions, development time increases,JOURNAL, Chiang HC, Hodson AC, An analytical study of population growth in Drosophila melanogaster, Ecological Monographs, 1950, 20, 173–206, 1948580, 10.2307/1948580, 3, while the emerging flies are smaller.JOURNAL, Bakker K, An analysis of factors which determine success in competition for food among larvae of Drosophila melanogaster, Archives Neerlandaises de Zoologie, 1961, 14, 200–281, 10.1163/036551661X00061, 2, Females lay some 400 eggs (embryos), about five at a time, into rotting fruit or other suitable material such as decaying mushrooms and sap fluxes. The eggs, which are about 0.5 mm long, hatch after 12–15 hours (at {{convert|25|C|F|disp=or}}). The resulting larvae grow for about 4 days (at 25 Â°C) while molting twice (into second- and third-instar larvae), at about 24 and 48 h after hatching. During this time, they feed on the microorganisms that decompose the fruit, as well as on the sugar of the fruit itself. The mother puts feces on the egg sacs to establish the same microbial composition in the larvae's guts that has worked positively for herself.JOURNAL, Blum JE, Fischer CN, Miles J, Handelsman J, Frequent replenishment sustains the beneficial microbiome of Drosophila melanogaster, mBio, 4, 6, e00860-13, November 2013, 24194543, 3892787, 10.1128/mBio.00860-13, Then the larvae encapsulate in the puparium and undergo a 4-day-long metamorphosis (at 25 Â°C), after which the adults eclose (emerge).The female fruit fly prefers a shorter duration when it comes to sex. Males, though, prefer it to last longer.NEWS, Female Flies Put Up a Fight to Keep Sex Short,weblink August 21, 2009, National Geographic News, August 21, 2009, Maggie, Koerth-Baker, vanc, Males perform a sequence of five behavioral patterns to court females. First, males orient themselves while playing a courtship song by horizontally extending and vibrating their wings. Soon after, the male positions himself at the rear of the female's abdomen in a low posture to tap and lick the female genitalia. Finally, the male curls his abdomen and attempts copulation. Females can reject males by moving away, kicking, and extruding their ovipositor.JOURNAL, Connolly, Kevin, Cook, Robert, vanc, Rejection Responses by Female Drosophila melanogaster: Their Ontogeny, Causality and Effects upon the Behaviour of the Courting Male, Behaviour, 1973, 44, 1/2, 142–166, 4533484, 10.1163/156853973x00364, Copulation lasts around 15–20 minutes,JOURNAL, Houot B, Svetec N, Godoy-Herrera R, Ferveur JF, Effect of laboratory acclimation on the variation of reproduction-related characters in Drosophila melanogaster, The Journal of Experimental Biology, 213, Pt 13, 2322–31, July 2010, 20543131, 10.1242/jeb.041566, during which males transfer a few hundred, very long (1.76 mm) sperm cells in seminal fluid to the female.BOOK, Developmental Biology, Gilbert SF, 2006, 9: Fertilization in Drosophila, Sinauer Associates,weblink 978-0-87893-250-4, 8th, yes,weblink" title="web.archive.org/web/20070207233537weblink">weblink 2007-02-07, Females store the sperm in a tubular receptacle and in two mushroom-shaped spermathecae; sperm from multiple matings compete for fertilization. A last male precedence is believed to exist; the last male to mate with a female sires about 80% of her offspring. This precedence was found to occur through both displacement and incapacitation.JOURNAL, Price CS, Dyer KA, Coyne JA, Sperm competition between Drosophila males involves both displacement and incapacitation, Nature, 400, 6743, 449–52, July 1999, 10440373, 10.1038/22755, 1999Natur.400..449P, The displacement is attributed to sperm handling by the female fly as multiple matings are conducted and is most significant during the first 1–2 days after copulation. Displacement from the seminal receptacle is more significant than displacement from the spermathecae. Incapacitation of first male sperm by second male sperm becomes significant 2–7 days after copulation. The seminal fluid of the second male is believed to be responsible for this incapacitation mechanism (without removal of first male sperm) which takes effect before fertilization occurs. The delay in effectiveness of the incapacitation mechanism is believed to be a protective mechanism that prevents a male fly from incapacitating his own sperm should he mate with the same female fly repetitively. Sensory neurons in the uterus of female D. melanogaster respond to a male protein, sex peptide, which is found in sperm. This protein makes the female reluctant to copulate for about 10 days after insemination. The signal pathway leading to this change in behavior has been determined. The signal is sent to a brain region that is a homolog of the hypothalamus and the hypothalamus then controls sexual behavior and desire. Gonadotropic hormones in Drosophila maintain homeostasis and govern reproductive output via a cyclic interrelationship, not unlike the mammalian estrous cycle.JOURNAL, Meiselman M, Lee SS, Tran RT, Dai H, Ding Y, Rivera-Perez C, Wijesekera TP, Dauwalder B, Noriega FG, Adams ME, Drosophila melanogaster, Proceedings of the National Academy of Sciences of the United States of America, 114, 19, E3849–E3858, May 2017, 28439025, 5441734, 10.1073/pnas.1620760114, Sex Peptide perturbs this homeostasis and dramatically shifts the endocrine state of the female by inciting juvenile hormone synthesis in the corpus allatum.JOURNAL, Moshitzky P, Fleischmann I, Chaimov N, Saudan P, Klauser S, Kubli E, Applebaum SW, Sex-peptide activates juvenile hormone biosynthesis in the Drosophila melanogaster corpus allatum, Archives of Insect Biochemistry and Physiology, 32, 3–4, 363–74, 1996, 8756302, 10.1002/(SICI)1520-6327(1996)32:3/43.0.CO;2-T, D. melanogaster is often used for life extension studies, such as to identify genes purported to increase lifespan when mutated.JOURNAL, Carnes MU, Campbell T, Huang W, Butler DG, Carbone MA, Duncan LH, Harbajan SV, King EM, Peterson KR, Weitzel A, Zhou S, Mackay TF, The Genomic Basis of Postponed Senescence in Drosophila melanogaster, PloS One, 10, 9, e0138569, 2015, 26378456, 4574564, 10.1371/journal.pone.0138569, 2015PLoSO..1038569C,

