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Polycomb-group proteins

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Polycomb-group proteins ( PcG proteins ) are a family of protein complexes first discovered in fruit flies that can remodel chromatin such that epigenetic silencing of genes takes place. Polycomb-group proteins are well known for silencing Hox genes through modulation of chromatin structure during embryonic development in fruit flies ( Drosophila melanogaster ). They derive their name from the fact that the first sign of a decrease in PcG function is often a homeotic transformation of posterior legs towards anterior legs, which have a characteristic comb-like set of bristles.

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117-908: In Drosophila , the Trithorax-group (trxG) and Polycomb-group (PcG) proteins act antagonistically and interact with chromosomal elements, termed Cellular Memory Modules (CMMs). Trithorax-group (trxG) proteins maintain the active state of gene expression while the Polycomb-group (PcG) proteins counteract this activation with a repressive function that is stable over many cell generations and can only be overcome by germline differentiation processes. Polycomb Gene complexes or PcG silencing consist of at least three kinds of multiprotein complex Polycomb Repressive Complex 1 (PRC1), PRC2 and PhoRC . These complexes work together to carry out their repressive effect. PcGs proteins are evolutionarily conserved and exist in at least two separate protein complexes;

234-429: A helical structure (i.e., shaped like a corkscrew). Their double-helix model had two strands of DNA with the nucleotides pointing inward, each matching a complementary nucleotide on the other strand to form what look like rungs on a twisted ladder. This structure showed that genetic information exists in the sequence of nucleotides on each strand of DNA. The structure also suggested a simple method for replication : if

351-534: A sexual process for transferring DNA from one cell to another cell (usually of the same species). Transformation requires the action of numerous bacterial gene products , and its primary adaptive function appears to be repair of DNA damages in the recipient cell. The diploid nature of chromosomes allows for genes on different chromosomes to assort independently or be separated from their homologous pair during sexual reproduction wherein haploid gametes are formed. In this way new combinations of genes can occur in

468-686: A white eyed mutant in 1910 to the academic community. He was in search of a model organism to study genetic heredity and required a species that could randomly acquire genetic mutation that would visibly manifest as morphological changes in the adult animal. His work on Drosophila earned him the 1933 Nobel Prize in Medicine for identifying chromosomes as the vector of inheritance for genes. This and other Drosophila species are widely used in studies of genetics , embryogenesis , chronobiology , speciation , neurobiology , and other areas. However, some species of Drosophila are difficult to culture in

585-582: A DNA molecule. In 1983, Kary Banks Mullis developed the polymerase chain reaction , providing a quick way to isolate and amplify a specific section of DNA from a mixture. The efforts of the Human Genome Project , Department of Energy, NIH, and parallel private efforts by Celera Genomics led to the sequencing of the human genome in 2003. At its most fundamental level, inheritance in organisms occurs by passing discrete heritable units, called genes , from parents to offspring. This property

702-493: A complex trait is called heritability . Measurement of the heritability of a trait is relative—in a more variable environment, the environment has a bigger influence on the total variation of the trait. For example, human height is a trait with complex causes. It has a heritability of 89% in the United States. In Nigeria, however, where people experience a more variable access to good nutrition and health care , height has

819-483: A different parent. Many species have so-called sex chromosomes that determine the sex of each organism. In humans and many other animals, the Y chromosome contains the gene that triggers the development of the specifically male characteristics. In evolution, this chromosome has lost most of its content and also most of its genes, while the X chromosome is similar to the other chromosomes and contains many genes. This being said, Mary Frances Lyon discovered that there

936-546: A diploid cell with paired chromosomes. Diploid organisms form haploids by dividing, without replicating their DNA, to create daughter cells that randomly inherit one of each pair of chromosomes. Most animals and many plants are diploid for most of their lifespan, with the haploid form reduced to single cell gametes such as sperm or eggs . Although they do not use the haploid/diploid method of sexual reproduction, bacteria have many methods of acquiring new genetic information. Some bacteria can undergo conjugation , transferring

1053-429: A gene is used to produce a specific amino acid sequence . This process begins with the production of an RNA molecule with a sequence matching the gene's DNA sequence, a process called transcription . This messenger RNA molecule then serves to produce a corresponding amino acid sequence through a process called translation . Each group of three nucleotides in the sequence, called a codon , corresponds either to one of

1170-482: A heritability of only 62%. The molecular basis for genes is deoxyribonucleic acid (DNA). DNA is composed of deoxyribose (sugar molecule), a phosphate group, and a base (amine group). There are four types of bases: adenine (A), cytosine (C), guanine (G), and thymine (T). The phosphates make phosphodiester bonds with the sugars to make long phosphate-sugar backbones. Bases specifically pair together (T&A, C&G) between two backbones and make like rungs on

1287-573: 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. The larvae of at least one species, D. suzukii , can also feed in fresh fruit and can sometimes be a pest. A few species have switched to being parasites or predators . Many species can be attracted to baits of fermented bananas or mushrooms, but others are not attracted to any kind of baits. Males may congregate at patches of suitable breeding substrate to compete for

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1404-435: A higher body temperature. In a low-temperature environment, however, the protein's structure is stable and produces dark-hair pigment normally. The protein remains functional in areas of skin that are colder—such as its legs, ears, tail, and face—so the cat has dark hair at its extremities. Environment plays a major role in effects of the human genetic disease phenylketonuria . The mutation that causes phenylketonuria disrupts

