Diese Präsentation wurde erfolgreich gemeldet.
Die SlideShare-Präsentation wird heruntergeladen. ×

Mitochondrial DNA Report - Rasol Sindy

Anzeige
Anzeige
Anzeige
Anzeige
Anzeige
Anzeige
Anzeige
Anzeige
Anzeige
Anzeige
Anzeige
Anzeige
Sunday, November 13, 2022
Kurdistan Regional Government – Iraq
Ministry of Higher Education & Scientific Research
Universi...
I
Abstract
Mitochondrial DNA (mtDNA) is the physical embodiment of the genetic
information encoded in the mitochondrion. T...
II
List of Contents
1. Introduction..........................................................................................
Anzeige
Anzeige
Anzeige
Anzeige
Wird geladen in …3
×

Hier ansehen

1 von 21 Anzeige

Mitochondrial DNA Report - Rasol Sindy

Herunterladen, um offline zu lesen

Mitochondrial DNA (mtDNA) is the physical embodiment of the genetic information encoded in the mitochondrion. Technically, the term ‘mitochondrial DNA’ encompasses not only the mitochondrial genome per se, but additional DNA types (e.g., small linear plasmid-like DNAs) that are present in the mitochondria of some organisms. As its name implies, mtDNA is compartmentalized within the mitochondrion and is therefore physically and transcriptionally separate from the main nuclear genome of the eukaryotic cell. As well as being transcriptionally distinct from the nuclear genome, the mitochondrial genome and other types of mtDNA molecules are distinct in evolutionary origin, with the main mitochondrial genome having been derived from a eubacterial ancestor through a process of endosymbiosis.

Mitochondrial DNA (mtDNA) is the physical embodiment of the genetic information encoded in the mitochondrion. Technically, the term ‘mitochondrial DNA’ encompasses not only the mitochondrial genome per se, but additional DNA types (e.g., small linear plasmid-like DNAs) that are present in the mitochondria of some organisms. As its name implies, mtDNA is compartmentalized within the mitochondrion and is therefore physically and transcriptionally separate from the main nuclear genome of the eukaryotic cell. As well as being transcriptionally distinct from the nuclear genome, the mitochondrial genome and other types of mtDNA molecules are distinct in evolutionary origin, with the main mitochondrial genome having been derived from a eubacterial ancestor through a process of endosymbiosis.

Anzeige
Anzeige

Weitere Verwandte Inhalte

Aktuellste (20)

