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20-September-2008 09:29:09 - Mitochondrial DNA Mitochondrial DNA. Mitochondrial DNA. Mitochondrial DNA mtDNA is the DNA located in organelles called mitochondria. Most other DNA present in eukaryotic organisms is found in the cell nucleus. Mitochondrial DNA was discovered by Margit M. K. Nass and Sylvan Nass by electron microscopy as DNAase-sensitive thread inside mitochondria1, and by Ellen Haslbrunner, Hans Tuppy and Gottfried Schatz by biochemical assays on highly purified mitochondrial fractions2. Nuclear and mitochondrial DNA are thought to be of separate evolutionary origin, with the mtDNA being derived from the circular genomes of the bacteria that were engulfed by the early ancestors of today's eukaryotic cells. Each mitochondrion is estimated to contain 2-10 mtDNA copies.3 In the cells of extant organisms, the vast majority of the proteins present in the mitochondria numbering approximately 1500 different types in mammals are coded for by nuclear DNA, but the genes for some of them, if not most, are thought to have originally been of bacterial origin, having since been transferred to the eukaryotic nucleus during evolution. In most multicellular organisms, mtDNA is inherited from the mother maternally inherited. Mechanisms for this include simple dilution an egg contains 100,000 to 1,000,000 mtDNA molecules, whereas a sperm contains only 100 to 1000, degradation of sperm mtDNA in the fertilized egg, and, at least in a few organisms, failure of sperm mtDNA to enter the egg. Whatever the mechanism, this single parent uniparental pattern of mtDNA inheritance is found in most animals, most plants and in fungi as well. mtDNA is particularly susceptible to reactive oxygen species generated by the respiratory chain due to its close proximity. Though mtDNA is packaged by proteins and harbors significant DNA repair capacity, these protective functions are less robust than those operating on nuclear DNA and therefore thought to contribute to enhanced susceptibility of mtDNA to oxidative damage. Mutations in mtDNA cause maternally inherited diseases and are thought to be a major contributor to aging and age-associated pathology. In humans and probably in metazoans in general, 100-10,000 separate copies of mtDNA are usually present per cell egg and sperm cells are exceptions. In mammals, each circular mtDNA molecule consists of 15,000-17,000 base pairs, which encode the same 37 genes: 13 for proteins polypeptides, 22 for transfer RNA tRNA and one each for the small and large subunits of ribosomal RNA rRNA. This pattern is also seen among most metazoans, although in some cases one or more of the 37 genes is absent and the mtDNA size range is greater. Even greater variation in mtDNA gene content and size exists among fungi and plants, although there appears to be a core subset of genes that are present in all eukaryotes except for the few that have no mitochondria at all. Some plant species have enormous mtDNAs as many as 2,500,000 base pairs per mtDNA molecule but, surprisingly, even those huge mtDNAs contain the same number and kinds of genes as related plants with much smaller mtDNAs. Contents 1 Use in identification 2 Mitochondrial inheritance 2.1 Female inheritance 2.2 Male inheritance 3 Genes 4 Genetic influence 4.1 Genetic illness 5 See also 6 References 7 External links Use in identification Unlike nuclear DNA, which is inherited from both parents and in which genes are rearranged in the process of recombination, there is usually no change in mtDNA from parent to offspring. Although mtDNA also recombines, it does so with copies of itself within the same mitochondrion. Because of this and because the mutation rate of animal mtDNA is higher than that of nuclear DNA,4 mtDNA is a powerful tool for tracking ancestry through females matrilineage and has been used in this role to track the ancestry of many species back hundreds of generations. Human