Drosophila melanogaster

3034: 1074:(having multiple sexual partners at the same time). 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. 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. 2891: 3017: 785:; 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. 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 619: 2525: 994: 1571: 1370: 1897:) undergoes rapid DNA replication and 13 nuclear divisions until about 5000 to 6000 nuclei accumulate in the unseparated cytoplasm of the embryo. By the end of the eighth division, most nuclei have migrated to the surface, surrounding the yolk sac (leaving behind only a few nuclei, which will become the yolk nuclei). After the 10th division, the pole cells form at the posterior end of the embryo, segregating the germ line from the syncytium. Finally, after the 13th division, cell membranes slowly invaginate, dividing the syncytium into individual somatic cells. Once this process is completed, 3365:

stitched and aligned together to create a 3D image volume. The methods used in reconstruction and initial analysis of the such datasets followed. Due to advancements in deep learning, automated methods for image segmentation have made large scale reconstruction providing dense reconstructions of all the neurites within the volume. Furthermore, the resolution of electron microscopy illuminates ultrastructural variations between neurons as well as the location of individual synapses, thereby providing a wiring diagram of synaptic connectivity between all neurites within the given dataset.

3280:. This means that three of the legs swing together while the other three remain stationary, or in stance. Specifically, the middle leg moves in phase with the contralateral front and hind legs. However, variability around the tripod configuration exists along a continuum, meaning that flies do not exhibit distinct transitions between different gaits. At fast walking speeds, the walking configuration is mostly tripod (3 legs in stance), but at slower walking speeds, flies are more likely to have four (tetrapod) or five legs in stance (wave). These transitions may help to optimize 2911: 2843: 1590:, there have been many discoveries of the vestigial gene in other vertebrates and their functions within the vertebrates. The vestigial gene is considered to be one of the most important genes for wing formation, but when it becomes over expressed the issue of ectopic wings begin to form. The vestigial gene acts to regulate the expression of the wing imaginal discs in the embryo and acts with other genes to regulate the development of the wings. A mutated vestigial allele removes an essential sequence of the DNA required for correct development of the wings. 44: 2851: 1654: 1674: 1113: 62: 3342: 1566:. The vermilion mutation is sex-linked and recessive. The gene that is defect lies on the X chromosome. The brown ommochromes are synthesised from kynurenine, which is made from tryptophane. Vermilion flies cannot convert tryptophane into kynurenine and thus cannot make ommochromes, either. Vermilion mutants live longer than wild-type flies. This longer life span may be associated with the reduced amount of tryptophan converted to kynurenine in vermilion flies. 2931:(345 nm). The R7p cells are strictly paired with the R8p cells that express Rh5, which maximally absorbs violet light (437 nm). The other, the yellow R7y cells express a blue-absorbing screening pigment and the opsin Rh4, which maximally absorbs UV-light (375 nm). The R7y cells are strictly paired with R8y cells that express Rh6, which maximally absorbs UV-light (508 nm). In a subset of omatidia both R7 and R8 cells express the opsin Rh3. 1215: 3016: 3265: 900: 2935:

absorption maxima of opsins differs and thus also the sensitivity of their photoreceptor cells. With screening pigment, the opsin Rh3 is short wave shifted from 345 nm to 330 nm and Rh4 from 375 nm to 355 nm. Whether screening pigment is present does not make a practical difference for the opsin Rh5 (435 nm and 437 nm), while the opsin R6 is long wave shifted by 92 nm from 508 nm to 600 nm.

941: 2530: 2529: 2526: 12371: 995: 762: 2531: 2816: 996: 9570: 764: 2528: 2558:, one occurring in the early embryo and one occurring during development from larva to adult. However Drosophila hemocytes do not renew over the adult lifespan, and so the fly has a finite number of hemocytes that decrease over the course of its lifespan. Hemocytes are also involved in regulating cell-cycle events and apoptosis of aberrant tissue (e.g. cancerous cells) by producing Eiger, a 998: 3033: 2839:, and a cornea. Wild-type flies have reddish pigment cells, which serve to absorb excess blue light so the fly is not blinded by ambient light. Eye color genes regulate cellular vesicular transport. The enzymes needed for pigment synthesis are then transported to the cell's pigment granule, which holds pigment precursor molecules. 2073:). Male sterility can be reversible if adults are returned to an optimal temperature after developing at stressful temperatures. Male flies are smaller and more successful at defending food/oviposition sites when reared at 25 °C versus 18 °C; thus smaller males will have increased mating success and reproductive output. 2089:. Unlike humans, the Y chromosome does not confer maleness; rather, it encodes genes necessary for making sperm. Sex is instead determined by the ratio of X chromosomes to autosomes. Furthermore, each cell "decides" whether to be male or female independently of the rest of the organism, resulting in the occasional occurrence of 927:. 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 naïve females (which have not observed the copulation of others). This behavior is sensitive to environmental conditions, and females copulate less in bad weather conditions. 2521:

peptide Listericin. The mechanisms through which many of these proteins act is still under investigation. For instance, the TEPs appear to promote phagocytosis of Gram-positive bacteria and the induction of the toll pathway. As a consequence, flies lacking TEPs are susceptible to infection by toll pathway challenges.

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18 °C (64 °F) it takes 19 days and at 12 °C (54 °F) it takes over 50 days. Under crowded conditions, development time increases, while the emerging flies are smaller. Females lay some 400 eggs (embryos), about five at a time, into rotting fruit or other suitable material such as decaying

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Popovych, Sergiy; Macrina, Thomas; Kemnitz, Nico; Castro, Manuel; Nehoran, Barak; Jia, Zhen; Bae, J. Alexander; Mitchell, Eric; Mu, Shang; Trautman, Eric T.; Saalfeld, Stephan; Li, Kai; Seung, Sebastian (March 27, 2022). "Petascale pipeline for precise alignment of images from serial section electron

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in 2020, and a dense connectome of the entire female adult nerve cord in 2021. Generally, these datasets are acquired by sectioning the tissue (e.g. the brain) into thin sections (on order of ten or hundreds of nanometers). Each section is then imaged using an electron microscope and these images are

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from returning to grooming behaviors that have already been accessed in the grooming sequence. The order of grooming behaviors in the suppression hierarchy is thought to be related to the priority of cleaning a specific body part. For example, the eyes and antennae are likely executed early on in the

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but lack an adaptive immune response. However, the core elements of this innate immune response are conserved between humans and fruit flies. As a result, the fruit fly offers a useful model of innate immunity for disentangling genetic interactions of signalling and effector function, as flies do not

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has a series of mating behaviors that enable them to copulate within a given environment and therefore contribute to their fitness. After Morgan's discovery of the white-eye mutation being sex-linked, a study led by Sturtevant (1915) concluded that white-eyed males were less successful than wild-type

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wild type typically expresses a brick red eye color. The white eye mutation in fruit flies is caused due to the absence of two pigments associated with red and brown eye colors; peridines (red) and ommochromes (brown). In January 1910, Thomas Hunt Morgan first discovered the white gene and denoted it

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Grooming behaviors have been shown to be executed in a suppression hierarchy. This means that grooming behaviors that occur at the beginning of the sequence prevent those that come later in the sequence from occurring simultaneously, as the grooming sequence consists of mutually exclusive behaviors.