1521-506: A ladder. The bases, phosphates, and sugars together make a nucleotide that connects to make long chains of DNA. Genetic information exists in the sequence of these nucleotides, and genes exist as stretches of sequence along the DNA chain. These chains coil into a double a-helix structure and wrap around proteins called Histones which provide the structural support. DNA wrapped around these histones are called chromosomes. Viruses sometimes use

1638-505: A living cell or organism may increase or decrease gene transcription. A classic example is two seeds of genetically identical corn, one placed in a temperate climate and one in an arid climate (lacking sufficient waterfall or rain). While the average height the two corn stalks could grow to is genetically determined, the one in the arid climate only grows to half the height of the one in the temperate climate due to lack of water and nutrients in its environment. The word genetics stems from

1755-477: A matter of putting in a small piece of the natural host to receive the eggs. The Drosophila Species Stock Center located at Cornell University in Ithaca , New York, maintains cultures of hundreds of species for researchers. Drosophila is considered one of the most valuable genetic model organisms; both adults and embryos are experimental models. Drosophila is a prime candidate for genetic research because

1872-456: A phenotype involves studying identical and fraternal twins , or other siblings of multiple births . Identical siblings are genetically the same since they come from the same zygote. Meanwhile, fraternal twins are as genetically different from one another as normal siblings. By comparing how often a certain disorder occurs in a pair of identical twins to how often it occurs in a pair of fraternal twins, scientists can determine whether that disorder

1989-403: A series of genes can be combined to form a linear linkage map that roughly describes the arrangement of the genes along the chromosome. Genes express their functional effect through the production of proteins, which are molecules responsible for most functions in the cell. Proteins are made up of one or more polypeptide chains, each composed of a sequence of amino acids . The DNA sequence of

2106-457: A single parent. Offspring that are genetically identical to their parents are called clones . Eukaryotic organisms often use sexual reproduction to generate offspring that contain a mixture of genetic material inherited from two different parents. The process of sexual reproduction alternates between forms that contain single copies of the genome ( haploid ) and double copies ( diploid ). Haploid cells fuse and combine genetic material to create

2223-448: A small circular piece of DNA to another bacterium. Bacteria can also take up raw DNA fragments found in the environment and integrate them into their genomes, a phenomenon known as transformation . These processes result in horizontal gene transfer , transmitting fragments of genetic information between organisms that would be otherwise unrelated. Natural bacterial transformation occurs in many bacterial species, and can be regarded as

2340-400: A smooth blend of traits from their parents. Mendel's work provided examples where traits were definitely not blended after hybridization, showing that traits are produced by combinations of distinct genes rather than a continuous blend. Blending of traits in the progeny is now explained by the action of multiple genes with quantitative effects . Another theory that had some support at that time

2457-419: A time, so that they can be laid together on one site. Others that breed in more-abundant but less nutritious substrates, such as leaves, may only lay one egg per day. The eggs have one or more respiratory filaments near the anterior end; the tips of these extend above the surface and allow oxygen to reach the embryo. Larvae feed not on the vegetable matter itself, but on the yeasts and microorganisms present on

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2574-452: A transition of heredity from its status as myth to that of a scientific discipline, by providing a fundamental theoretical basis for genetics in the twentieth century. Other theories of inheritance preceded Mendel's work. A popular theory during the 19th century, and implied by Charles Darwin 's 1859 On the Origin of Species , was blending inheritance : the idea that individuals inherit

2691-412: A unique three-dimensional structure for that protein, and the three-dimensional structures of proteins are related to their functions. Some are simple structural molecules, like the fibers formed by the protein collagen . Proteins can bind to other proteins and simple molecules, sometimes acting as enzymes by facilitating chemical reactions within the bound molecules (without changing the structure of

2808-400: A variety of hereditary characteristics that replicate and remain active throughout generations. While haploid organisms have only one copy of each chromosome, most animals and many plants are diploid , containing two of each chromosome and thus two copies of every gene. The two alleles for a gene are located on identical loci of the two homologous chromosomes , each allele inherited from

2925-488: Is X-chromosome inactivation during reproduction to avoid passing on twice as many genes to the offspring. Lyon's discovery led to the discovery of X-linked diseases. When cells divide, their full genome is copied and each daughter cell inherits one copy. This process, called mitosis , is the simplest form of reproduction and is the basis for asexual reproduction. Asexual reproduction can also occur in multicellular organisms, producing offspring that inherit their genome from

3042-599: Is a genus of fly , belonging to the family Drosophilidae , whose members are often called "small fruit flies" or pomace flies, vinegar flies, or wine flies, a reference to the characteristic of many species to linger around overripe or rotting fruit. They should not be confused with the Tephritidae , a related family, which are also called fruit flies (sometimes referred to as "true fruit flies"); tephritids feed primarily on unripe or ripe fruit , with many species being regarded as destructive agricultural pests, especially

3159-498: Is a connection between the number of time females choose to mate and chromosomal variants of the third chromosome. It is believed that the presence of the inverted polymorphism is why re-mating by females occurs. The stability of these polymorphisms may be related to the sex-ratio meiotic drive. However, for Drosophila subobscura , the main mating system is monandry, not normally seen in Drosophila. The following section

3276-462: Is an accepted version of this page Genetics is the study of genes , genetic variation , and heredity in organisms . It is an important branch in biology because heredity is vital to organisms' evolution . Gregor Mendel , a Moravian Augustinian friar working in the 19th century in Brno , was the first to study genetics scientifically. Mendel studied "trait inheritance", patterns in

3393-438: Is an attractive behaviour. Females respond via their perception of the behavior portrayed by the male. Male and female Drosophila use a variety of sensory cues to initiate and assess courtship readiness of a potential mate. 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