Females

(File:Fruit_flies.jpg|thumb|left|Females prefer to mate with their own brothers over unrelated males.JOURNAL, Loyau A, Cornuau JH, Clobert J, Danchin E, Incestuous sisters: mate preference for brothers over unrelated males in Drosophila melanogaster, PloS One, 7, 12, e51293, 2012, 23251487, 3519633, 10.1371/journal.pone.0051293, 2012PLoSO...751293L, )Females become receptive to courting males about 8–12 hours after emergence.JOURNAL, Pitnick S, Investment in testes and the cost of making long sperm in Drosophila, American Naturalist, 1996, 148, 57–80, 10.1086/285911, Specific neuron groups in females have been found to affect copulation behavior and mate choice. One such group in the abdominal nerve cord allows the female fly to pause her body movements to copulate.WEB, Fruit fly research may reveal what happens in female brains during courtship, mating,weblink October 5, 2014, Activation of these neurons induces the female to cease movement and orient herself towards the male to allow for mounting. If the group is inactivated, the female remains in motion and does not copulate. Various chemical signals such as male pheromones often are able to activate the group.Also, females exhibit mate choice copying. When virgin females are shown other females copulating with a certain type of male, they tend to copulate more with this type of male afterwards than naive females (which have not observed the copulation of others). This behavior is sensitive to environmental conditions, and females copulate less in bad weather conditions.JOURNAL, Dagaeff AC, Pocheville A, Nöbel S, Loyau A, Isabel G, Danchin E, 2016, Drosophila mate copying correlates with atmospheric pressure in a speed learning situation., Animal Behaviour, 121, 163–174, 10.1016/j.anbehav.2016.08.022,weblink {{clear}}

Males

{{refimprove section|date=October 2015}}D. melanogaster males exhibit a strong reproductive learning curve. That is, with sexual experience, these flies tend to modify their future mating behavior in multiple ways. These changes include increased selectivity for courting only intraspecifically, as well as decreased courtship times.Sexually naïve D. melanogaster males are known to spend significant time courting interspecifically, such as with D. simulans flies. Naïve D. melanogaster will also attempt to court females that are not yet sexually mature, and other males. D. melanogaster males show little to no preference for D. melanogaster females over females of other species or even other male flies. However, after D. simulans or other flies incapable of copulation have rejected the males’ advances, D. melanogaster males are much less likely to spend time courting nonspecifically in the future. This apparent learned behavior modification seems to be evolutionarily significant, as it allows the males to avoid investing energy into futile sexual encounters.JOURNAL, Reuven, Dukas, vanc, Male fruit flies learn to avoid interspecific courtship, Behavioral Ecology, 2004, 15, 4, 695–698, 10.1093/beheco/arh068, In addition, males with previous sexual experience modify their courtship dance when attempting to mate with new females — the experienced males spend less time courting, so have lower mating latencies, meaning that they are able to reproduce more quickly. This decreased mating latency leads to a greater mating efficiency for experienced males over naïve males.JOURNAL, Saleem S, Ruggles PH, Abbott WK, Carney GE, Sexual experience enhances Drosophila melanogaster male mating behavior and success, PloS One, 9, 5, e96639, 2014, 24805129, 4013029, 10.1371/journal.pone.0096639, 2014PLoSO...996639S, This modification also appears to have obvious evolutionary advantages, as increased mating efficiency is extremely important in the eyes of natural selection.