Anzeige

Mitochondrial DNA Report - Rasol Sindy

  1. 1. Sunday, November 13, 2022 Kurdistan Regional Government – Iraq Ministry of Higher Education & Scientific Research University of Zakho Faculty of Science Biology Department Mitochondrial DNA Subject (Genetics I) 5th Semester Prepared By Rahma Hishyar Rasol haji Ahmed Dunya Miqdad Ruwayda Abdullqadir Supervised By Prof. Dr. Yousif Mohammed
  2. 2. I Abstract Mitochondrial DNA (mtDNA) is the physical embodiment of the genetic information encoded in the mitochondrion. Technically, the term ‘mitochondrial DNA’ encompasses not only the mitochondrial genome per se, but additional DNA types (e.g., small linear plasmid-like DNAs) that are present in the mitochondria of some organisms. As its name implies, mtDNA is compartmentalized within the mitochondrion and is therefore physically and transcriptionally separate from the main nuclear genome of the eukaryotic cell. As well as being transcriptionally distinct from the nuclear genome, the mitochondrial genome and other types of mtDNA molecules are distinct in evolutionary origin, with the main mitochondrial genome having been derived from a eubacterial ancestor through a process of endosymbiosis.
  3. 3. II List of Contents 1. Introduction..........................................................................................................1 2. Genome structure and diversity...........................................................................2 2.1 Animal...........................................................................................................3 2.2 Plants and fung ..............................................................................................3 2.3 Protists...........................................................................................................4 3. Replication...........................................................................................................4 4. Genes on the human mtDNA and their transcription..........................................5 5. Regulation of transcription ..................................................................................5 6. Mitochondrial inheritance....................................................................................6 6.1 Female inheritance ........................................................................................7 6.2 Male inheritance............................................................................................8 7. The mitochondrial bottleneck..............................................................................9 8. Mitochondrial donation .....................................................................................10 9. Mutations and disease........................................................................................10 10. Causes:............................................................................................................11 11. Symptoms:......................................................................................................11 12. Test and diagnosis ..........................................................................................12 13. Susceptibility ..................................................................................................12 14. Genetic illness.................................................................................................13 15. Use in evolutionary biology and systematic biology mtDNA in nuclear DNA 13 16. Mitochondrial vs. Nuclear DNA ....................................................................14 17. Reference........................................................................................................16
  4. 4. III Table of Figure No. Figure Figure Page No. 1 Illustration of the location of mitochondrial DNA in human cells 2 2 mtDNA replication and transcription 6 3 Mitochondria inheritance 8 4 Mitochondrial bottleneck 9 5 Mutation of mtDNA 11
  5. 5. 1 1. Introduction Mitochondrial DNA (mtDNA or mDNA) isthe DNA locatedin mitochondria, cellular organelleswithin eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial DNA is only a small portion of the DNA in a eukaryotic cell; most of the DNA can be found in the cell nucleusand, in plants and algae, also in plastids such as chloroplasts. Nuclear and mitochondrial DNA are thought to be of separate evolutionary origin, with the mtDNA being derived from the circular genomes of bacteriaengulfed by the early ancestors of today's eukaryotic cells. This theory is called the endosymbiotic theory. 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, if not most, of them are thought to have originally been of bacterial origin, having since been transferred to the eukaryotic nucleus during evolution. The reasons mitochondria have retained some genes are debated. The existence in some species of mitochondrion-derived organelles lacking a genome suggests that complete gene loss is possible, and transferring mitochondrial genes to the nucleus has several advantages. The difficulty of targeting remotely- produced hydrophobic protein products to the mitochondrion is one hypothesis for why some genes are retained in mtDNA colocalisation for redox regulation is another, citing the desirability of localised control over mitochondrial machinery Recent analysis of a wide range of mtDNA genomes suggests that both these features may dictate mitochondrial gene retention.( Johnston & Williams,2016)