3510-417: Is based on the following Drosophila species: Drosophila serrata , Drosophila pseudoobscura , Drosophila melanogaster , and Drosophila neotestacea . Polyandry is a prominent mating system among Drosophila . Females mating with multiple sex partners has been a beneficial mating strategy for Drosophila . The benefits include both pre and post copulatory mating. Pre-copulatory strategies are

3627-494: Is based on the following Drosophila species: Drosophila melanogaster , Drosophila simulans , and Drosophila mauritiana . Sperm competition is a process that polyandrous Drosophila females use to increase the fitness of their offspring. The female Drosophila has two sperm storage organs, the spermathecae and seminal receptacle, that allows her to choose the sperm that will be used to inseminate her eggs. However, some species of Drosophila have evolved to only use one or

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3744-482: Is caused by genetic or postnatal environmental factors. One famous example involved the study of the Genain quadruplets , who were identical quadruplets all diagnosed with schizophrenia . The genome of a given organism contains thousands of genes, but not all these genes need to be active at any given moment. A gene is expressed when it is being transcribed into mRNA and there exist many cellular methods of controlling

3861-407: Is explored as a mechanism of controlling viral diseases ( e.g. Dengue fever) by transferring these Wolbachia to disease-vector mosquitoes. The S. poulsonii strain of Drosophila neotestacea protects its host from parasitic wasps and nematodes using toxins that preferentially attack the parasites instead of the host. Since the Drosophila species is one of the most used model organisms, it

3978-505: Is inactivated in the young embryo . The Polycomb gene FIE is expressed in unfertilised egg cells of the moss Physcomitrella patens and expression ceases after fertilisation in the developing diploid sporophyte. It has been shown that unlike in mammals the PcG are necessary to keep the cells in a differentiated state. Consequently, loss of PcG causes de-differentiation and promotes embryonic development. Polycomb-group proteins also intervene in

4095-453: Is known to regulate ink4 locus (p16, p19). Regulation of Polycomb-group proteins at bivalent chromatin sites is performed by SWI/SNF complexes, which oppose the accumulation of Polycomb complexes through ATP-dependent eviction. In Physcomitrella patens the PcG protein FIE is specifically expressed in stem cells such as the unfertilized egg cell . Soon after fertilisation the FIE gene

4212-557: Is one step in the immune responses to some pathogens. Dudzic et al. 2019 additionally find a large number of shared serine protease messengers between Spätzle/Toll and melanization and a large amount of crosstalk between these pathways.   immigrans-tripunctata  radiation   D. quadrilineata  species group   Samoaia   Zaprionus   D. tumiditarsus  species group   Liodrosophila   Dichaetophora   Hirtodrosophila Genetics This

4329-441: Is responsible for the development of structures within multicellular organisms, these patterns arise from the complex interactions between many cells. Within eukaryotes , there exist structural features of chromatin that influence the transcription of genes, often in the form of modifications to DNA and chromatin that are stably inherited by daughter cells. These features are called " epigenetic " because they exist "on top" of

4446-705: Is seen during courtship and mating . Drosophila species are prey for many generalist predators, such as robber flies . In Hawaii , the introduction of yellowjackets from mainland United States has led to the decline of many of the larger species. The larvae are preyed on by other fly larvae, staphylinid beetles , and ants . Fruit flies use several fast-acting neurotransmitters, similar to those found in humans, which allow neurons to communicate and coordinate behavior. Acetylcholine, glutamate, gamma-aminobutyric acid (GABA), dopamine, serotonin, and histamine are all neurotransmitters that can be found in humans, but Drosophila also have another neurotransmitter, octopamine,

4563-497: Is the physical basis for inheritance: DNA replication duplicates the genetic information by splitting the strands and using each strand as a template for synthesis of a new partner strand. Genes are arranged linearly along long chains of DNA base-pair sequences. In bacteria , each cell usually contains a single circular genophore , while eukaryotic organisms (such as plants and animals) have their DNA arranged in multiple linear chromosomes. These DNA strands are often extremely long;

4680-694: Is the same as that which Mendel published. In his third law, he developed the basic principles of mutation (he can be considered a forerunner of Hugo de Vries ). Festetics argued that changes observed in the generation of farm animals, plants, and humans are the result of scientific laws. Festetics empirically deduced that organisms inherit their characteristics, not acquire them. He recognized recessive traits and inherent variation by postulating that traits of past generations could reappear later, and organisms could produce progeny with different attributes. These observations represent an important prelude to Mendel's theory of particulate inheritance insofar as it features

4797-407: Is very diverse in appearance, behavior, and breeding habitat. The term " Drosophila ", meaning "dew-loving", is a modern scientific Latin adaptation from Greek words δρόσος , drósos , " dew ", and φιλία , philía , "lover". Drosophila species are small flies, typically pale yellow to reddish brown to black, with red eyes. When the eyes (essentially a film of lenses) are removed,

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4914-486: The Blue-eyed Mary ( Omphalodes verna ), for example, there exists a gene with alleles that determine the color of flowers: blue or magenta. Another gene, however, controls whether the flowers have color at all or are white. When a plant has two copies of this white allele, its flowers are white—regardless of whether the first gene has blue or magenta alleles. This interaction between genes is called epistasis , with