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was found to stimulate gustatory receptor neurons, which was necessary to stimulate aggression. However, once the amount of food becomes greater than a certain amount, the competition between males lowers. This is possibly due to an over-abundance of food resources. On a larger scale, food was found

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develop at cold temperatures they will have greater cold tolerance, but if cold-reared flies are maintained at warmer temperatures their cold tolerance decreases and heat tolerance increases over time. Because insects typically only mate in a specific range of temperatures, their cold/heat tolerance

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grow inside the larva. Imaginal discs develop to form most structures of the adult body, such as the head, legs, wings, thorax, and genitalia. Cells of the imaginal disks are set aside during embryogenesis and continue to grow and divide during the larval stages—unlike most other cells of the larva,

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allows the female fly to pause her body movements to copulate. 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

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Sharkey et al. give the absorption maximum of Rh3 as 334 nm in their result section. However, in the introduction and the material and methods section they give it as 345 nm. For both values, they cite Feiler et al., who reported 345 nm only. Therefore, this seems to be a mistake and

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is sometimes referred to as a pest due to its tendency to live in human settlements where fermenting fruit is found. Flies may collect in homes, restaurants, stores, and other locations. The name and behavior of this species of fly have led to the misconception that it is a biological security risk

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proteins, which specifically bind the C termini of target proteins. Disruption of the complex by mutations in either the PDZ domains or the target proteins reduces the efficiency of signaling. For example, disruption of the interaction between InaC, the protein kinase C, and InaD results in a delay

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Circulating hemocytes are key regulators of infection. This has been demonstrated both through genetic tools to generate flies lacking hemocytes, or through injecting microglass beads or lipid droplets that saturate hemocyte ability to phagocytose a secondary infection. Flies treated like this fail

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pathway genes. JAK-STAT signalling is induced upon various organismal stresses such as heat stress, dehydration, or infection. JAK-STAT induction leads to the production of a number of stress response proteins including Thioester-containing proteins (TEPs), Turandots, and the putative antimicrobial

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wings are 1.5x shorter than wild-type but are believed to have the same number of cells. This is due to the lack of complete flattening by these cells, making the overall structure of the wing seem shorter in comparison. The pathway of wing expansion is regulated by a signal-receptor pathway, where

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has two more opsins: The ocelli express the opsin Rh2, which maximally absorbs violet light (~420 nm). And the opsin Rh7, which maximally absorbs UV-light (350 nm) with an unusually long wavelength tail up to 500 nm. The long tail disappears if a lysine at position 90 is replaced by

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with a frequency of about a day (24 hours). They found mutants with faster and slower rhythms, as well as broken rhythms—flies that move and rest in random spurts. Work over the following 30 years has shown that these mutations (and others like them) affect a group of genes and their products that

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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

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species. The shortest development time (egg to adult), 7 days, is achieved at 28 °C (82 °F). Development times increase at higher temperatures (11 days at 30 °C or 86 °F) due to heat stress. Under ideal conditions, the development time at 25 °C (77 °F) is 8.5 days, at

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In May 2017 a paper published in bioRxiv presented an electron microscopy image stack of the whole adult female brain at synaptic resolution. The volume is available for sparse tracing of selected circuits. Since then, multiple datasets have been collected including a dense connectome of half the

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signaling, which are important pathways for regulating arousal in insects. Due to reduced aggression, sleep-deprived male flies were found to be disadvantaged at mating compared to normal flies. However, when octopamine agonists were administered upon these sleep-deprived flies, aggression levels

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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

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These genes and their modes of action form a complex regulatory network that guides the embryonic development of Drosophila melanogaster. They influence cell differentiation, segment formation, and axial patterning in the embryo, ultimately leading to the development of a fully formed adult fruit

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encodes seven opsins, five of those are expressed in the omatidia of the eye. The photoreceptor cells R1-R6 express the opsin Rh1, which absorbs maximally blue light (around 480 nm), however the R1-R6 cells cover a broader range of the spectrum than an opsin would allow due to a sensitising

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As with most insects, aggressive behaviors between male flies commonly occur in the presence of courting a female and when competing for resources. Such behaviors often involve raising wings and legs towards the opponent and attacking with the whole body. Thus, it often causes wing damage, which

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Presence or absence of functional sex-lethal proteins now go on to affect the transcription of another protein known as doublesex. In the absence of sex-lethal, doublesex will have the fourth exon removed and be translated up to and including exon 6 (DSX-M), while in its presence the fourth exon

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Hunt eventually discovered that the gene followed a similar pattern of inheritance related to the meiotic segregation of the X chromosome. He discovered that the gene was located on the X chromosome with this information. This led to the discovery of sex-linked genes and also to the discovery of

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These genes are encoded in the female fruit fly and are present in the early stages of embryo development. They determine the embryo's main features and early development. For example, the gene called Bicoid regulates the formation of the embryo's anterior end, and its absence leads to an embryo

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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

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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

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Winding, Michael; Pedigo, Benjamin D.; Barnes, Christopher L.; Patsolic, Heather G.; Park, Youngser; Kazimiers, Tom; Fushiki, Akira; Andrade, Ingrid V.; Khandelwal, Avinash; Valdes-Aleman, Javier; Li, Feng; Randel, Nadine; Barsotti, Elizabeth; Correia, Ana; Fetter, Richard D. (March 10, 2023).

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of the fly body, but the opposite case with inertia as the dominant force may occur. However, subsequent work showed that while the viscous effects on the insect body during flight may be negligible, the aerodynamic forces on the wings themselves actually cause fruit flies' turns to be damped

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comparing the fruit fly and human genome estimated that about 60% of genes are conserved between the two species. About 75% of known human disease genes have a recognizable match in the genome of fruit flies, and 50% of fly protein sequences have mammalian homologs . An online database called

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Phelps, Jasper S.; Hildebrand, David Grant Colburn; Graham, Brett J.; Kuan, Aaron T.; Thomas, Logan A.; Nguyen, Tri M.; Buhmann, Julia; Azevedo, Anthony W.; Sustar, Anne; Agrawal, Sweta; Liu, Mingguan; Shanny, Brendan L.; Funke, Jan; Tuthill, John C.; Lee, Wei-Chung Allen (February 2, 2021).

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However, these absorption maxima of the opsins where measured in white eyed flies without screening pigments (Rh3-Rh6), or from the isolated opsin directly (Rh1). Those pigments reduce the light that reaches the opsins depending on the wavelength. Thus in fully pigmented flies, the effective

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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

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During oogenesis, cytoplasmic bridges called "ring canals" connect the forming oocyte to nurse cells. Nutrients and developmental control molecules move from the nurse cells into the oocyte. In the figure to the left, the forming oocyte can be seen to be covered by follicular support cells.

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designation refers to the lengths of their wings, which do not stretch beyond their body and, thus, are notably shorter than the wild-type length. He also noted its inheritance is connected to the sex of the fly and could be paired with the inheritance of other sex-determined traits such as

2006:, and territorial success, while development at 18 °C increases body mass, wing size, all of which are tied to fitness. Moreover, developing at certain low temperatures produces proportionally large wings which improve flight and reproductive performance at similarly low temperatures ( 725:

Drosophila melanogaster, commonly known as the fruit fly, has been a significant model organism in embryonic development research. Many of its genes that regulate embryonic development and their mechanisms of action have been crucial in understanding the fundamental principles of embryonic

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technique. This technique made it possible to assign a particular characteristic to a specific anatomical location. For example, this technique showed that male courtship behavior is controlled by the brain. Mosaic fate mapping also provided the first indication of the existence of

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also possesses several immune mechanisms to both shape the microbiota and prevent excessive immune responses upon detection of microbial stimuli. For instance, secreted PGRPs with amidase activity scavenge and degrade immunostimulatory DAP-type PGN in order to block Imd activation.