  6. 6. 2 2. Genome structure and diversity Across all organisms, there are six main genome types found in mitochondrial genomes, classified by their structure (i.e. circular versus linear), size, presence of introns or plasmid like structures, and whether the genetic material is a singular molecule or collection of homogeneous or heterogeneousmolecules. In many unicellular organisms (e.g., the ciliateTetrahymena and the green alga Chlamydomonas reinhardtii), and in rare cases also in multicellular organisms (e.g. in some species of Cnidaria), the mtDNA is found as linearly organized DNA. Most of these linear mtDNAs possess telomerase-independent telomeres (i.e., the ends of the linear DNA) with different modes of replication, which have made them interesting objects of research because many of these unicellular organisms with linear mtDNA are known pathogens. (Manicom et al., 1987). Mitochondrial DNA is the small circular chromosome found inside mitochondria. These organelles, found in all eukaryotic cells, are the powerhouse of the cell. The mitochondria, and thus mitochondrial DNA, are passed exclusively from mother to offspring through the egg cell. 0:33 Illustration of the location of mitochondrial DNA in human cells Figure 1 mtDNA
  7. 7. 3 2.1 Animal Most animals, specifically bilaterian animals, have a circular mitochondrial genome. Medusozoa and calcarea clades however have species with linear mitochondrial chromosomes. In terms of base pairs, the anemone Isarachnanthus nocturnus has the largest mitochondrial genome of any animal at 80,923 bp. In February 2020, a jellyfish-related parasite – Henneguya salminicola – was discovered that lacks mitochondrial genome but retains structures deemed mitochondrion-related organelles. Moreover, nuclear DNA genes involved in aerobic respiration and in mitochondrial DNA replication and transcription were either absent or present only as pseudogenes. This is the first multicellular organism known to have this absence of aerobic respiration and lives completely free of oxygen dependency. 2.2 Plants and fung There are three different mitochondrial genome types found in plants and fungi. The first type is a circular genome that has introns and may range from 19 to 1000 kbp in length. The second genome type is a circular genome (about 20–1000 kbp) that also has a plasmid-like structure (1 kb) The final genome type that can be found in plants and fungi is a linear genome made up of homogeneous DNA molecules Great 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).in Fungi, however, there is no single gene shared among all mitogenomes. Some plant species have enormous mitochondrial genomes, with Silene conica mtDNA containing as many as 11,300,000 base pairs. Surprisingly, even those huge mtDNAs contain the same number and kinds of genes as related plants with much smaller mtDNAs. The genome of the mitochondrion of the cucumber (Cucumis sativus) consists of three
  8. 8. 4 circular chromosomes (lengths 1556, 84 and 45 kilobases), which are entirely or largely autonomous with regard to their replication. 2.3 Protists Protists contain the most diverse mitochondrial genomes, with five different types found in this kingdom. Type 2, type 3 and type 5 mentioned in the plant and fungal genomes also exist in some protists, as do two unique genome types. One of these unique types is a heterogeneous collection of circular DNA molecules while the other is a heterogeneous collection of linear molecules Genome each range from 1–200 kbp in size.The smallest mitochondrial genome sequenced to date is the 5,967 bp mtDNA of the parasite Plasmodium falciparum. Endosymbiotic gene transfer, the process by which genes that were coded in the mitochondrial genome are transferred to the cell's main genome, likely explains why more complex organisms such as humans have smaller mitochondrial genomes than simpler organisms such as protists. 3. Replication Mitochondrial DNA is replicated by the DNA polymerase gamma complex which is composed of a 140 kDa catalytic DNA polymerase encoded by the POLG gene and two 55 kDa accessory subunits encoded by the POLG2 gene. The replisome machinery is formed by DNA polymerase, TWINKLE And mitochondrial SSB proteins. TWINKLE is a helicase, which unwinds short stretches of dsDNA in the 5' to 3' direction All these polypeptides are encoded in the nuclear genome. During embryogenesis, replication of mtDNA is strictly down-regulated from the fertilized oocyte through the preimplantation embryo The resulting reduction in per- cell copy number of mtDNA plays a role in the mitochondrial bottleneck, exploiting cell-to-cell variability to ameliorate the inheritance of damaging mutations.
  9. 9. 5 According to Justin St. John and colleagues, "At the blastocyststage, the onset of mtDNA replication is specific to the cells of the trophectoderm. In contrast, the cells of the inner cell mass restrict mtDNA replication until they receive the signals to differentiate to specific cell types." 4. Genes on the human mtDNA and their transcription The two strands of the human mitochondrial DNA are distinguished as the heavy strand and the light strand. The heavy strand is rich in guanine and encodes 12 subunits of the oxidative phosphorylation system, two ribosomal RNAs (12S and 16S), and 14 transfer RNAs (tRNAs). The light strand encodes one subunit, and 8 tRNAs. So, altogether mtDNA encodes for two rRNAs, 22 tRNAs, and 13 protein subunits, all of which are involved in the oxidative phosphorylation process. Between most (but not all) protein-coding regions, tRNAs are present (see the human mitochondrial genome map). During transcription, the tRNAs acquire their characteristic L-shape that gets recognized and cleaved by specific enzymes. With the mitochondrial RNA processing, individual mRNA, rRNA, and tRNA sequences are released from the primary transcript. Folded tRNAs therefore act as secondary structure punctuations. 5. Regulation of transcription The promoters for the initiation of the transcription of the heavy and light strands are located in the main non-coding region of the mtDNA called the displacement loop, the D-loop. There is evidence that the transcription of the mitochondrial rRNAs is regulated by the heavy-strand promoter 1 (HSP1), and the transcription of the polycistronic transcripts coding for the protein subunits are regulated by HSP2. Measurement of the levels of the mtDNA-encoded RNAs in bovine tissues has