5031-497: The D. melanogaster Spiroplasma poulsonii (named MSRO). The male-killing factor of the D. melanogaster MSRO strain was discovered in 2018, solving a decades-old mystery of the cause of male-killing. This represents the first bacterial factor that affects eukaryotic cells in a sex-specific fashion, and is the first mechanism identified for male-killing phenotypes. Alternatively, they may protect theirs hosts from infection. Drosophila Wolbachia can reduce viral loads upon infection, and

5148-608: The Drosophila DNA damage response, including deficiencies in expression of genes involved in DNA damage repair , accelerates intestinal stem cell ( enterocyte ) aging. Sharpless and Depinho reviewed evidence that stem cells undergo intrinsic aging and speculated that stem cells grow old, in part, as a result of DNA damage. The following section is based on the following Drosophila species: Drosophila simulans and Drosophila melanogaster . Courtship behavior of male Drosophila

5265-465: The Mediterranean fruit fly . One species of Drosophila in particular, Drosophila melanogaster , has been heavily used in research in genetics and is a common model organism in developmental biology . The terms "fruit fly" and " Drosophila " are often used synonymously with D. melanogaster in modern biological literature. The entire genus, however, contains more than 1,500 species and

5382-413: The ancient Greek γενετικός genetikos meaning "genitive"/"generative", which in turn derives from γένεσις genesis meaning "origin". The observation that living things inherit traits from their parents has been used since prehistoric times to improve crop plants and animals through selective breeding . The modern science of genetics, seeking to understand this process, began with

5499-494: The fru mutation leads back to the typical courtship behavior. A novel class of pheromones was found to be conserved across the subgenus Drosophila in 11 desert dwelling species. 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. The following section

5616-415: The neutral theory of molecular evolution through publishing the nearly neutral theory of molecular evolution . In this theory, Ohta stressed the importance of natural selection and the environment to the rate at which genetic evolution occurs. One important development was chain-termination DNA sequencing in 1977 by Frederick Sanger . This technology allows scientists to read the nucleotide sequence of

5733-421: The DNA sequence and retain inheritance from one cell generation to the next. Because of epigenetic features, different cell types grown within the same medium can retain very different properties. Although epigenetic features are generally dynamic over the course of development, some, like the phenomenon of paramutation , have multigenerational inheritance and exist as rare exceptions to the general rule of DNA as

5850-632: The F1 offspring mate with each other, the offspring are called the "F2" (second filial) generation. One of the common diagrams used to predict the result of cross-breeding is the Punnett square . When studying human genetic diseases, geneticists often use pedigree charts to represent the inheritance of traits. These charts map the inheritance of a trait in a family tree. Organisms have thousands of genes, and in sexually reproducing organisms these genes generally assort independently of each other. This means that

5967-504: The Greek word genesis —γένεσις, "origin", predates the noun and was first used in a biological sense in 1860. Bateson both acted as a mentor and was aided significantly by the work of other scientists from Newnham College at Cambridge, specifically the work of Becky Saunders , Nora Darwin Barlow , and Muriel Wheldale Onslow . Bateson popularized the usage of the word genetics to describe

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6084-428: The PcG repressive complex 1 (PRC1) and the PcG repressive complex 2–4 (PRC2/3/4). PRC2 catalyzes trimethylation of lysine 27 on histone H3 (H3K27me2/3), while PRC1 mono- ubiquitinates histone H2A on lysine 119 (H2AK119Ub1). In mammals Polycomb Group gene expression is important in many aspects of development like homeotic gene regulation and X chromosome inactivation , being recruited to the inactive X by Xist RNA ,

6201-448: The ability of the body to break down the amino acid phenylalanine , causing a toxic build-up of an intermediate molecule that, in turn, causes severe symptoms of progressive intellectual disability and seizures. However, if someone with the phenylketonuria mutation follows a strict diet that avoids this amino acid, they remain normal and healthy. A common method for determining how genes and environment ("nature and nurture") contribute to

6318-444: The adaptive function of repair of DNA damages. The first cytological demonstration of crossing over was performed by Harriet Creighton and Barbara McClintock in 1931. Their research and experiments on corn provided cytological evidence for the genetic theory that linked genes on paired chromosomes do in fact exchange places from one homolog to the other. The probability of chromosomal crossover occurring between two given points on

6435-546: The analog of norepinephrine. Acetylcholine is the primary excitatory neurotransmitter and GABA is the primary inhibitory neurotransmitter utilized in the drosophila central nervous system. In Drosophila, the effects of many neurotransmitters can vary depending on the receptors and signaling pathways involved, allowing them to act as excitatory or inhibitory signals under different contexts. This versatility enables complex neural processing and behavioral flexibility. Glutamate can serve as an excitatory neurotransmitter, specifically at

6552-440: The basis for inheritance. During the process of DNA replication, errors occasionally occur in the polymerization of the second strand. These errors, called mutations, can affect the phenotype of an organism, especially if they occur within the protein coding sequence of a gene. Error rates are usually very low—1 error in every 10–100 million bases—due to the "proofreading" ability of DNA polymerases . Processes that increase

6669-501: The behaviours associated with mate choice and the genetic contributions, such as production of gametes, that are exhibited by both male and female Drosophila regarding mate choice. Post copulatory strategies include sperm competition, mating frequency, and sex-ratio meiotic drive. These lists are not inclusive. Polyandry among the Drosophila pseudoobscura in North America vary in their number of mating partners. There

6786-434: The brain is revealed. Drosophila brain structure and function develop and age significantly from larval to adult stage. Developing brain structures make these flies a prime candidate for neuro-genetic research. 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