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differentially expresses a core group of 252 genes upon infection with most bacteria. This core group of genes is associated with gene ontology categories such as antimicrobial response, stress response, secretion, neuron-like, reproduction, and metabolism among others.

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classes: the mutants that show a complete loss of pigmentation from the cuticle (y-type) and other mutants that show a mosaic pigment pattern with some regions of the cuticle (wild type, y2-type). The role of the yellow gene is diverse and is responsible for changes in

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sequenced genome of 139.5 million base pairs has been annotated and contains around 15,682 genes according to Ensemble release 73. More than 60% of the genome appears to be functional non-protein-coding DNA involved in gene expression control. Determination of sex in

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female genitalia. Finally, the male curls his abdomen and attempts copulation. Females can reject males by moving away, kicking, and extruding their ovipositor. Copulation lasts around 15–20 minutes, during which males transfer a few hundred, very long (1.76 mm)

763: 1915:

The gene network (transcriptional and protein interactions) governing the early development of the fruit fly embryo is one of the best understood gene networks to date, especially the patterning along the anteroposterior (AP) and dorsoventral (DV) axes (See under

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Courtship behavior in male. The male first showed wing extension (stage 1), and later other steps such as abdomen bending (stage 2), then frequent attempted copulation, licking and even ejaculation (stage 3), finally the male fell over and was on its back (stage

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temperature-induced developmental plasticity can be beneficial and/or detrimental. Most often lower developmental temperatures reduce growth rates which influence many other physiological factors. For example, development at 25 °C increases walking speed,

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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

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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

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This gene family regulates segmentation and axial patterning in development. They act as regulatory factors that determine cell fate in embryonic development. For example, the gene called Antennapedia regulates the formation of anterior limbs in the embryo.

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These are molecules that form gradients in embryonic development and regulate cell fate depending on their position in the gradient. For example, the Hedgehog morphogen regulates the differentiation of segments and segment identity in the fruit fly embryo.

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multiple mutants (clockwise from top): brown eyes and black cuticle (2 mutations), cinnabar eyes and wildtype cuticle (1 mutation), sepia eyes and ebony cuticle, vermilion eyes and yellow cuticle, white eyes and yellow cuticle, wildtype eyes and yellow

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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.

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opsins in white eyed flies. The sensitivities of Rh3–R6 are modelled with opsin templates and sensitivity estimates from Salcedo et al. (1999). The opsin Rh1 (redrawn from Salcedo et al.) has a characteristic shape as it is coupled to a UV-sensitising

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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. Then the larvae encapsulate in the

611:; females are about 2.5 mm (0.10 in) long; males are slightly smaller. 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 1476:, a beta amino acid. The phenotypic expression of this mutation varies based on the genotype of the individual; for example, whether the specimen is homozygotic or heterozygotic results in a darker or less dark appearance. This genetic mutation is 1514:

the neurohormone bursicon interacts with its complementary G protein-coupled receptor; this receptor drives one of the G-protein subunits to signal further enzyme activity and results in development in the wing, such as apoptosis and growth.

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insect, so it undergoes a full metamorphosis. Their life cycle is broken down into 4 stages: embryo, larva, pupa, adult. The eggs, which are about 0.5 mm long, hatch after 12–15 hours (at 25 °C or 77 °F). The resulting

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larval brain at the synapse level, and an analysis of its architecture was published. The larval brain consists of 3016 neurons and 548,000 synaptic sites, whereas the adult brain has about 150,000 neurons and 150 million synapses.

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pigment that adds two sensitivity maxima in the UV-range (355 and 370 nm). The R7 cells come in two types with yellow and pale rhabdomeres (R7y and R7p). The pale R7p cells express the opsin Rh3, which maximally absorbs

2906:: The top row shows two of the six outer photoreceptor cells (R1-R6) and the inner R7 and R8 cells. The bottom row shows the different opsins (Rh1, Rh3, Rh4, Rh5, and Rh6) the cells express. Figure from Sharkey et al. (2020). 5868:

Bilousov OO, Katanaev VL, Demydov SV, Kozeretska IA (March–April 2013). "The downregulation of the miniature gene does not replicate miniature loss-of-function phenotypes in Drosophila melanogaster wing to the full extent".

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Prudhommeau C, Proust J. UV irradiation of poplar cells of Drosophila melanogaster embryos. V. A study of the meiotic recombination in females with chromosomes of different structure. Mutat Res. 1974 Apr;23(1):63-6. PMID

2769:, an anti-aphrodisiac pheromone used by males to mark females after copulation and to deter other males from mating. This male-specific pheromone causes an increase in male-male aggression when detected by another male's 632:

can be distinguished from related species by the following combination of features: gena ~1/10 diameter of eye at greatest vertical height; wing hyaline and with costal index 2.4; male protarsus with a single row of ~12

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in this species. Males distinguish between conspecific males and females and direct persistent courtship preferentially toward females thanks to a female-specific sex pheromone which is mostly produced by the female's

1536:, which is encoded on chromosome 3L. The gene has a premature stop codon in sepia flies, so that the flies cannot produce the pyrimidodiazepine synthase and thus no red pigment, so that the eyes stay sepia. The sepia 739:

These genes are activated in later stages of embryo development when the fruit fly embryo begins to produce its own genetic products. For example, the hunchback gene regulates the formation of segments in the embryo.

2055:) is maximized when reared at 25 °C (versus 17 °C and 29 °C) regardless of adult temperature. Across a wide range of developmental temperatures, females tend to have greater heat tolerance than males. 806:

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

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greater mating efficiency for experienced males over naïve males. This modification also appears to have obvious evolutionary advantages, as increased mating efficiency is extremely important in the eyes of

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Schewe MJ, Suzuki DT, Erasmus U. The genetic effects of mitomycin C in Drosophila melanogaster. II. Induced meiotic recombination. Mutat Res. 1971 Jul;12(3):269-79. doi: 10.1016/0027-5107(71)90015-7. PMID

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Paloma Álvarez-Rendón, Jéssica; Manuel Murillo-Maldonado, Juan; Rafael Riesgo-Escovar, Juan (2023). "The insulin signaling pathway a century after its discovery: Sexual dimorphism in insulin signaling".

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flies can sense air currents with the hairs on their backs. Their eyes are sensitive to slight differences in light intensity and will instinctively fly away when a shadow or other movement is detected.

1586:- A spontaneous mutation, discovered in 1919 by Thomas Morgan and Calvin Bridges. Vestigial wings are those not fully developed and that have lost function. Since the discovery of the vestigial gene in 2474:

The immune response to infection can involve up to 2,423 genes, or 13.7% of the genome. Although the fly's transcriptional response to microbial challenge is highly specific to individual pathogens,

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has been extensively studied, as its small size, short generation time, and large brood size makes it ideal for genetic studies. It is also unique among model organisms in that cleavage occurs in a

1320:

Genetic transformation techniques have been available since 1987. One approach of inserting foreign genes into the Drosophila genome involves P elements. The transposable P elements, also known as

1275:, "puffs", which indicate regions of transcription, hence gene activity. The under-replication of rDNA occurs resulting in only 20% of DNA compared to the brain. Compare to the 47%, less rDNA in 861:

also cause increased physiologic signs of aging, such as shorter lifespan, higher tumor incidence, muscle degeneration, reduced climbing ability, altered behavior and reduced locomotor activity.