mtDNA can be used to identify individuals.5 Forensic laboratories occasionally use mtDNA comparison to identify human remains, and especially to identify older unidentified skeletal remains. Although unlike nuclear DNA mtDNA is not specific to one individual, it can be used in combination with other evidence anthropological evidence, circumstantial evidence, and the like to establish identification. mtDNA is also used to exclude possible matches between missing persons and unidentified remains.6 Many researchers believe that mtDNA is better suited to identification of older skeletal remains than nuclear DNA because it is often easier to harvest from older remains because of the greater number of copies of mtDNA per cell, and because a match with a living relative is possible even if numerous maternal generations separate the two. American outlaw Jesse James's remains were identified using a comparison between mtDNA extracted from his remains and the mtDNA of the son of the female-line great-granddaughter of his sister.7 Because the base sequence of animal mtDNA changes rapidly, it is useful for assessing genetic relationships of individuals or groups within a species and also for identifying and quantifying the phylogeny evolutionary relationships; see phylogenetics among different species, provided they are not too distantly related. To do this, biologists determine and then compare the mtDNA sequences from different individuals or species. Data from the comparisons is used to construct a network of relationships among the sequences, which provides an estimate of the relationships among the individuals or species from which the mtDNAs were taken. This approach has limits that are imposed by the rate of mtDNA sequence change. In animals, the rapid rate of change makes mtDNA most useful for comparisons of individuals within species and for comparisons of species that are closely or moderately-closely related, among which the number of sequence differences can be easily counted. As the species become more distantly related, the number of sequence differences becomes very large; changes begin to accumulate on changes until an accurate count becomes impossible. Mitochondrial inheritance Female inheritance In sexually reproducing organisms, mitochondria are normally inherited exclusively from the mother. The mitochondria in mammalian sperm are usually destroyed by the egg cell after fertilization. Also, most mitochondria are present at the base of the sperm's tail, which is used for propelling the sperm cells. Sometimes the tail is lost during fertilization. In 1999 it was reported that paternal sperm mitochondria containing mtDNA are marked with ubiquitin to select them for later destruction inside the embryo.8 Some in vitro fertilization techniques, particularly injecting a sperm into an oocyte, may interfere with this. The fact that mitochondrial DNA is maternally inherited enables researchers to trace maternal lineage far back in time. Y chromosomal DNA, paternally inherited, is used in an analogous way to trace the agnate lineage. This is accomplished in humans by sequencing one or more of the hypervariable control regions HVR1 or HVR2 of the mitochondrial DNA. HVR1 consists of about 440 base pairs. These 440 base pairs are then compared to the control regions of other individuals either specific people or subjects in a database to determine maternal lineage. Most often, the comparison is made to the revised. Vilà et al have published studies tracing the matrilineal descent of domestic dogs to wolves.9 The concept of the Mitochondrial Eve is based on the same type of analysis, attempting to discover the origin of humanity by tracking the lineage back in time. Because mtDNA is not highly conserved and has a rapid mutation rate, it is useful for studying the evolutionary relationships - phylogeny - of organisms. Biologists can determine and then compare mtDNA sequences among different