6903-404: The cat plays the role of the environment. The cat's genes code for dark hair, thus the hair-producing cells in the cat make cellular proteins resulting in dark hair. But these dark hair-producing proteins are sensitive to temperature (i.e. have a mutation causing temperature-sensitivity) and denature in higher-temperature environments, failing to produce dark-hair pigment in areas where the cat has

7020-429: The cell, these genes for tryptophan synthesis are no longer needed. The presence of tryptophan directly affects the activity of the genes—tryptophan molecules bind to the tryptophan repressor (a transcription factor), changing the repressor's structure such that the repressor binds to the genes. The tryptophan repressor blocks the transcription and expression of the genes, thereby creating negative feedback regulation of

7137-421: The chromosome is related to the distance between the points. For an arbitrarily long distance, the probability of crossover is high enough that the inheritance of the genes is effectively uncorrelated. For genes that are closer together, however, the lower probability of crossover means that the genes demonstrate genetic linkage; alleles for the two genes tend to be inherited together. The amounts of linkage between

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7254-422: The combined DNA sequences of all chromosomes) is called the genome . DNA is most often found in the nucleus of cells, but Ruth Sager helped in the discovery of nonchromosomal genes found outside of the nucleus. In plants, these are often found in the chloroplasts and in other organisms, in the mitochondria. These nonchromosomal genes can still be passed on by either partner in sexual reproduction and they control

7371-504: The components of the SNARE complex are known to be somewhat substitutable: Although the loss of SNAP-25 - a component of neuronal SNAREs - is lethal, SNAP-24 can fully replace it. For another example, an R-SNARE not normally found in synapses can substitute for synaptobrevin . The Spätzle protein is a ligand of Toll . In addition to melanin 's more commonly known roles in the endoskeleton and in neurochemistry , melanization

7488-472: The control of flowering by silencing the Flowering Locus C gene. This gene is a central part of the pathway that inhibits flowering in plants and its silencing during winter is suspected to be one of the main factors intervening in plant vernalization . Drosophila Oinopota Kirby & Spence, 1815 Drosophila ( / d r ə ˈ s ɒ f ɪ l ə , d r ɒ -, d r oʊ -/ )

7605-475: The decaying breeding substrate. Development time varies widely between species (between 7 and more than 60 days) and depends on the environmental factors such as temperature , breeding substrate, and crowding. Fruit flies lay eggs in response to environmental cycles. Eggs laid at a time (e.g., night) during which likelihood of survival is greater than in eggs laid at other times (e.g., day) yield more larvae than eggs that were laid at those times. Ceteris paribus ,

7722-417: The expression of genes such that proteins are produced only when needed by the cell. Transcription factors are regulatory proteins that bind to DNA, either promoting or inhibiting the transcription of a gene. Within the genome of Escherichia coli bacteria, for example, there exists a series of genes necessary for the synthesis of the amino acid tryptophan . However, when tryptophan is already available to

7839-561: 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 a house fly . Environmental challenge by natural toxins helped to prepare Drosophila e to detox DDT , by shaping the glutathione S -transferase mechanism that metabolizes both. The Drosophila genome is subject to a high degree of selection, especially unusually widespread negative selection compared to other taxa . A majority of

7956-421: The females, or form leks , conducting courtship in an area separate from breeding sites. Several Drosophila species, including Drosophila melanogaster , D. immigrans , and D. simulans , are closely associated with humans, and are often referred to as domestic species. These and other species ( D. subobscura , and from a related genus Zaprionus indianus ) have been accidentally introduced around

8073-531: The findings can be applied to deduce genetic trends in humans. Research conducted on Drosophila help determine the ground rules for transmission of genes in many organisms. Drosophila is a useful in vivo tool to analyze Alzheimer's disease. Rhomboid proteases were first detected in Drosophila but then found to be highly conserved across eukaryotes , mitochondria , and bacteria . Melanin's ability to protect DNA against ionizing radiation has been most extensively demonstrated in Drosophila , including in

8190-417: The flies were more cold resistant and also had the highest concentration of Wolbachia. The microbiome in the gut can also be transplanted among organisms. It was found that Drosophila melanogaster became more cold-tolerant when the gut microbiota from Drosophila melanogaster that were reared at low temperatures. This depicted that the gut microbiome is correlated to physiological processes. Moreover,

8307-523: The following three stages: insemination, sperm storage, and fertilizable sperm. Among the preceding species there are variations at each stage that play a role in the natural selection process. This sperm competition has been found to be a driving force in the establishment of reproductive isolation during speciation. Parthenogenesis does not occur in D. melanogaster , but in the gyn-f9 mutant, gynogenesis occurs at low frequency. The natural populations of D. mangebeirai are entirely female, making it

8424-608: The formative study by Hopwood et al. 1985. Like other animals, Drosophila is associated with various bacteria in its gut. The fly gut microbiota or microbiome seems to have a central influence on Drosophila fitness and life history characteristics. The microbiota in the gut of Drosophila represents an active current research field. Drosophila species also harbour vertically transmitted endosymbionts, such as Wolbachia and Spiroplasma . These endosymbionts can act as reproductive manipulators, such as cytoplasmic incompatibility induced by Wolbachia or male-killing induced by

8541-669: The function and behavior of genes. Gene structure and function, variation, and distribution are studied within the context of the cell , the organism (e.g. dominance ), and within the context of a population. Genetics has given rise to a number of subfields, including molecular genetics , epigenetics , and population genetics . Organisms studied within the broad field span the domains of life ( archaea , bacteria , and eukarya ). Genetic processes work in combination with an organism's environment and experiences to influence development and behavior , often referred to as nature versus nurture . The intracellular or extracellular environment of