  10. 10. 6 shown that there are major differences in the expression of the mitochondrial RNAs relative to total tissue RNA. Among the 12 tissues examined the highest level of expression was observed in heart, followed by brain and steroidogenic tissue samples. As demonstrated by the effect of the trophic hormone ACTH on adrenal cortex cells, the expression of the mitochondrial genes may be strongly regulated by external factors, apparently to enhance the synthesis of mitochondrial proteins necessary for energy production. Interestingly, while the expression of protein- encoding genes was stimulated by ACTH, the levels of the mitochondrial 16S rRNA showed no significant change. 6. Mitochondrial inheritance In most multicellular organisms, mtDNA is inherited from the mother (maternally inherited). Mechanisms for this include simple dilution (an egg contains on average 200,000 mtDNA molecules, whereas a healthy human sperm has been reported to contain on average 5 molecules), degradation of sperm mtDNA in the male genital tract and 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 inheritance) pattern of mtDNA inheritance is found in most animals, most plants and Figure2 mtDNA replication and transcription
  11. 11. 7 also in fungi.In a study published in 2018, human babies were reported to inherit mtDNA from both their fathers and their mothers resulting in mtDNA heteroplasmy. 6.1 Female inheritance In sexual reproduction, mitochondria are normally inherited exclusively from the mother; the mitochondria in mammalian sperm are usually destroyed by the egg cell after fertilization. Also, mitochondria are only in the sperm tail, which is used for propelling the sperm cells and 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. Some in vitro fertilization techniques, particularly injecting a sperm into an oocyte, may interfere with this.The fact that mitochondrial DNA is mostly maternally inherited enables genealogical researchers to trace maternal lineage far back in time. (Y-chromosomal DNA, paternally inherited, is used in an analogous way to determine the patrilineal history.) This is usually accomplished on human mitochondrial DNAby sequencing the hypervariable control regions(HVR1 or HVR2), and sometimes the complete molecule of the mitochondrial DNA, as a genealogical DNA test HVR1, for example, consists of about 440 base pairs. These 440 base pairs are compared to the same regions of other individuals (either specific people or subjects in a database) to determine maternal lineage. Most often, the comparison is made with the revised Cambridge Reference Sequence. Vilà et al. have published studies tracing the matrilineal descent of domestic dogs from wolves. 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.
  12. 12. 8 6.2 Male inheritance Male mitochondrial DNA inheritance has been discovered in Plymouth Rock chickens. Evidence supports rare instances of male mitochondrial inheritance in some mammals as well. Specifically, documented occurrences exist for mice, where the male-inherited mitochondria were subsequently rejected. It has also been found in sheep and in cloned cattle. Rare cases of male mitochondrial inheritance have been documented in humans. Although many of these cases involve cloned embryos or subsequent rejection of the paternal mitochondria, others document in vivo inheritance and persistence under lab conditions doubly uniparental inheritance of mtDNA is observed in bivalve mollusks. In those species, females have only one type of mtDNA (F), whereas males have F type mtDNA in their somatic cells, but M type of mtDNA (which can be as much as 30% divergent) in germline cells. Paternally inherited mitochondria have additionally been reported in some insects such as fruit flies, honeybees, and periodical cicadas. Figure3 mitochondria inheritance
  13. 13. 9 7. The mitochondrial bottleneck Entities subject to uniparental inheritance and with little to no recombination may be expected to be subject to Muller's ratchet, the accumulation of deleterious mutations until functionality is lost. Animal populations of mitochondria avoid this through a developmental process known as the mtDNA bottleneck. The bottleneck exploits random processes in the cell to increase the cell-to-cell variability in mutant load as an organism develops: a single egg cell with some proportion of mutant mtDNA thus produces an embryo in which different cells have different mutant loads. Cell-level selection may then act to remove those cells with more mutant mtDNA, leading to a stabilisation or reduction in mutant load between generations. The mechanism underlying the bottleneck is debated, with a recent mathematical and experimental metastudy providing evidence for a combination of the random partitioning of mtDNAs at cell divisions and the random turnover of mtDNA molecules within the cell. Figure 4 mitochondrial bottleneck
  14. 14. 10 8. Mitochondrial donation An IVF technique known as mitochondrial donation or mitochondrial replacement therapy (MRT) results in offspring containing mtDNA from a donor female, and nuclear DNA from the mother and father. In the spindle transfer procedure, the nucleus of an egg is inserted into the cytoplasm of an egg from a donor female which has had its nucleus removed, but still contains the donor female's mtDNA. The composite egg is then fertilized with the male's sperm. The procedure is used when a woman with genetically defective mitochondria wishes to procreate and produce offspring with healthy mitochondria. The first known child to be born as a result of mitochondrial donation was a boy born to a Jordanian couple in Mexico on 6 April 2016. 9. Mutations and disease Mitochondrial disorders impair the function of mitochondria, the tiny compartments in every cell of the body that produce the energy needed by cells. Depending on which cells have fewer or lower-functioning mitochondria, different symptoms may occur. Organs and other parts of the body that require more energy, such as the heart, muscles and brain, are often affected. Mitochondrial disease is the name for a large number of mitochondrial disorders, with different genetic causes and presentations.Mitochondrial disease may be caused by genetic mutations in the body’s nuclear DNA (the DNA found in the nucleus of cells) or by genetic mutations or deletions in the body’s mitochondrial DNA (mtDNA < the DNA found in cells’ mitochondria). mtDNA common mutation syndromes are mitochondrial disorders caused by recurrent mtDNA mutations in unrelated families and populations. Despite the name, these are rare conditions; “common” means that these specific mutations recur across families and ethnic groups.