species and use the comparisons to build an evolutionary tree for the species examined. Male inheritance It has been reported that mitochondria can occasionally be inherited from the father in some species such as mussels.1011 Paternally inherited mitochondria have also been reported in some insects such as the fruit fly12 and the honeybee.13 Evidence supports rare instances of male mitochondrial inheritance in some mammals as well. Specifically, documented occurrences exist for mice,1415 where the male-inherited mitochondria was subsequently rejected. It has also been found in sheep,16 and in cloned cattle.17 It has been found in a single case in a human male and was linked to infertility.18 While many of these cases involve cloned embryos or subsequent rejection of the paternal mitochondria, others document in vivo inheritance and persistence under lab conditions. Genes ATP synthase: MT-ATP6, MT-ATP8 cytochrome c oxidase: MT-CO1, MT-CO2, MT-CO3, MT-CYB NADH dehydrogenase: MT-ND1, MT-ND2, MT-ND3, MT-ND4, MT-ND4L, MT-ND5, MT-ND6 12S, 16S: MT-RNR1, MT-RNR2 tRNA: MT-TA, MT-TC, MT-TD, MT-TE, MT-TF, MT-TG, MT-TH, MT-TI, MT-TK, MT-TL1, MT-TL2, MT-TM, MT-TN, MT-TP, MT-TQ, MT-TR, MT-TS1, MT-TS2, MT-TT, MT-TV, MT-TW, MT-TY, MT1X Genetic influence Genetic illness Mutations of mitochondrial DNA can lead to a number of illnesses including exercise intolerance and Kearns-Sayre syndrome KSS, which causes a person to lose full function of their heart, eye, and muscle movements. See also Mitochondrial disease. See also Mitochondrial disease Human mitochondrial genetics Paternal mtDNA transmission Single origin theory Mitochondrial Eve Mitochondrial CRS References ^ Nass, M.M. Nass, S. 1963 at the Wenner-Gren Institute for Experimental Biology, Stockholm University, Stockholm, Sweden: Intramitochondrial Fibers with DNA characteristics PDF. In: J. Cell. Biol. Bd. 19, S. 593-629. PMID 14086138 ^ Ellen Haslbrunner, Hans Tuppy and Gottfried Schatz 1964 at the Institut for Biochemistry at the Medical Faculty of the University of Vienna in Vienna, Austria: Deoxyribonucleic Acid Associated with Yeast Mitochondria PDF Biochem. Biophys. Res. Commun. 15, 127 - 132. ^ Wiesner RJ, Ruegg JC, Morano I 1992. Counting target molecules by exponential polymerase chain reaction, copy number of mitochondrial DNA in rat tissues. Biochim Biophys Acta. 183: 553-559. PMID 1550563. ^ Brown WM, George M Jr., Wilson AC 1979. Rapid evolution of mitochondrial DNA. Proc Natl Acad Sci USA 76: 1967-1971. doi:10.1073/pnas.76.4.1967. PMID 109836. ^ Brown WM 1980. Polymorphism in mitochondrial DNA of humans as revealed by restriction endonuclease analysis. Proc Natl Acad Sci USA 77: 3605-3609. doi:10.1073/pnas.77.6.3605. PMID 6251473. ^ Paleo-DNA Laboratory - Forensic Services ^ http://www.eva.mpg.de/genetics/pdf/Stone.JFS.2001.pdf ^ Sutovsky, P., et. al Nov. 25, 1999. Ubiquitin tag for sperm mitochondria. Nature 402: 371-372. doi:10.1038/46466. PMID 10586873. Discussed in 1. ^ Vilà C, Savolainen P, Maldonado JE, and Amorin IR 13. Multiple and Ancient Origins of the Domestic Dog. Science 276: 1687-1689. doi:10.1126/science.276.5319.1687. ISSN 0036-8075. PMID 9180076. ^ Hoeh WR, Blakley KH, Brown WM 1991. Heteroplasmy suggests limited biparental inheritance of Mytilus mitochondrial DNA. Science 251: 1488-1490. doi:10.1126/science.1672472. PMID 1672472. ^ Penman, Danny 23 August 2002. Mitochondria can be inherited from both parents, NewScientist.com. Retrieved on 2008-02-05. ^ Kondo R, Matsuura ET, Chigusa SI 1992. Further observation of paternal transmission of Drosophila mitochondrial DNA by PCR selective amplification method. Genet. Res. 59 2: 81-4. PMID 1628820. ^ Meusel MS, Moritz RF 1993. Transfer of paternal mitochondrial DNA during fertilization of honeybee Apis mellifera L. eggs. Curr. Genet. 