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Drosophila

From Wikipedia, the free encyclopedia
This article is about the entire genus. For the most commonly used laboratory species, often just calledDrosophila,seeDrosophila melanogaster.For the type-subgenus, seeDrosophila (subgenus).For a mushroom that once shared this name, seePsathyrella candolleana.

Drosophila
Drosophila pseudoobscura
Scientific classificationEdit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Diptera
Family: Drosophilidae
Subfamily: Drosophilinae
Genus: Drosophila
Fallén,1823
Type species
Musca funebris
Subgenera
Synonyms

OinopotaKirby & Spence, 1815

Drosophila(/drəˈsɒfɪlə,drɒ-,droʊ-/[1][2]) is agenusoffly,belonging to thefamilyDrosophilidae,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 theTephritidae,a related family, which are also called fruit flies (sometimes referred to as "true fruit flies" ); tephritids feed primarily on unripe or ripefruit,with many species being regarded as destructive agricultural pests, especially theMediterranean fruit fly.

One species ofDrosophilain particular,Drosophila melanogaster,has been heavily used in research ingeneticsand is a commonmodel organismindevelopmental biology.The terms "fruit fly" and "Drosophila"are often used synonymously withD. melanogasterin modern biological literature. The entire genus, however, contains more than 1,500 species[3]and is very diverse in appearance, behavior, and breeding habitat.

Etymology

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The term "Drosophila",meaning" dew-loving ", is a modern scientificLatinadaptation fromGreekwordsδρόσος,drósos,"dew",andφιλία,philía,"lover".

Morphology

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Drosophilaspecies 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.Drosophilabrain structure and function develop and age significantly fromlarvalto adult stage. Developing brain structures make these flies a prime candidate for neuro-genetic research.[4]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 2–4 millimetres (0.079–0.157 in) long, but some, especially many of the Hawaiian species, are larger than ahouse fly.

Evolution

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Detoxification mechanisms

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Environmental challenge by natural toxins helped to prepareDrosophilae todetoxDDT,[5]: Abstract [5]: 1365 [5]: 1369 by shaping theglutathioneS-transferasemechanism[5]: 1365 [5]: 1369 that metabolizes both.[5]: Abstract [6]

Selection

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TheDrosophilagenome is subject to a high degree of selection, especially unusually widespreadnegative selectioncompared to othertaxa.A majority of the genome is under selection of some sort, and a supermajority of this is occurring innon-coding DNA.[7]

Effective population sizehas been credibly suggested to positively correlate with the effect size of both negative andpositive selection.Recombinationis likely to be a significant source ofdiversity.There is evidence thatcrossoveris positively correlated withpolymorphisminD.populations.[7]

Biology

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Habitat

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Drosophilaspecies are found all around the world, with more species in the tropical regions.Drosophilamade their way to the Hawaiian Islands andradiatedinto over 800 species.[8]They can be found indeserts,tropical rainforest,cities,swamps,andalpine zones.Some northern specieshibernate.The northern speciesD. montanais the best cold-adapted,[9]and is primarily found at high altitudes.[10]Most species breed in various kinds of decaying plant andfungalmaterial, includingfruit,bark,slime fluxes,flowers,andmushrooms.Drosophilaspecies that are fruit-breeding are attracted to various products of fermentation, especiallyethanolandmethanol.Fruits exploited byDrosophilaspecies 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.[11]

The larvae of at least one species,D. suzukii,can also feed in fresh fruit and can sometimes be a pest.[12]A few species have switched to beingparasitesorpredators.Many species can be attracted to baits of fermentedbananasor 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 formleks,conducting courtship in an area separate from breeding sites.[citation needed]

SeveralDrosophilaspecies, includingDrosophila melanogaster,D. immigrans,andD. simulans,are closely associated with humans, and are often referred to asdomesticspecies. These and other species (D. subobscura,and from a related genusZaprionus indianus[13][14][15]) have been accidentally introduced around the world by human activities such as fruit transports.