8658-467: The genome is under selection of some sort, and a supermajority of this is occurring in non-coding DNA . Effective population size has been credibly suggested to positively correlate with the effect size of both negative and positive selection . Recombination is likely to be a significant source of diversity . There is evidence that crossover is positively correlated with polymorphism in D. populations. Drosophila species are found all around

8775-460: The habit of laying eggs at this 'advantageous' time would yield more surviving offspring, and more grandchildren, than the habit of laying eggs during other times. This differential reproductive success would cause D. melanogaster to adapt to environmental cycles, because this behavior has a major reproductive advantage. Their median lifespan is 35–45 days. DNA damage accumulates in Drosophila intestinal stem cells with age. Deficiencies in

8892-469: The information an organism uses to function, the environment plays an important role in determining the ultimate phenotypes an organism displays. The phrase " nature and nurture " refers to this complementary relationship. The phenotype of an organism depends on the interaction of genes and the environment. An interesting example is the coat coloration of the Siamese cat . In this case, the body temperature of

9009-406: The inheritance of an allele for yellow or green pea color is unrelated to the inheritance of alleles for white or purple flowers. This phenomenon, known as " Mendel's second law " or the "law of independent assortment," means that the alleles of different genes get shuffled between parents to form offspring with many different combinations. Different genes often interact to influence the same trait. In

9126-404: The laboratory, often because they breed on a single specific host in the wild. For some, it can be done with particular recipes for rearing media, or by introducing chemicals such as sterols that are found in the natural host; for others, it is (so far) impossible. In some cases, the larvae can develop on normal Drosophila lab medium, but the female will not lay eggs; for these it is often simply

9243-427: The largest human chromosome, for example, is about 247 million base pairs in length. The DNA of a chromosome is associated with structural proteins that organize, compact, and control access to the DNA, forming a material called chromatin ; in eukaryotes, chromatin is usually composed of nucleosomes , segments of DNA wound around cores of histone proteins. The full set of hereditary material in an organism (usually

9360-434: The longest. D. melanogaster sperm cells are a more modest 1.8 mm long, although this is still about 35 times longer than a human sperm. Several species in the D. melanogaster species group are known to mate by traumatic insemination . Drosophila species vary widely in their reproductive capacity. Those such as D. melanogaster that breed in large, relatively rare resources have ovaries that mature 10–20 eggs at

9477-505: The male and female. Recent experiments explore the role of fruitless ( fru ) and doublesex ( dsx ), a group of sex-behaviour linked genes. The fruitless ( fru ) gene in Drosophila helps regulate the network for male courtship behavior; when a mutation to this gene occurs altered same sex sexual behavior in males is observed. Male Drosophila with the fru mutation direct their courtship towards other males as opposed to typical courtship, which would be directed towards females. Loss of

9594-526: The master regulator of XCI or embryonic stem cell self-renewal. The Bmi1 polycomb ring finger protein promotes neural stem cell self-renewal. Murine null mutants in PRC2 genes are embryonic lethals while most PRC1 mutants are live born homeotic mutants that die perinatally. In contrast overexpression of PcG proteins correlates with the severity and invasiveness of several cancer types. The mammalian PRC1 core complexes are very similar to Drosophila. Polycomb Bmi1

9711-495: The microbiome plays a role in aggression, immunity, egg-laying preferences, locomotion and metabolism . As for aggression, it plays a role to a certain degree during courtship. It was observed that germ-free flies were not as competitive compared to the wild-type males. Microbiome of the Drosophila species is also known to promote aggression by octopamine OA signalling. The microbiome has been shown to impact these fruit flies' social interactions, specifically aggressive behaviour that

9828-571: The nature of inheritance in plants. In his paper " Versuche über Pflanzenhybriden " (" Experiments on Plant Hybridization "), presented in 1865 to the Naturforschender Verein (Society for Research in Nature) in Brno , Mendel traced the inheritance patterns of certain traits in pea plants and described them mathematically. Although this pattern of inheritance could only be observed for a few traits, Mendel's work suggested that heredity

9945-437: The neuromuscular junction in fruit flies. This differs from vertebrates, where acetylcholine is used at these junctions. In Drosophila, histamine primarily functions as a neurotransmitter in the visual system. It is released by photoreceptor cells to transmit visual information from the eye to the brain, making it essential for vision. As with many Eukaryotes, this genus is known to express SNAREs , and as with several others

10062-500: The offspring of a mating pair. Genes on the same chromosome would theoretically never recombine. However, they do, via the cellular process of chromosomal crossover . During crossover, chromosomes exchange stretches of DNA, effectively shuffling the gene alleles between the chromosomes. This process of chromosomal crossover generally occurs during meiosis , a series of cell divisions that creates haploid cells. Meiotic recombination , particularly in microbial eukaryotes , appears to serve

10179-412: The only obligate parthenogenetic species of Drosophila. Parthenogenesis is facultative in parthenogenetica and mercatorum . D. melanogaster is a popular experimental animal because it is easily cultured en masse out of the wild, has a short generation time, and mutant animals are readily obtainable. In 1906, Thomas Hunt Morgan began his work on D. melanogaster and reported his first finding of

10296-435: The original sequence. A particularly important source of DNA damages appears to be reactive oxygen species produced by cellular aerobic respiration , and these can lead to mutations. In organisms that use chromosomal crossover to exchange DNA and recombine genes, errors in alignment during meiosis can also cause mutations. Errors in crossover are especially likely when similar sequences cause partner chromosomes to adopt