Polygamy

Both male and female D. melanogaster flies act polygamously (having multiple sexual partners at the same time).JOURNAL, Haartman, Lars von, vanc, Successive Polygamy, Behaviour, 1951, 3, 1, 256–273, 10.1163/156853951x00296, In both males and females, polygamy results in a decrease in evening activity compared to virgin flies, more so in males than females. Evening activity consists of those in which the flies participate other than mating and finding partners, such as finding food.JOURNAL, Vartak VR, Varma V, Sharma VK, Effects of polygamy on the activity/rest rhythm of male fruit flies Drosophila melanogaster, Die Naturwissenschaften, 102, 1–2, 1252, February 2015, 25604736, 10.1007/s00114-014-1252-5, 2015SciNa.102....3V, The reproductive success of males and females varies, because a female only needs to mate once to reach maximum fertility. Mating with multiple partners provides no advantage over mating with one partner, so females exhibit no difference in evening activity between polygamous and monogamous individuals. For males, however, mating with multiple partners increases their reproductive success by increasing the genetic diversity of their offspring. This benefit of genetic diversity is an evolutionary advantage because it increases the chance that some of the offspring will have traits that increase their fitness in their environment.The difference in evening activity between polygamous and monogamous male flies can be explained with courtship. For polygamous flies, their reproductive success increases by having offspring with multiple partners, and therefore they spend more time and energy on courting multiple females. On the other hand, monogamous flies only court one female, and expend less energy doing so. While it requires more energy for male flies to court multiple females, the overall reproductive benefits it produces has kept polygamy as the preferred sexual choice.The mechanism that affects courtship behavior in Drosophila is controlled by the oscillator neurons DN1s and LNDs.JOURNAL, Bateman AJ, Intra-sexual selection in Drosophila, Heredity, 2, Pt. 3, 349–68, December 1948, 18103134, 10.1038/hdy.1948.21, Oscillation of the DN1 neurons was found to be effected by sociosexual interactions, and is connected to mating-related decrease of evening activity.

Model organism in genetics

D. melanogaster remains one of the most studied organisms in biological research, particularly in genetics and developmental biology.

History of use in genetic analysis

File:Drosophila Gene Linkage Map.svg|thumb|upright=1.5|Thomas Hunt Morgan's Drosophila melanogaster genetic linkage map: This was the first successful gene mapping work and provides important evidence for the chromosome theory of inheritance. The map shows the relative positions of allelic characteristics on the second Drosophila chromosome. The distance between the genes (map units) are equal to the percentage of crossing-over events that occurs between different alleles.]]D. melanogaster was among the first organisms used for genetic analysis, and today it is one of the most widely used and genetically best-known of all eukaryotic organisms. All organisms use common genetic systems; therefore, comprehending processes such as transcription and replication in fruit flies helps in understanding these processes in other eukaryotes, including humans.BOOK, Pierce, Benjamin A, vanc, Genetics: A Conceptual Approach, 2nd, W. H. Freeman, 2004, 978-0-7167-8881-2, Thomas Hunt Morgan began using fruit flies in experimental studies of heredity at Columbia University in 1910 in a laboratory known as the Fly Room. The Fly Room was cramped with eight desks, each occupied by students and their experiments. They started off experiments using milk bottles to rear the fruit flies and handheld lenses for observing their traits. The lenses were later replaced by microscopes, which enhanced their observations. Morgan and his students eventually elucidated many basic principles of heredity, including sex-linked inheritance, epistasis, multiple alleles, and gene mapping.