2773:. However, upon inserting a mutation that makes the flies irresponsive to cVA, no aggressive behaviors were seen. This shows how there are multiple modalities for promoting aggression in flies. 8779:"The 2017 Nobel Prize in Physiology or Medicine jointly to Jeffrey C. Hall, Michael Rosbash and Michael W. Young for their discoveries of molecular mechanisms controlling the circadian rhythm" 4849:

Moshitzky P, Fleischmann I, Chaimov N, Saudan P, Klauser S, Kubli E, Applebaum SW (1996). "Sex-peptide activates juvenile hormone biosynthesis in the Drosophila melanogaster corpus allatum".

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Since then, Benzer and others have used behavioral screens to isolate genes involved in vision, olfaction, audition, learning/memory, courtship, pain, and other processes, such as longevity.

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development regulation in many multicellular organisms, including humans. Here are some important genes regulating embryonic development in Drosophila melanogaster and their modes of action:

6004:"Purification and properties of the enzymes from Drosophila melanogaster that catalyze the conversion of dihydroneopterin triphosphate to the pyrimidodiazepine precursor of the drosopterins" 2051:). Under stressful temperatures, these structures will develop to smaller ultimate sizes and decrease a female's reproductive output. Early fecundity (total eggs laid in first 10 days post- 1636:. This mutation can be easily identified by the atypical yellow pigment observed in the cuticle of the adult flies and the mouth pieces of the larva. The y mutation comprises the following 1877: 1614:

The white-eye mutation leads to several disadvantages in flies, such as a reduced climbing ability, shortened life span, and lowered resistance to stress when compared to wild type flies.

2721:

In order for aggression to occur, male flies produce sounds to communicate their intent. A 2017 study found that songs promoting aggression contain pulses occurring at longer intervals.

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embryo happens so quickly, no proper checkpoints exist, so mistakes may be made in division of the DNA. To get around this problem, the nuclei that have made a mistake detach from their

12341: 1324:, are segments of bacterial DNA that are transferred into the fly genome. Transgenic flies have already contributed to many scientific advances, e.g., modeling such human diseases as 10271:

Huber A, Schulz S, Bentrop J, Groell C, Wolfrum U, Paulsen R (April 1997). "Molecular cloning of Drosophila Rh6 rhodopsin: the visual pigment of a subset of R8 photoreceptor cells".

5901:"Identification and characteristics of the structural gene for the Drosophila eye colour mutant sepia, encoding PDA synthase, a member of the omega class glutathione S-transferases" 661: 11164:

Mathis A, Mamidanna P, Cury KM, Abe T, Murthy VN, Mathis MW, Bethge M (September 2018). "DeepLabCut: markerless pose estimation of user-defined body parts with deep learning".

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Zamudio KR, Huey RB, Crill WD (1995). "Bigger isn't always better: body size, developmental and parental temperature and male territorial success in Drosophila melanogaster".

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Abram PK, Boivin G, Moiroux J, Brodeur J (November 2017). "Behavioural effects of temperature on ectothermic animals: unifying thermal physiology and behavioural plasticity".

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of odors, with additional features including similarity of novel odor to that of previously experienced examples, and time elapsed since previous experience of the same odor.

9803:"Blue- and green-absorbing visual pigments of Drosophila: ectopic expression and physiological characterization of the R8 photoreceptor cell-specific Rh5 and Rh6 rhodopsins" 2656:

Male flies sing to the females during courtship using their wings to generate sound, and some of the genetics of sexual behavior have been characterized. In particular, the

592:

fruit flies are yellow-brown, with brick-red eyes and transverse black rings across the abdomen. The black portions of the abdomen are the inspiration for the species name (

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Xu CS, Januszewski M, Lu Z, Takemura SY, Hayworth K, Huang G, Shinomiya K, Maitin-Shepard J, Ackerman D, Berg S, Blakely T, et al. (2020). "A connectome of the adult

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opsins Rh1, Rh3, Rh4, Rh5, and Rh6 measured in their native photoreceptor cells in red eye flies with screening pigment. Each spectral curve is the average from six flies.

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Raghu P, Colley NJ, Webel R, James T, Hasan G, Danin M, et al. (May 2000). "Normal phototransduction in Drosophila photoreceptors lacking an InsP(3) receptor gene".

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reprogramming. The y gene is an ideal gene to study as it is visibly clear when an organism has this gene, making it easier to understand the passage of DNA to offspring.

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A high-level connectome, at the level of brain compartments and interconnecting tracts of neurons, exists for the full fly brain. A version of this is available online.

6498:

Wittkopp PJ, True JR, Carroll SB (April 2002). "Reciprocal functions of the Drosophila yellow and ebony proteins in the development and evolution of pigment patterns".

3292:

Flies fly via straight sequences of movement interspersed by rapid turns called saccades. During these turns, a fly is able to rotate 90° in less than 50 milliseconds.

3284:. Because flies are so small, inertial forces are negligible compared with the elastic forces of their muscles and joints or the viscous forces of the surrounding air. 2649:

for 2017 was awarded to Jeffrey C. Hall, Michael Rosbash, Michael W. Young for their works using fruit flies in understanding the "molecular mechanisms controlling the

3170:(PKC) known as InaC. These proteins interact with other proteins and have been shown to be necessary for shut off of the light response. In addition, proteins called 2781:

Specifically, when competing for food, aggression occurs based on amount of food available and is independent of any social interactions between males. Specifically,

2807:

were seen to be increased and sexual fitness was subsequently restored. Therefore, this finding implicates the importance of sleep in aggression between male flies.

2003: 2634:, etc.) were isolated by William "Chip" Quinn while in Benzer's lab, and were eventually shown to encode components of an intracellular signaling pathway involving 1268:

Males and females are readily distinguished, and virgin females can be easily identified by their light-colored, translucent abdomen, facilitating genetic crossing.

4597:

Cook R, Connolly K (1973). "Rejection Responses by Female Drosophila melanogaster: Their Ontogeny, Causality and Effects upon the Behaviour of the Courting Male".

1188:

microscopes, which enhanced their observations. Morgan and his students eventually elucidated many basic principles of heredity, including sex-linked inheritance,

7384:

Crill WD, Huey RB, Gilchrist GW (June 1996). "Within- and Between-Generation Effects of Temperature on the Morphology and Physiology of Drosophila Melanogaster".

10801:

Kumbalasiri T, Rollag MD, Isoldi MC, Castrucci AM, Provencio I (March 2007). "Melanopsin triggers the release of internal calcium stores in response to light".

3088:

to all-trans-3-hydroxyretinal. Rh undergoes a conformational change into its active form, metarhodopsin. Metarhodopsin activates Gq, which in turn activates a

2599:. This clock is found in a wide range of fly cells, but the clock-bearing cells that control activity are several dozen neurons in the fly's central brain. 11489: 10368:

Feiler R, Harris WA, Kirschfeld K, Wehrhahn C, Zuker CS (June 1988). "Targeted misexpression of a Drosophila opsin gene leads to altered visual function".

1619:

males in terms of mating with females. It was found that the greater the density in eye pigmentation, the greater the success in mating for the males of Dr

1472:. The black mutation results in a darker colored body, wings, veins, and segments of the fruit fly's leg. This occurs due to the fly's inability to create 9696:

Ostroy SE, Wilson M, Pak WL (August 1974). "Drosophila rhodopsin: photochemistry, extraction and differences in the norp AP12 phototransduction mutant".

7334:

Sheeba V, Chandrashekaran MK, Joshi A, Sharma VK (January 2002). "Developmental plasticity of the locomotor activity rhythm of Drosophila melanogaster".