10413-423: The other. Females have little control when it comes to cryptic female choice . Female Drosophila through cryptic choice, one of several post-copulatory mechanisms, which allows for the detection and expelling of sperm that reduces inbreeding possibilities. Manier et al. 2013 has categorized the post copulatory sexual selection of Drosophila melanogaster , Drosophila simulans , and Drosophila mauritiana into

10530-470: The phenotype of the organism, while the other allele is called recessive as its qualities recede and are not observed. Some alleles do not have complete dominance and instead have incomplete dominance by expressing an intermediate phenotype, or codominance by expressing both alleles at once. When a pair of organisms reproduce sexually , their offspring randomly inherit one of the two alleles from each parent. These observations of discrete inheritance and

10647-543: The process of protein production . It was discovered that the cell uses DNA as a template to create matching messenger RNA , molecules with nucleotides very similar to DNA. The nucleotide sequence of a messenger RNA is used to create an amino acid sequence in protein; this translation between nucleotide sequences and amino acid sequences is known as the genetic code . With the newfound molecular understanding of inheritance came an explosion of research. A notable theory arose from Tomoko Ohta in 1973 with her amendment to

10764-435: The product rule, the sum rule, and more. Geneticists use diagrams and symbols to describe inheritance. A gene is represented by one or a few letters. Often a "+" symbol is used to mark the usual, non-mutant allele for a gene. In fertilization and breeding experiments (and especially when discussing Mendel's laws) the parents are referred to as the "P" generation and the offspring as the "F1" (first filial) generation. When

10881-519: The properties of a protein by destabilizing the structure or changing the surface of the protein in a way that changes its interaction with other proteins and molecules. For example, sickle-cell anemia is a human genetic disease that results from a single base difference within the coding region for the β-globin section of hemoglobin, causing a single amino acid change that changes hemoglobin's physical properties. Sickle-cell versions of hemoglobin stick to themselves, stacking to form fibers that distort

10998-426: The protein itself). Protein structure is dynamic; the protein hemoglobin bends into slightly different forms as it facilitates the capture, transport, and release of oxygen molecules within mammalian blood. A single nucleotide difference within DNA can cause a change in the amino acid sequence of a protein. Because protein structures are the result of their amino acid sequences, some changes can dramatically change

11115-406: The rate of changes in DNA are called mutagenic : mutagenic chemicals promote errors in DNA replication, often by interfering with the structure of base-pairing, while UV radiation induces mutations by causing damage to the DNA structure. Chemical damage to DNA occurs naturally as well and cells use DNA repair mechanisms to repair mismatches and breaks. The repair does not, however, always restore

11232-686: The relationship between human and fruit fly genes is very close. 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 in Drosophila is higher than the human genome. Low and manageable number of chromosomes make Drosophila species easier to study. These flies also carry genetic information and pass down traits throughout generations, much like their human counterparts. The traits can then be studied through different Drosophila lineages and

11349-408: The same allele of a given gene are called homozygous at that gene locus , while organisms with two different alleles of a given gene are called heterozygous . The set of alleles for a given organism is called its genotype , while the observable traits of the organism are called its phenotype . When organisms are heterozygous at a gene, often one allele is called dominant as its qualities dominate

11466-401: The second gene epistatic to the first. Many traits are not discrete features (e.g. purple or white flowers) but are instead continuous features (e.g. human height and skin color ). These complex traits are products of many genes. The influence of these genes is mediated, to varying degrees, by the environment an organism has experienced. The degree to which an organism's genes contribute to

11583-509: The segregation of alleles are collectively known as Mendel's first law or the Law of Segregation. However, the probability of getting one gene over the other can change due to dominant, recessive, homozygous, or heterozygous genes. For example, Mendel found that if you cross heterozygous organisms your odds of getting the dominant trait is 3:1. Real geneticist study and calculate probabilities by using theoretical probabilities, empirical probabilities,

11700-653: The shape of red blood cells carrying the protein. These sickle-shaped cells no longer flow smoothly through blood vessels , having a tendency to clog or degrade, causing the medical problems associated with this disease. Some DNA sequences are transcribed into RNA but are not translated into protein products—such RNA molecules are called non-coding RNA . In some cases, these products fold into structures which are involved in critical cell functions (e.g. ribosomal RNA and transfer RNA ). RNA can also have regulatory effects through hybridization interactions with other RNA molecules (such as microRNA ). Although genes contain all

11817-446: The similar molecule RNA instead of DNA as their genetic material. DNA normally exists as a double-stranded molecule, coiled into the shape of a double helix . Each nucleotide in DNA preferentially pairs with its partner nucleotide on the opposite strand: A pairs with T, and C pairs with G. Thus, in its two-stranded form, each strand effectively contains all necessary information, redundant with its partner strand. This structure of DNA

11934-500: The single celled alga Acetabularia . The Hershey–Chase experiment in 1952 confirmed that DNA (rather than protein) is the genetic material of the viruses that infect bacteria, providing further evidence that DNA is the molecule responsible for inheritance. James Watson and Francis Crick determined the structure of DNA in 1953, using the X-ray crystallography work of Rosalind Franklin and Maurice Wilkins that indicated DNA has

12051-405: The stomach, attempt to copulate, and the copulatory act itself. The songs of Drosophila melanogaster and Drosophila simulans have been studied extensively. These luring songs are sinusoidal in nature and varies within and between species. The courtship behavior of Drosophila melanogaster has also been assessed for sex-related genes, which have been implicated in courtship behavior in both