Reasons for use in laboratories

(File:EyeColors.jpg|thumb|right|D. melanogaster types (clockwise): Brown eyes with black body, cinnabar eyes, sepia eyes with ebony body, vermilion eyes, white eyes, and wildtype eyes with yellow body)There are many reasons the fruit fly is a popular choice as a model organism:
  • Its care and culture require little equipment, space, and expense even when using large cultures.
  • It can be safely and readily anesthetized (usually with ether, carbon dioxide gas, by cooling, or with products such as FlyNap).
  • Its morphology is easy to identify once anesthetized.
  • It has a short generation time (about 10 days at room temperature), so several generations can be studied within a few weeks.
  • It has a high fecundity (females lay up to 100 eggs per day, and perhaps 2000 in a lifetime).
  • Males and females are readily distinguished, and virgin females are easily isolated, facilitating genetic crossing.
  • The mature larva has giant chromosomes in the salivary glands called polytene chromosomes, "puffs", which indicate regions of transcription, hence gene activity.
  • It has only four pairs of chromosomes – three autosomes, and one pair of sex chromosomes.
  • Males do not show meiotic recombination, facilitating genetic studies.
  • Recessive lethal "balancer chromosomes" carrying visible genetic markers can be used to keep stocks of lethal alleles in a heterozygous state without recombination due to multiple inversions in the balancer.
  • The development of this organism—from fertilized egg to mature adult—is well understood.
  • Genetic transformation techniques have been available since 1987.
  • Its complete genome was sequenced and first published in 2000.JOURNAL, Adams MD, Celniker SE, Holt RA, Evans CA, Gocayne JD, Amanatides PG, etal, The genome sequence of Drosophila melanogaster, Science, 287, 5461, 2185–95, March 2000, 10731132, 10.1126/science.287.5461.2185, 10.1.1.549.8639, 2000Sci...287.2185.,
  • Sexual mosaics can be readily produced, providing an additional tool for studying the development and behavior of these flies.JOURNAL, Hotta Y, Benzer S, Mapping of behaviour in Drosophila mosaics, Nature, 240, 5383, 527–35, December 1972, 4568399, 10.1038/240527a0, 1972Natur.240..527H,

Genetic markers

Genetic markers are commonly used in Drosophila research, for example within balancer chromosomes or P-element inserts, and most phenotypes are easily identifiable either with the naked eye or under a microscope. In the list of a few common markers below, the allele symbol is followed by the name of the gene affected and a description of its phenotype. (Note: Recessive alleles are in lower case, while dominant alleles are capitalised.)
  • Cy1: Curly; the wings curve away from the body, flight may be somewhat impaired
  • e1: Ebony; black body and wings (heterozygotes are also visibly darker than wild type)
  • Sb1: Stubble; bristles are shorter and thicker than wild type
  • w1: White; eyes lack pigmentation and appear white
  • bw: Brown; eye color determined by various pigments combined.
  • y1: Yellow; body pigmentation and wings appear yellow, the fly analog of albinism
Drosophila genes are traditionally named after the phenotype they cause when mutated. For example, the absence of a particular gene in Drosophila will result in a mutant embryo that does not develop a heart. Scientists have thus called this gene tinman, named after the Oz character of the same name.JOURNAL, Azpiazu N, Frasch M, tinman and bagpipe: two homeo box genes that determine cell fates in the dorsal mesoderm of Drosophila, Genes & Development, 7, 7B, 1325–40, July 1993, 8101173, 10.1101/gad.7.7b.1325, Likewise changes in the Shavenbaby gene cause the loss of dorsal cuticular hairs in Drosophila sechellia larvae.JOURNAL, Stern DL, Frankel N, The structure and evolution of cis-regulatory regions: the shavenbaby story, Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences, 368, 1632, 20130028, December 2013, 24218640, 3826501, 10.1098/rstb.2013.0028, This system of nomenclature results in a wider range of gene names than in other organisms.

Classic Genetic Mutations

  • Adh: Alcohol dehydrogenase- Drosophila melanogaster can express the alcohol dehydrogenase (ADH) mutation, thereby preventing the breakdown of toxic levels of alcohols into aldehydes and ketones.JOURNAL, Winberg JO, McKinley-McKee JS, February 1998, Drosophila melanogaster alcohol dehydrogenase: mechanism of aldehyde oxidation and dismutation, The Biochemical Journal, 329 ( Pt 3), Pt 3, 561–70, 1219077, 9445383, While ethanol produced by decaying fruit is a natural food source and location for oviposit for Drosophila at low concentrations (


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