Side view of head showing characteristic bristles above the eye

Reproduction

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Males of this genus are known to have the longestsperm cellsof any studied organism on Earth, including one species,Drosophila bifurca,that has sperm cells that are 58 mm (2.3 in) long.[16]The cells mostly consist of a long, thread-like tail, and are delivered to the females in tangled coils. The other members of the genusDrosophilaalso make relatively few giant sperm cells, with that ofD. bifurcabeing the longest.[17]D. melanogastersperm cells are a more modest 1.8 mm long, although this is still about 35 times longer than a human sperm. Several species in theD. melanogasterspecies group are known to mate bytraumatic insemination.[18]

Drosophilaspecies vary widely in their reproductive capacity. Those such asD. melanogasterthat breed in large, relatively rare resources haveovariesthat 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 theyeastsandmicroorganismspresent 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 astemperature,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 causeD. melanogasterto adapt to environmental cycles, because this behavior has a major reproductive advantage.[19]

Their median lifespan is 35–45 days.[20]

Lifecycle ofDrosophila
Egg
Larva
Pupae (brown specimens are older than the white ones)
AdultD. melanogaster

Aging

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DNA damageaccumulates inDrosophilaintestinalstem cellswith age.[21]Deficiencies in theDrosophilaDNA damage response, including deficiencies in expression of genes involved inDNA damage repair,accelerates intestinal stem cell (enterocyte) aging.[22]Sharpless and Depinho[23]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

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Courtship behavior

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The following section is based on the followingDrosophilaspecies:Drosophila simulansandDrosophila melanogaster.

Courtship behavior of maleDrosophilais an attractive behaviour.[24]Females respond via their perception of the behavior portrayed by the male.[25]Male and femaleDrosophilause a variety of sensory cues to initiate and assess courtship readiness of a potential mate.[24][25][26]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.[27][24][25][26]The songs ofDrosophila melanogasterandDrosophila simulanshave been studied extensively. These luring songs are sinusoidal in nature and varies within and between species.[26]

The courtship behavior ofDrosophila melanogasterhas also been assessed for sex-related genes, which have been implicated in courtship behavior in both the male and female.[24]Recent experiments explore the role of fruitless (fru) and doublesex (dsx), a group of sex-behaviour linked genes.[28][24]

Thefruitless(fru) gene inDrosophilahelps regulate the network for male courtship behavior; when a mutation to this gene occurs altered same sex sexual behavior in males is observed.[29]MaleDrosophilawith thefrumutation direct their courtship towards other males as opposed to typical courtship, which would be directed towards females.[30]Loss of thefrumutation leads back to the typical courtship behavior.[30]

Pheromones

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A novel class ofpheromoneswas found to be conserved across the subgenusDrosophilain 11 desert dwelling species.[31]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

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The following section is based on the followingDrosophilaspecies:Drosophila serrata,Drosophila pseudoobscura,Drosophila melanogaster,andDrosophila neotestacea.Polyandryis a prominent mating system amongDrosophila.[32][33][34][35]Females mating with multiple sex partners has been a beneficial mating strategy forDrosophila.[32][33][34][35]The benefits include both pre and post copulatory mating. Pre-copulatory strategies are the behaviours associated withmate choiceand the genetic contributions, such as production of gametes, that are exhibited by both male and femaleDrosophilaregarding mate choice.[32][33]Post copulatory strategies include sperm competition, mating frequency, and sex-ratio meiotic drive.[32][33][34][35]

These lists are not inclusive. Polyandry among theDrosophila pseudoobscurain North America vary in their number of mating partners.[34]There is a connection between the number of time females choose to mate and chromosomal variants of the third chromosome.[34]It is believed that the presence of the invertedpolymorphismis why re-mating by females occurs.[34]The stability of these polymorphisms may be related to the sex-ratio meiotic drive.[35]

However, forDrosophila subobscura,the main mating system is monandry, not normally seen inDrosophila.[36]

Sperm competition

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The following section is based on the followingDrosophilaspecies:Drosophila melanogaster,Drosophila simulans,andDrosophila mauritiana.Sperm competitionis a process that polyandrousDrosophilafemales use to increase the fitness of their offspring.[37][38][39][40][41]The femaleDrosophilahas two sperm storage organs, the spermathecae and seminal receptacle, that allows her to choose the sperm that will be used to inseminate her eggs.[41]However, some species ofDrosophilahave evolved to only use one or the other.[42]Females have little control when it comes tocryptic female choice.[40][38]FemaleDrosophilathrough cryptic choice, one of several post-copulatory mechanisms, which allows for the detection and expelling of sperm that reduces inbreeding possibilities.[39][38]Manier et al. 2013 has categorized the post copulatory sexual selection ofDrosophila melanogaster,Drosophila simulans,andDrosophila mauritianainto the following three stages: insemination, sperm storage, and fertilizable sperm.[40]Among the preceding species there are variations at each stage that play a role in the natural selection process.[40]This sperm competition has been found to be a driving force in the establishment of reproductive isolation during speciation.[43][44]