3445: 10167:"Ectopic expression of ultraviolet-rhodopsins in the blue photoreceptor cells of Drosophila: visual physiology and photochemistry of transgenic animals" 8589:"Adult Drosophila Lack Hematopoiesis but Rely on a Blood Cell Reservoir at the Respiratory Epithelia to Relay Infection Signals to Surrounding Tissues" 7046:

Weigmann K, Klapper R, Strasser T, Rickert C, Technau G, Jäckle H, et al. (June 2003). "FlyMove--a new way to look at development of Drosophila".

2998:

in Rh1, then Rh1 loses light sensitivity but still responds to aristolochic acid. Thus, the lysine is not needed for Rh1 to function as chemoreceptor.

7847:

Cohet Y, David J (January 1978). "Control of the adult reproductive potential by preimaginal thermal conditions: A study in Drosophila melanogaster".

3205:

Unlike vertebrate metarhodopsin, invertebrate metarhodopsin can be converted back into rhodopsin by absorbing a photon of orange light (580 nm).

2467:, which are analogous to mammalian monocytes/macrophages. Hemocytes also possess a significant role in mediating humoral immune responses such as the 2028:

While the traits described above are expected to manifest similarly across sexes, developmental temperature can also produce sex-specific effects in

1364: 12065:

Scheffer LK, Xu CS, Januszewski M, Lu Z, Takemura SY, Hayworth KJ, Huang G, Shinomiya K, Maitlin-Shepard J, Berg S, Clements J, et al. (2020).

10314:

Pollock JA, Benzer S (June 1988). "Transcript localization of four opsin genes in the three visual organs of Drosophila; RH2 is ocellus specific".

3199: 11672:

Meinertzhagen IA, O'Neil SD (March 1991). "Synaptic organization of columnar elements in the lamina of the wild type in Drosophila melanogaster".

4388: 2662:

gene has several different splice forms, and male flies expressing female splice forms have female-like behavior and vice versa. The TRP channels

1771:

of X chromosomes to autosomes, not because of the presence of a Y chromosome as in human sex determination. Although the Y chromosome is entirely

7710:"Different mechanisms underlie phenotypic plasticity and interspecific variation for a reproductive character in drosophilids (Insecta: Diptera)" 2878:

molecules, the visual protein that absorbs light. The other visual proteins are also tightly packed into the microvilli, leaving little room for

2407:

immune system can be divided into two responses: humoral and cell-mediated. The former is a systemic response mediated in large part through the

8363:"Cooperative regulation of the induction of the novel antibacterial Listericin by peptidoglycan recognition protein LE and the JAK-STAT pathway" 8261:"Thioester-containing proteins regulate the Toll pathway and play a role in Drosophila defence against microbial pathogens and parasitoid wasps" 5352:

Kilbey BJ, MacDonald DJ, Auerbach C, Sobels FH, Vogel EW (June 1981). "The use of Drosophila melanogaster in tests for environmental mutagens".

3707: 2890: 5221:

Vartak VR, Varma V, Sharma VK (February 2015). "Effects of polygamy on the activity/rest rhythm of male fruit flies Drosophila melanogaster".

2496:

have to contend with interference of adaptive immune mechanisms that could confuse results. Various genetic tools, protocols, and assays make

1951:, inside which the larval tissues are reabsorbed and the imaginal tissues undergo extensive morphogenetic movements to form adult structures. 13010: 3190: 1691: 1529: 795: 6175:

Simon E, Faucheux C, Zider A, Thézé N, Thiébaud P (July 2016). "From vestigial to vestigial-like: the Drosophila gene that has taken wing".

1486:- The brown eye mutation results from inability to produce or synthesize pteridine (red) pigments, due to a point mutation on chromosome II. 586:

Female fruit flies are substantially larger than male fruit flies, with females having bodies that are up to 30% larger than an adult male.

457:, and large number of offspring per generation. It was originally an African species, with all non-African lineages having a common origin. 13117: 11880: 9589:"Phylogenomics of Opsin Genes in Diptera Reveals Lineage-Specific Events and Contrasting Evolutionary Dynamics in Anopheles and Drosophila" 2387:

which encodes a stop codon will produce a truncated version of the protein (DSX-F). DSX-F causes transcription of Yolk proteins 1 and 2 in

802:. 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 9135:

Sengupta S, Smith DP (2014). "How Drosophila Detect Volatile Pheromones: Signaling, Circuits, and Behavior". In Mucignat-Caretta C (ed.).

4219: 1970:, also referred to as developmental plasticity. As in all insects, environmental factors can influence several aspects of development in 1206:

also has importance in environmental mutagenesis research, allowing researchers to study the effects of specific environmental mutagens.

1104:

in biological research, particularly in genetics and developmental biology. It is also employed in studies of environmental mutagenesis.

7287:"Developmental plasticity and stability in the tracheal networks supplying Drosophila flight muscle in response to rearing oxygen level" 4862: 4020:"Quantitative Genetics of Drosophila Melanogaster. II. Heritabilities and Genetic Correlations between Sexes for Head and Thorax Traits" 2463:

or inhibiting intracellular processes. The cellular immune response instead refers to the direct activity of blood cells (hemocytes) in

4497:

Fernández-Moreno MA, Farr CL, Kaguni LS, Garesse R (2007). "Drosophila melanogaster as a Model System to Study Mitochondrial Biology".

4070:

Ewart GD, Howells AJ (January 1, 1998). "ABC transporters involved in transport of eye pigment precursors in Drosophila melanogaster".

3510: 3235:

consistently begin a grooming sequence by using their front legs to clean the eyes, then the head and antennae. Using their hind legs,

2638:, protein kinase A, and a transcription factor known as CREB. These molecules were shown to be also involved in synaptic plasticity in 1947:

which have differentiated to perform specialized functions and grow without further cell division. At metamorphosis, the larva forms a

12528: 12623: 7568:"Reversibility of developmental heat and cold plasticity is asymmetric and has long-lasting consequences for adult thermal tolerance" 4719:

Price CS, Dyer KA, Coyne JA (July 1999). "Sperm competition between Drosophila males involves both displacement and incapacitation".

7479:

French V, Feast M, Partridge L (November 1998). "Body size and cell size in Drosophila: the developmental response to temperature".

12945: 2554:

to phagocytose bacteria upon infection, and are correspondingly susceptible to infection. These hemocytes derive from two waves of

1795: 958: 190: 175: 7432:"Male sterility at extreme temperatures: a significant but neglected phenomenon for understanding Drosophila climatic adaptations" 599:). The brick-red color of the eyes of the wild type fly are due to two pigments: xanthommatin, which is brown and is derived from 12971: 8507:"Identification of lipoteichoic acid as a ligand for draper in the phagocytosis of Staphylococcus aureus by Drosophila hemocytes" 8122:"Negative regulation by amidase PGRPs shapes the Drosophila antibacterial response and protects the fly from innocuous infection" 6128:"The extended life span of Drosophila melanogaster eye-color (white and vermilion) mutants with impaired formation of kynurenine" 3780: 3096: 2663: 1509:

eyes. The wings may also demonstrate other characteristics deviant from the wild-type wing, such as a duller and cloudier color.

10004:

Kirschfeld K, Franceschini N (June 1977). "Photostable pigments within the membrane of photoreceptors and their possible role".

1990:, where flies raised under constant dark or light are less active as adults than those raised under a 12-hour light/dark cycle. 10049:"A rhodopsin gene expressed in photoreceptor cell R7 of the Drosophila eye: homologies with other signal-transducing molecules" 7190:"Locomotor performance of Drosophila melanogaster: interactions among developmental and adult temperatures, age, and geography" 11207:

Strauss R, Heisenberg M (August 1990). "Coordination of legs during straight walking and turning in Drosophila melanogaster".