12168-441: The strands are separated, new partner strands can be reconstructed for each based on the sequence of the old strand. This property is what gives DNA its semi-conservative nature where one strand of new DNA is from an original parent strand. Although the structure of DNA showed how inheritance works, it was still not known how DNA influences the behavior of cells. In the following years, scientists tried to understand how DNA controls

12285-580: The study of inheritance in his inaugural address to the Third International Conference on Plant Hybridization in London in 1906. After the rediscovery of Mendel's work, scientists tried to determine which molecules in the cell were responsible for inheritance. In 1900, Nettie Stevens began studying the mealworm. Over the next 11 years, she discovered that females only had the X chromosome and males had both X and Y chromosomes. She

12402-536: The tryptophan synthesis process. Differences in gene expression are especially clear within multicellular organisms , where cells all contain the same genome but have very different structures and behaviors due to the expression of different sets of genes. All the cells in a multicellular organism derive from a single cell, differentiating into variant cell types in response to external and intercellular signals and gradually establishing different patterns of gene expression to create different behaviors. As no single gene

12519-482: The twenty possible amino acids in a protein or an instruction to end the amino acid sequence ; this correspondence is called the genetic code . The flow of information is unidirectional: information is transferred from nucleotide sequences into the amino acid sequence of proteins, but it never transfers from protein back into the sequence of DNA—a phenomenon Francis Crick called the central dogma of molecular biology . The specific sequence of amino acids results in

12636-545: The two is responsible for inheritance. In 1928 , Frederick Griffith discovered the phenomenon of transformation : dead bacteria could transfer genetic material to "transform" other still-living bacteria. Sixteen years later, in 1944, the Avery–MacLeod–McCarty experiment identified DNA as the molecule responsible for transformation. The role of the nucleus as the repository of genetic information in eukaryotes had been established by Hämmerling in 1943 in his work on

12753-437: The way traits are handed down from parents to offspring over time. He observed that organisms (pea plants) inherit traits by way of discrete "units of inheritance". This term, still used today, is a somewhat ambiguous definition of what is referred to as a gene. Trait inheritance and molecular inheritance mechanisms of genes are still primary principles of genetics in the 21st century, but modern genetics has expanded to study

12870-538: The work of the Augustinian friar Gregor Mendel in the mid-19th century. Prior to Mendel, Imre Festetics , a Hungarian noble, who lived in Kőszeg before Mendel, was the first who used the word "genetic" in hereditarian context, and is considered the first geneticist. He described several rules of biological inheritance in his work The genetic laws of nature (Die genetischen Gesetze der Natur, 1819). His second law

12987-489: The world by human activities such as fruit transports. Males of this genus are known to have the longest sperm cells of any studied organism on Earth, including one species, Drosophila bifurca , that has sperm cells that are 58 mm (2.3 in) long. The cells mostly consist of a long, thread-like tail, and are delivered to the females in tangled coils. The other members of the genus Drosophila also make relatively few giant sperm cells, with that of D. bifurca being

13104-788: The world, with more species in the tropical regions. Drosophila made their way to the Hawaiian Islands and radiated into over 800 species. They can be found in deserts , tropical rainforest , cities , swamps , and alpine zones . Some northern species hibernate . The northern species D. montana is the best cold-adapted, and is primarily found at high altitudes. Most species breed in various kinds of decaying plant and fungal material, including fruit , bark , slime fluxes , flowers , and mushrooms . Drosophila species that are fruit-breeding are attracted to various products of fermentation, especially ethanol and methanol . Fruits exploited by Drosophila species include those with

13221-518: Was able to conclude that sex is a chromosomal factor and is determined by the male. In 1911, Thomas Hunt Morgan argued that genes are on chromosomes , based on observations of a sex-linked white eye mutation in fruit flies . In 1913, his student Alfred Sturtevant used the phenomenon of genetic linkage to show that genes are arranged linearly on the chromosome. Although genes were known to exist on chromosomes, chromosomes are composed of both protein and DNA, and scientists did not know which of

13338-591: Was first observed by Gregor Mendel, who studied the segregation of heritable traits in pea plants, showing for example that flowers on a single plant were either purple or white—but never an intermediate between the two colors. The discrete versions of the same gene controlling the inherited appearance (phenotypes) are called alleles . In the case of the pea, which is a diploid species, each individual plant has two copies of each gene, one copy inherited from each parent. Many species, including humans, have this pattern of inheritance. Diploid organisms with two copies of

13455-421: Was particulate, not acquired, and that the inheritance patterns of many traits could be explained through simple rules and ratios. The importance of Mendel's work did not gain wide understanding until 1900, after his death, when Hugo de Vries and other scientists rediscovered his research. William Bateson , a proponent of Mendel's work, coined the word genetics in 1905. The adjective genetic , derived from

13572-529: Was the inheritance of acquired characteristics : the belief that individuals inherit traits strengthened by their parents. This theory (commonly associated with Jean-Baptiste Lamarck ) is now known to be wrong—the experiences of individuals do not affect the genes they pass to their children. Other theories included Darwin's pangenesis (which had both acquired and inherited aspects) and Francis Galton 's reformulation of pangenesis as both particulate and inherited. Modern genetics started with Mendel's studies of

13689-480: Was vastly used in genetics. However, the effect abiotic factors , such as temperature, has on the microbiome on Drosophila species has recently been of great interest. Certain variations in temperature have an impact on the microbiome. It was observed that higher temperatures (31 °C) lead to an increase of Acetobacter populations in the gut microbiome of Drosophila melanogaster as compared to lower temperatures (13 °C). In low temperatures (13 °C),

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