Parthenogenesis and gynogenesis

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Parthenogenesisdoes not occur inD. melanogaster,but in thegyn-f9mutant,gynogenesisoccurs at low frequency. The natural populations ofD. mangebeiraiare entirely female, making it the only obligate parthenogenetic species of Drosophila. Parthenogenesis is facultative inparthenogeneticaandmercatorum.[45][46]

Laboratory-cultured animals

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D. melanogasteris 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 Morganbegan his work onD. melanogasterand reported his first finding of awhite eyed mutantin 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 onDrosophilaearned him the 1933Nobel Prizein Medicine for identifyingchromosomesas the vector of inheritance for genes. This and otherDrosophilaspecies are widely used in studies ofgenetics,embryogenesis,chronobiology,speciation,neurobiology,and other areas.[citation needed]

However, some species ofDrosophilaare 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 assterolsthat are found in the natural host; for others, it is (so far) impossible. In some cases, the larvae can develop on normalDrosophilalab 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.[47]

TheDrosophila Species Stock Centerlocated atCornell UniversityinIthaca,New York, maintains cultures of hundreds of species for researchers.[48]

Use in genetic research

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Drosophilais considered one of the most valuable genetic model organisms; both adults and embryos are experimental models.[49]Drosophilais a prime candidate for genetic research because the relationship between human and fruit fly genes is very close.[50]Human and fruit fly genes are so similar, that disease-producing genes in humans can be linked to those in flies. The fly has approximately 15,500 genes on its four chromosomes, whereas humans have about 22,000 genes among their 23 chromosomes. Thus the density of genes per chromosome inDrosophilais higher than the human genome.[51]Low and manageable number of chromosomes makeDrosophilaspecies easier to study. These flies also carry genetic information and pass down traits throughout generations, much like their human counterparts.[clarification needed]The traits can then be studied through differentDrosophilalineages and the findings can be applied to deduce genetic trends in humans. Research conducted onDrosophilahelp determine the ground rules for transmission of genes in many organisms.[52][4]Drosophilais a useful in vivo tool to analyze Alzheimer's disease.[53]Rhomboid proteaseswere first detected inDrosophilabut then found to behighly conservedacrosseukaryotes,mitochondria,andbacteria.[54][55]Melanin's ability to protect DNA againstionizing radiationhas been most extensively demonstrated inDrosophila,including in the formative study by Hopwood et al. 1985.[56]

Microbiome

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Like other animals,Drosophilais associated with various bacteria in its gut. The fly gut microbiota or microbiome seems to have a central influence onDrosophilafitness and life history characteristics. Themicrobiota in the gut ofDrosophilarepresents an active current research field.

Drosophilaspecies also harbour vertically transmitted endosymbionts, such asWolbachiaandSpiroplasma.These endosymbionts can act as reproductive manipulators, such ascytoplasmic incompatibilityinduced byWolbachiaor male-killing induced by theD. melanogaster Spiroplasma poulsonii(named MSRO). The male-killing factor of theD. melanogasterMSRO 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.[57]Alternatively, they may protect theirs hosts from infection.Drosophila Wolbachiacan reduce viral loads upon infection, and is explored as a mechanism of controlling viral diseases (e.g.Dengue fever) by transferring theseWolbachiato disease-vector mosquitoes.[58]TheS. poulsoniistrain ofDrosophila neotestaceaprotects its host from parasitic wasps and nematodes using toxins that preferentially attack the parasites instead of the host.[59][60][61]

Since theDrosophilaspecies is one of the most used model organisms, it was vastly used in genetics. However, the effectabiotic factors,[62]such as temperature, has on themicrobiomeon 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 ofAcetobacterpopulations in thegut microbiomeofDrosophila melanogasteras compared to lower temperatures (13 °C). In low temperatures (13 °C), the flies were more cold resistant and also had the highest concentration ofWolbachia.[63]