4139:"Identification key for drosophilid species (Diptera, Drosophilidae) exotic to the Neotropical Region and occurring in Brazil" 3744: 1917: 1354:

Sexual mosaics can be readily produced, providing an additional tool for studying the development and behavior of these flies.

12654: 12287: 12264: 9146: 8762: 7962: 6929: 5336: 4694: 4682: 4514: 4355: 4087: 3792: 3728: 2786:

to determine the boundaries of a territory since flies were observed to be more aggressive at the food's physical perimeter.

2646: 2424: 1939:(gut), as well as extensive body segmentation until finally hatching from the surrounding cuticle into a first-instar larva. 160: 12976: 5394:; Holt RA; Evans CA; Gocayne JD; Amanatides PG; et al. (March 2000). "The genome sequence of Drosophila melanogaster". 4462: 3674: 1605:. The discovery of the white-eye mutation by Morgan brought about the beginnings of genetic experimentation and analysis of 61: 8687:

Sturtevant AH (1929). "The claret mutant type of Drosophila simulans: a study of chromosome elimination and cell-lineage".

2726: 9969:

Kirschfeld K, Feiler R, Franceschini N (1978). "A photostable pigment within the rhabdomere of fly photoreceptors no. 7".

7979: 4188:"A Review of the Species of Drosophila (Diptera: Drosophilidae) and Genera of Drosophilidae of Northeastern North America" 2447:

to the human liver. The fat body is the primary secretory organ and produces key immune molecules upon infection, such as

1754:

labeled 2, 3, and 4. The fourth chromosome is relatively very small and therefore often ignored, aside from its important

10421:"Drosophila melanogaster rhodopsin Rh7 is a UV-to-visible light sensor with an extraordinarily broad absorption spectrum" 2982:. Opsins without it are not light sensitive. In particular, the Drosophila opsins Rh1, Rh4, and Rh7 function not only as 9179:"Drosophila melanogaster females restore their attractiveness after mating by removing male anti-aphrodisiac pheromones" 3243:

periodically rub their legs together to get rid of excess dust and debris that accumulates during the grooming process.

2044:

Egg size is also affected by developmental temperature, and exacerbated when both parents develop at warm temperatures (

1050:

seems to be evolutionarily significant, as it allows the males to avoid investing energy into futile sexual encounters.

13107: 12906: 8851:"Sound response mediated by the TRP channels NOMPC, NANCHUNG, and INACTIVE in chordotonal organs of Drosophila larvae" 709:

twice (into second- and third-instar larvae), at about 24 and 48 h after hatching. During this time, they feed on the

13092: 12796: 3479:"Vinegar fly" is preferred by a handful of recent publications as being a more accurate description than "fruit fly". 980: 9858:

Kirschfeld K, Franceschini N, Minke B (September 1977). "Evidence for a sensitising pigment in fly photoreceptors".

6043: 1986:

are also subject to developmental plasticity. Light conditions during development affect daily activity patterns in

13087: 13059: 12924: 8464: 8121: 4378: 3410: 2419:

pathways, which are parallel systems for detecting microbes. Other pathways including the stress response pathways

1131: 881: 17: 12591: 1085:

is controlled by the oscillator neurons DN1s and LNDs. Oscillation of the DN1 neurons was found to be effected by

13015: 12937: 12221: 11035:

Dawkins R, Dawkins M (1976). "Hierarchical organization and postural facilitation: rules for grooming in flies".

4642:"Effect of laboratory acclimation on the variation of reproduction-related characters in Drosophila melanogaster" 3405: 12823: 11536: 7609:"Parental and developmental temperature effects on the thermal dependence of fitness in Drosophila melanogaster" 3946:

Kelley, Darcy B.; Bayer, Emily A. (2021). "Sexual dimorphism: Neural circuit switches in the Drosophila brain".

453:

is typically used in research owing to its rapid life cycle, relatively simple genetics with only four pairs of

12487: 12018:"Reconstruction of motor control circuits in adult Drosophila using automated transmission electron microscopy" 3148: 2725:

from fly mutants displaying over-aggressive behaviors found more than 50 auditory-related genes (important for

2635: 1558:- The vermilion mutants cannot produce the brown ommochromes leaving the red drosopterins so that the eyes are 1477: 962: 618: 12553:

of the "Small fly: BIG impact" educational videos explaining the history and importance of the model organism

10216:"A second opsin gene expressed in the ultraviolet-sensitive R7 photoreceptor cells of Drosophila melanogaster" 9911:"The contribution of a sensitizing pigment to the photosensitivity spectra of fly rhodopsin and metarhodopsin" 7800:"Metabolic and functional characterization of effects of developmental temperature in Drosophila melanogaster" 811:. Sex peptide perturbs this homeostasis and dramatically shifts the endocrine state of the female by inciting 408:

are attracted to rotting fruit and fermenting beverages, and are often found in orchards, kitchens and pubs.

12997: 12616: 10111:"Identification of a novel Drosophila opsin reveals specific patterning of the R7 and R8 photoreceptor cells" 8778: 6618:"Ubiquitous selective constraints in the Drosophila genome revealed by a genome-wide interspecies comparison" 4111: 12198: 11344:"Quantification of gait parameters in freely walking wild type and sensory deprived Drosophila melanogaster" 11715:

Rivera-Alba M, Vitaladevuni SN, Mishchenko Y, Mischenko Y, Lu Z, Takemura SY, et al. (December 2011).

8164:

Troha K, Buchon N (September 2019). "Methods for the study of innate immunity in Drosophila melanogaster".

3435: 3136: 2444: 1376:

which carries the Cy allele (right), hence showing a characteristic phenotype of curly wings in adult flies

12386: 8022:"Comparative transcriptomics reveals CrebA as a novel regulator of infection tolerance in D. melanogaster" 2066:

males; although the upper temperature limit can be increased by maintaining strains at high temperatures (

1071: 13112: 12841: 12818: 5632:"The Drosophila black enigma: the molecular and behavioural characterization of the black1 mutant allele" 4456:

Bakker K (1961). "An analysis of factors which determine success in competition for food among larvae of

3420: 2591: 1293: 12846: 11596:"Three-dimensional reconstruction of brain-wide wiring networks in Drosophila at single-cell resolution" 9081:

Versteven M, Vanden Broeck L, Geurten B, Zwarts L, Decraecker L, Beelen M, et al. (February 2017).

8587:

Sanchez Bosch P, Makhijani K, Herboso L, Gold KS, Baginsky R, Woodcock KJ, et al. (December 2019).

5700:

Sherald AF (September 1981). "Intergenic suppression of the black mutation of Drosophila melanogaster".

5539:"tinman and bagpipe: two homeo box genes that determine cell fates in the dorsal mesoderm of Drosophila" 2794:

However, like most behaviors requiring arousal and wakefulness, aggression was found to be impaired via

4382: 3281: 2517: 2420: 1533: 1443: 1046:

males are much less likely to spend time courting nonspecifically in the future. This apparent learned

911:

groups in females have been found to affect copulation behavior and mate choice. One such group in the

798:, which is found in semen. This protein makes the female reluctant to copulate for about 10 days after 6595: 1809:

is being used as a genetic model for several human diseases including the neurodegenerative disorders

13097: 13064: 13025: 12679: 12661: 1982:

produces smaller flight muscles, suggesting negative developmental effects of extreme oxygen levels.

1500:

in 1911. He described the wings as having a similar shape as the wild-type phenotype. However, their

458: 8414:"Drosophila as a model to study the role of blood cells in inflammation, innate immunity and cancer" 8310:

Srinivasan N, Gordon O, Ahrens S, Franz A, Deddouche S, Chakravarty P, et al. (November 2016).