24 6: 539-43. doi:10.1007/BF00351719. PMID 8299176. ^ Gyllensten U, Wharton D, Josefsson A, Wilson AC 1991. Paternal inheritance of mitochondrial DNA in mice. Nature 352 6332: 255-7. doi:10.1038/352255a0. PMID 1857422. ^ Shitara H, Hayashi JI, Takahama S, Kaneda H, Yonekawa H 1998. Maternal inheritance of mouse mtDNA in interspecific hybrids: segregation of the leaked paternal mtDNA followed by the prevention of subsequent paternal leakage. Genetics 148 2: 851-7. PMID 9504930. ^ Zhao X, Li N, Guo W, et al 2004. Further evidence for paternal inheritance of mitochondrial DNA in the sheep Ovis aries. Hery 93 4: 399-403. doi:10.1038/sj.hdy.6800516. PMID 15266295. ^ Steinborn R, Zakhartchenko V, Jelyazkov J, et al 1998. Composition of parental mitochondrial DNA in cloned bovine embryos. FEBS Lett. 426 3: 352-6. doi:10.1016/S0014-57939800350-0. PMID 9600265. ^ Schwartz M, Vissing J 2002. Paternal inheritance of mitochondrial DNA. N. Engl. J. Med. 347 8: 576-80. doi:10.1056/NEJMoa020350. PMID 12192017. External links Mitomap - a human mitochondrial genome database 2 Defining mutations of mtDNA haplogroups and subclades MitoSearch : public mtDNA database mtDNA mutation rates A polymorphism in mitochondrial DNA associated with IQ? mtDNA and the global diaspora of modern humans Professor Stephen Oppenheimer's Genetic Map EMPOP - Mitochondrial DNA Control Region Database v d e Types of nucleic acids Constituents Nucleobases | Nucleosides | Nucleotides | Deoxynucleotides Ribonucleic acids RNA | mRNA pre-mRNA/hnRNA | tRNA | rRNA | aRNA | gRNA | miRNA | ncRNA | piRNA | shRNA | siRNA | snRNA | snoRNA | stRNA | ta-siRNA | tmRNA Deoxyribonucleic acids DNA | cDNA | gDNA | msDNA | mtDNA Nucleic acid analogues GNA | LNA | PNA | TNA | morpholino Cloning vectors phagemid | plasmid | lambda phage | cosmid | P1 phage | fosmid | BAC | YAC | HAC Major families of biochemicals Saccharides | Carbohydrates | Glycosides | | Amino acids | Peptides | Proteins | Glycoproteins | | Lipids | Terpenes | Steroids | Carotenoids Alkaloids | Nucleobases | Nucleic acids | | Enzyme cofactors | Flavonoids | Polyketides | Tetrapyrroles v d e Mitochondrial enzymes and transporters Outer membrane fatty acid degradation Carnitine palmitoyltransferase I, Long fatty acyl CoA synthetase tryptophan metabolism Kynureninase monoamine neurotransmitter metabolism Monoamine oxidase Intermembrane space Adenylate kinase - Creatine kinase Inner membrane oxidative phosphorylation Coenzyme Q - cytochrome c reductase, Cytochrome c, NADH dehydrogenase, Succinate dehydrogenase pyrimidine metabolism Dihydroorotate dehydrogenase mitochondrial shuttle Malate-aspartate shuttle, Glycerol phosphate shuttle other Glutamate aspartate transporter, Glycerol-3-phosphate dehydrogenase, ATP synthase, Carnitine palmitoyltransferase II Matrix citric acid cycle Citrate synthase, Aconitase, Isocitrate dehydrogenase, Oxoglutarate dehydrogenase, Succinyl coenzyme A synthetase, Fumarase, Malate dehydrogenase anaplerotic reactions Aspartate transaminase, Glutamate dehydrogenase, Pyruvate dehydrogenase complex urea cycle Carbamoyl phosphate synthetase I, Ornithine transcarbamylase, N-Acetylglutamate synthase alcohol metabolism ALDH2 Other/to be sorted Cholesterol side-chain cleavage enzyme Mitochondrial DNA Complex I 7 units - Complex III 1 unit - Complex IV 3 units - ATP synthase 2 units Retrieved from http://en..org/wiki/Mitochondrial_DNA Categories: DNA | Mitochondrial genetics Views Article Discussion this page History Personal tools Log in / create account Navigation Main page Contents Featured content Current events Random article Search Go Search Interaction Community portal Recent changes Contact Donate to Help Toolbox What links here Related changes Upload file Special pages Printable version Permanent link Cite this page Languages БългарÑ?ки Català Česky Deutsch Eesti Español Français Bahasa Indonesia Italiano Nederlands 日本語 ‪Norsk bokmÃ¥l‬ Polski Português Română РуÑ?Ñ?кий Svenska УкраїнÑ?ька ä¸æ–‡ This page was last modified on 6 August 2008, at 00:49
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