The microbiome in the gut can also be transplanted among organisms. It was found thatDrosophila melanogasterbecame more cold-tolerant when the gut microbiota fromDrosophila melanogasterthat were reared at low temperatures. This depicted that the gut microbiome is correlated to physiological processes.[64]

Moreover, the microbiome plays a role in aggression, immunity, egg-laying preferences, locomotion andmetabolism.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 theDrosophilaspecies 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 duringcourtshipandmating.[65]

Predators

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Drosophilaspecies are prey for many generalist predators, such asrobber flies.InHawaii,the introduction ofyellowjacketsfrom mainland United States has led to the decline of many of the larger species. The larvae are preyed on by other fly larvae,staphylinidbeetles,andants.[66]

Neurochemistry

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As with many Eukaryotes, this genus is known to expressSNAREs,and as with several others the components of the SNARE complex are known to be somewhat substitutable: Although the loss ofSNAP-25- a component of neuronal SNAREs - is lethal,SNAP-24can fully replace it. For another example, anR-SNAREnot normally found insynapsescan substitute forsynaptobrevin.[67]

Immunity

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TheSpätzleprotein is aligandofToll.[68][69]In addition tomelanin's more commonly known roles in theendoskeletonand inneurochemistry,melanization is one step in the immune responses to some pathogens.[68][69]Dudzic et al. 2019 additionally find a large number of sharedserine proteasemessengers between Spätzle/Toll and melanization and a large amount ofcrosstalkbetween these pathways.[68][69]

Systematics

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Old WorldSophophora

New WorldSophophora

Lordiphosa

Hirtodrosophila duncani

D. setosimentum,a species of Hawaiian picture-wing fly

ThegenusDrosophilaas currently defined isparaphyletic(see below) and contains 1,450 described species,[3][70]while the total number of species is estimated at thousands.[71]The majority of thespeciesare members of two subgenera:Drosophila(about 1,100 species) andSophophora(includingD. (S.) melanogaster;around 330 species).

The Hawaiian species ofDrosophila(estimated to be more than 500, with roughly 380 species described) are sometimes recognized as a separate genus or subgenus,Idiomyia,[3][72]but this is not widely accepted. About 250 species are part of the genusScaptomyza,which arose from the HawaiianDrosophilaand later recolonized continental areas.

Evidence fromphylogeneticstudies suggests these genera arose from within the genusDrosophila:[73][74]

Several of the subgeneric and generic names are based on anagrams ofDrosophila,includingDorsilopha,Lordiphosa,Siphlodora,Phloridosa,andPsilodorha.

Genetics

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Drosophilaspecies 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 drosphilid genomes. The genomes of these species have been fully sequenced:[75]

The data have been used for many purposes, including evolutionary genome comparisons.D. simulansandD. sechelliaare sister species, and provide viable offspring when crossed, whileD. melanogasterandD. simulansproduce infertilehybridoffspring. TheDrosophilagenome is often compared with the genomes of more distantly related species such as the honeybeeApis melliferaor the mosquitoAnopheles gambiae.

The modEncode consortium is currently sequencing eight moreDrosophilagenomes,[76]and even more genomes are being sequenced by thei5Kconsortium.[77]

Curated data are available atFlyBase.

TheDrosophila 12 Genomes Consortium– led byAndrew G. Clark,Michael Eisen,Douglas Smith, Casey Bergman, Brian Oliver,Therese Ann Markow,Thomas Kaufman,Manolis Kellis,William Gelbart,Venky Iyer, Daniel Pollard, Timothy Sackton, Amanda Larracuente, Nadia Singh, and includingWojciech Makalowski,Mohamed Noor,Temple F. Smith,Craig Venter,Peter Keightley,andLeonid Boguslavskyamong its contributors – presents ten new genomes and combines those with previously released genomes forD. melanogasterandD. pseudoobscurato analyse the evolutionary history and common genomic structure of the genus. This includes the discovery oftransposable elementsand illumination of their evolutionary history.[78]Bartolomé et al. 2009 find at least1⁄3of the TEs inD. melanogaster,D. simulansandD. yakubahave been acquired byhorizontal transfer.They find an average of0.035 HT TEs⁄TE family⁄million years.Bartolomé also finds HT TEs follow other relatedness metrics, withD. melanogaster⇔D. simulansevents being twice as common as either of them ⇔D. yakuba.[78]

See also

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