5416: 1461:

larvae. This system of nomenclature results in a wider range of gene names than in other organisms.

1438:

will result in a mutant embryo that does not develop a heart. Scientists have thus called this gene

13102: 12674: 12609: 12536: 3179: 3175: 1854: 1570: 1532:(red) give the eyes the typical red color. The drosopterins are made via a pathway that involves a 1202:

had historically been used in laboratories to study genetics and patterns of inheritance. However,

1166: 850: 12989: 10590:

Vogt K, Kirschfeld K (April 1984). "Chemical identity of the chromophores of fly visual pigment".

8505:

Hashimoto Y, Tabuchi Y, Sakurai K, Kutsuna M, Kurokawa K, Awasaki T, et al. (December 2009).

7900:"The Sex Determination Gene transformer Regulates Male-Female Differences in Drosophila Body Size" 3980: 3159:, and cause release of calcium, but this process does not seem to be essential for normal vision. 12442: 11115:"A suppression hierarchy among competing motor programs drives sequential grooming in Drosophila" 5745:"The development of pigment granules in the eyes of wild type and mutant Drosophila melanogaster" 4777: 4008:"... adult females are 30 % bigger than males; these differences happen during larval life." 3924:. Department of Entomology, College of Agricultural Sciences, Pennsylvania State University. 2017 3353: 1975: 1685:, centimorgan distances are approximate and estimated from the locations of selected mapped loci. 1682: 1233:

Its care and culture require little equipment, space, and expense even when using large cultures.

951: 719: 12810: 7293:. The limits of respiratory function: external and internal constraints on insect gas exchange. 4943:

Cassidy D, Epiney DG, Salameh C, Zhou LT, Salomon RN, Schirmer AE, et al. (November 2019).

4223: 1966:

factors experienced during development will affect developmental resource allocation leading to

1369: 1165:

organisms. All organisms use common genetic systems; therefore, comprehending processes such as

637:

forming a sex comb; male epandrial posterior lobe small and nearly triangular; female abdominal

12758: 12711: 12570: 9638:"Genetic dissection of the photoreceptor system in the compound eye of Drosophila melanogaster" 8207:

Gilbert R, Torres M, Clemens R, Hateley S, Hosamani R, Wade W, Bhattacharya S (February 2020).

5411: 3327: 2452: 2059: 1822: 1818: 1325: 604: 10165:

Feiler R, Bjornson R, Kirschfeld K, Mismer D, Rubin GM, Smith DP, et al. (October 1992).

6947:"Immunity in Drosophila melanogaster--from microbial recognition to whole-organism physiology" 2910: 2745:. This signifies the major role of hearing as a sensory modality in communicating aggression. 1258:(about 10 days at room temperature), so several generations can be studied within a few weeks. 12503: 11905:

Zheng Z, Lauritzen JS, Perlman E, Robinson CG, Nichols M, Milkie D, et al. (July 2018).

11772:

Takemura SY, Bharioke A, Lu Z, Nern A, Vitaladevuni S, Rivlin PK, et al. (August 2013).

7998: 6773:"A systematic analysis of human disease-associated gene sequences in Drosophila melanogaster" 4413: 3356:

column, both in the visual system of the fruit fly, and the alpha lobe of the mushroom body.

3317: 3156: 3112: 2983: 2947:. This mutant then absorbs maximally violet light (450 nm). The opsin Rh7 entrains with 2559: 1967: 1300: 1143: 1047: 229: 213: 13051: 12563:

of the "Small fly: BIG impact" educational videos explaining how research is carried out in

10684:

Katana R, Guan C, Zanini D, Larsen ME, Giraldo D, Geurten BR, et al. (September 2019).

10109:

Chou WH, Hall KJ, Wilson DB, Wideman CL, Townson SM, Chadwell LV, Britt SG (December 1996).

9733:"Characteristics of Drosophila rhodopsin in wild-type and norpA vision transduction mutants" 7798:

Schou MF, Kristensen TN, Pedersen A, Karlsson BG, Loeschcke V, Malmendal A (February 2017).

7524:"Cold rearing improves cold-flight performance in Drosophila via changes in wing morphology" 5504: 5055:"Drosophila mate copying correlates with atmospheric pressure in a speed learning situation" 3519: 3239:

proceed to groom their abdomen, and finally the wings and thorax. Throughout this sequence,

12880: 12805: 12697: 12525: 12128: 11918: 11785: 11728: 11607: 11504: 11447: 10991: 10757: 10697: 10599: 10497: 10432: 10377: 10323: 9867: 9525: 9463: 9301: 9190: 8932: 8907: 8862: 8719: 7856: 7669: 7488: 7343: 7298: 7094: 6549: 6426: 6080: 5801: 5675: 5461: 5403: 5230: 5132: 4897: 4728: 4422: 3415: 2866:, while the 100-μm-long rhabdomere is made up of toothbrush-like stacks of membrane called 2501: 2347:

gene. A secondary promoter causes transcription in both males and females. Analysis of the

1775:, it contains at least 16 genes, many of which are thought to have male-related functions. 1457: 1321: 1307: 778: 412: 12598:—WhyFiles.org article describes how the fruit fly tastes a larva-killing chemical in food. 11717:"Wiring economy and volume exclusion determine neuronal placement in the Drosophila brain" 10939:"Light activation, adaptation, and cell survival functions of the Na+/Ca2+ exchanger CalX" 9801:

Salcedo E, Huber A, Henrich S, Chadwell LV, Chou WH, Paulsen R, Britt SG (December 1999).

9349:"Interaction between sleep and metabolism in Drosophila with altered octopamine signaling" 8708:"A new behavioral bioassay for an analysis of sexual attraction and pheromones in insects" 6271:"Control of Drosophila wing and haltere development by the nuclear vestigial gene product" 593: 471:

are also called "fruit flies". This can cause confusion, especially in the Mediterranean,

8: 13122: 12725: 12704: 12519: 10419:

Sakai K, Tsutsui K, Yamashita T, Iwabe N, Takahashi K, Wada A, Shichida Y (August 2017).

9587:

Feuda R, Goulty M, Zadra N, Gasparetti T, Rosato E, Pisani D, et al. (August 2021).

8802:"Distinct roles of TRP channels in auditory transduction and amplification in Drosophila" 8073:"The Toll and Imd pathways are the major regulators of the immune response in Drosophila" 5018: 3913: 3679: 3512:

Systematische Beschreibung der bekannten europäischen zweiflügeligen Insekten. (Volume 6)

3217: 2994:. These opsins still have the lysine like other opsins. However, if it is replaced by an 2738: 2675: 1928: 1826: 1314:

in a heterozygous state without recombination due to multiple inversions in the balancer.

1272: 1184: 924: 436: 12376: 11789: 11732: 11611: 11508: 11451: 10995: 10761: 10701: 10603: 10501: 10436: 10381: 10327: 9871: 9529: 9467: 9305: 9194: 8936: 8866: 8723: 7860: 7673: 7492: 7347: 7302: 7098: 6886: 6553: 6430: 6084: 5805: 5584:

Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences

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While certain effects of developmental temperature, like body size, are irreversible in

1912:

and fall into the centre of the embryo (yolk sac), which will not form part of the fly.

1317:

The development of this organism—from fertilized egg to mature adult—is well understood.

12323: 12298: 12249: 12175: 12150: 12108: 12095: 12066: 12042: 12017: 11983: 11943: 11906: 11857: 11830: 11806: 11773: 11749: 11716: 11697: 11633: 11528: 11370: 11343: 11278: 11251: 11232: 11189: 11141: 11114: 11113:

Seeds AM, Ravbar P, Chung P, Hampel S, Midgley FM, Mensh BD, Simpson JH (August 2014).

11092: 11052: 11017: 10919: 10872: 10826: 10778: 10745: 10723: 10661: 10634: 10615: 10572: 10531: 10518: 10485: 10466: 10453: 10420: 10401: 10347: 10296: 10253: 10240: 10231: 10215: 10191: 10182: 10166: 10140: 10086: 10073: 10064: 10048: 10029: 9986: 9948: 9935: 9910: 9891: 9840: 9827: 9818: 9802: 9770: 9757: 9732: 9675: 9662: 9637: 9613: 9588: 9559: 9546: 9513: 9487: 9431: 9404: 9375: 9348: 9324: 9289: 9260: 9235: 9211: 9178: 9109: 9082: 9053: 9028: 9004: 8979: 8955: 8920: 8885: 8850: 8826: 8801: 8664: 8637: 8613: 8588: 8440: 8413: 8389: 8362: 8338: 8311: 8287: 8260: 8233: 8208: 8189: 8048: 8021: 7926: 7899: 7880: 7824: 7799: 7739: 7726: 7709: 7638: 7625: 7608: 7461: 7401: 7267: 7219: 7206: 7189: 7170: 7117: 7082: 7020: 6995: 6971: 6946: 6895: 6870: 6734: 6705: 6642: 6617: 6449: 6414: 6387: 6360: 6336: 6311: 6246: 6219: 6200: 6152: 6127: 6103: 6068: 5979: 5955:"An Eye on Trafficking Genes: Identification of Four Eye Color Mutations in Drosophila" 5954: 5953:

Grant P, Maga T, Loshakov A, Singhal R, Wali A, Nwankwo J, et al. (October 2016).

5925: 5900: 5817: 5769: 5744: 5725: 5604: 5579: 5485: 5321: 5254: 5155: 5120: 5035: 4969: 4944: 4920: 4885: 4826: 4799: 4752: 4622: 4614: 4574: 4549: 4525: 4479: 4438: 4267: 4246: 4168: 4052: 4019: 3887: 3860: 3670: 3647: 3622: 3213: 2915: 2895: 2832: 1859:

The life cycle of this insect has four stages: fertilized egg, larva, pupa, and adult.

1497: 1469: 1406: 1277: 1180: 1086: 912: 645:

small, pale, without dorsodistal depression and with 12-13 peg-like outer ovisensilla.

481: 238: 56: 11594:

Chiang AS, Lin CY, Chuang CC, Chang HM, Hsieh CH, Yeh CW, et al. (January 2011).

11048: 11004: 10979: 10891: 10284: 10127: 10110: 9709: 8097: 8072: 7500: 7430:

David JR, Araripe LO, Chakir M, Legout H, Lemos B, Pétavy G, et al. (July 2005).

7355: 7059: 6797: 6772: 6678: 6572: 6537: 6359:

Ferreiro MJ, Pérez C, Marchesano M, Ruiz S, Caputi A, Aguilera P, et al. (2018).

6020: 6003: 4884:

Carnes MU, Campbell T, Huang W, Butler DG, Carbone MA, Duncan LH, et al. (2015).

4778:"Fruit fly research may reveal what happens in female brains during courtship, mating" 4550:"Frequent replenishment sustains the beneficial microbiome of Drosophila melanogaster" 4079: 2842: 1142:

chromosome. The distance between the genes (map units) are equal to the percentage of

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will also attempt to court females that are not yet sexually mature, and other males.

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Females become receptive to courting males about 8–12 hours after emergence. Specific

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that encodes a protein with essential roles in repair of DNA damage. Mutations in the

13046: 12867: 12685: 12328: 12283: 12276: 12260: 12180: 12112: 12100: 12047: 11987: 11948: 11907:"A Complete Electron Microscopy Volume of the Brain of Adult Drosophila melanogaster" 11862: 11811: 11754: 11689: 11625: 11576: 11520: 11465: 11416: 11375: 11324: 11283: 11224: 11181: 11146: 11088: 11009: 10960: 10923: 10911: 10864: 10818: 10783: 10727: 10715: 10666: 10523: 10458: 10393: 10339: 10288: 10245: 10196: 10132: 10078: 10021: 9940: 9883: 9832: 9762: 9713: 9667: 9618: 9563: 9551: 9479: 9436: 9380: 9329: 9265: 9236:"Hierarchical chemosensory regulation of male-male social interactions in Drosophila" 9216: 9152: 9142: 9114: 9058: 9009: 8960: 8890: 8831: 8758: 8735: 8669: 8618: 8569: 8528: 8487: 8445: 8394: 8343: 8292: 8238: 8193: 8181: 8146: 8102: 8053: 8002: 7958: 7951: 7931: 7872: 7829: 7780: 7731: 7687: 7630: 7589: 7545: 7504: 7453: 7448: 7431: 7409: 7359: 7316: 7259: 7255: 7211: 7162: 7122: 7063: 7025: 6976: 6925: 6900: 6851: 6846: 6821: 6802: 6739: 6682: 6647: 6577: 6515: 6454: 6392: 6341: 6292: 6251: 6192: 6157: 6108: 6025: 5984: 5930: 5878: 5825: 5774: 5717: 5654: 5609: 5560: 5477: 5429: 5369: 5365: 5332: 5298: 5246: 5160: 5121:"Sexual experience enhances Drosophila melanogaster male mating behavior and success" 4974: 4925: 4866: 4831: 4798:

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e: Ebony; black body and wings (heterozygotes are also visibly darker than wild type)

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in fruit flies helps in understanding these processes in other eukaryotes, including

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Carvalho AB (December 2002). "Origin and evolution of the Drosophila Y chromosome".

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cell death signalling are all involved in key physiological responses to infection.

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are known to exhibit grooming behaviors that are executed in a predictable manner.

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males are known to spend significant time courting interspecifically, such as with

812: 583:. The appearance and sex of fruit flies is determined only by genetic information. 12513: 8636:

Parvy JP, Yu Y, Dostalova A, Kondo S, Kurjan A, Bulet P, et al. (July 2019).

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American Journal of Physiology. Regulatory, Integrative and Comparative Physiology

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Harrison JF, Waters JS, Biddulph TA, Kovacevic A, Klok CJ, Socha JJ (April 2018).

5425: 5071: 5054: 4700: 2371:, causing a truncated form to be produced. In the female version, the presence of 1803:

is available to search for human disease gene homologues in flies and vice versa.

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Hotta Y, Benzer S (December 1972). "Mapping of behaviour in Drosophila mosaics".

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is attracted to fruit that is already rotting, rather than causing fruit to rot.

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Top view of a walking Drosophila (left) with legs tracked with DeepLabCut (right)

2831:, and are one of the most advanced among insects. Each ommatidium contains eight 2734: 2492: 2448: 2070: 2048: 2010: 1814: 1772: 1255: 1170: 1123: 842: 377: 13002: 10486:"A rhodopsin in the brain functions in circadian photoentrainment in Drosophila" 9044: 8483: 6327: 4960: 4506: 3995: 1673: 1653: 1161:, and today it is one of the most widely used and genetically best-known of all 12932: 12919: 12736: 12633: 12033: 11933: 10907: 10444: 9574: 9537: 9512:

Sharkey CR, Blanco J, Leibowitz MM, Pinto-Benito D, Wardill TJ (October 2020).

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