  15. 15. 11 10.Causes: mtDNA common mutation syndromes are often inherited from the mother. Only women pass mtDNA mutations on to their children through the oocyte. Men with mtDNA mutations may be affected by the conditions, but do not pass them on to their children.In some cases, mtDNA mutations are spontaneous (de novo) in the affected individual and are not inherited. 11.Symptoms: mtDNA common mutation syndromes vary widely by type and from case to case. There is also a great deal of overlap in symptoms among different mitochondrial conditions. Doctors now consider them to represent a broad spectrum of features and severity in diagnosis and treatment. Two patients may have the same symptoms with different genetic causes, and will therefore be diagnosed with different mitochondrial disorders. And two patients with the same genetic disorder may have very different symptoms and require different treatment.Listed here are the most common symptoms for each of the classical clinical mtDNA mutation syndromes. Figure 5 mutation of mtDNA
  16. 16. 12 12.Test and diagnosis If a mtDNA common mutation syndrome is suspected based on the patient’s physical symptoms and history, additional tests are performed to make a diagnosis. These tests may include: -Blood tests to look for high concentrations of lactic acid and other abnormalities -Cerebrospinal fluid (CSF) analysis to look for elevated protein levels MRI of the brain and/or spinal cord -Electrocardiography (ECG or EKG) to detect heart rhythm abnormalities -Echocardiography (Echo) to detect cardiomyopathy -Muscle biopsy to obtain tissue in which to measure biochemical function, perform DNA testing, and detect ragged red fibers characteristic of MELAS syndrome and MERRF -Genetic testing using blood, muscle, skin, urinary sediment or other tissue samples to look for mutations in mtDNA and nuclear DNA 13.Susceptibility The concept that mtDNA is particularly susceptible to reactive oxygen species generated by the respiratory chain due to its proximity remains controversial. mtDNA does not accumulate any more oxidative base damage than nuclear DNA. It has been reported that at least some types of oxidative DNA damage are repaired more efficiently in mitochondria than they are in the nucleus. mtDNA is packaged with proteins which appear to be as protective as proteins of the nuclear chromatin. Moreover, mitochondria evolved a unique mechanism which maintains mtDNA integrity through degradation of excessively damaged genomes followed by replication of intact/repaired mtDNA. This mechanism is not present in
  17. 17. 13 the nucleus and is enabled by multiple copies of mtDNA present in mitochondria The outcome of mutation in mtDNA may be an alteration in the coding instructions for some proteins, which may have an effect on organism metabolism and/or fitness. 14.Genetic illness Mutations of mitochondrial DNA can lead to a number of illnesses including exercise intoleranceand Kearns–Sayre syndrome (KSS), which causes a person to lose full function of heart, eye, and muscle movements. Some evidence suggests that they might be major contributors to the aging process and age-associated pathologies. Particularly in the context of disease, the proportion of mutant mtDNA molecules in a cell is termed heteroplasmy. The within-cell and between-cell distributions of heteroplasmy dictate the onset and severity of diseaseand are influenced by complicated stochastic processes within the cell and during development. Mutations in mitochondrial tRNAs can be responsible for severe diseases like the MELAS and MERRF syndromes. Mutations in nuclear genes that encode proteins that mitochondria use can also contribute to mitochondrial diseases. These diseases do not follow mitochondrial inheritance patterns, but instead follow Mendelian inheritance patterns. 15.Use in evolutionary biology and systematic biology mtDNA in nuclear DNA Sequencing the genomes of more than 66,000 people revealed that most of them had some mitochondrial DNA inserted into their nucleargenomes. More than 90% of these nuclear-mitochondrial segments (NUMTs) were inserted into the nuclear genome within the last 5 or 6 million years, that is, after humans diverged from apes. It appears that organellar DNA is much more often transferred to nuclear DNA than
  18. 18. 14 previously thought. This observation also supports the idea of the endosymbiont theory that eukaryotes have evolved from endosymbionts which turned into organelles while transferring most of their DNA to the nucleus so that the organellar genome shrunk in the process. 16.Mitochondrial vs. Nuclear DNA Similar to the nuclear genome, the mitochondrial genome is built of double-stranded DNA, and it encodes genes However, the mitochondrial genome differs from the nuclear genome in several ways Many interesting features distinguish human mitochondrial DNA from its nuclear counterpart, including the following: • The mitochondrial genome is circular, whereas the nuclear genome is linear • The mitochondrial genome is built of 16,569 DNA base pairs, whereas the nuclear genome is made of 3.3 billion DNA base pairs. • The mitochondrial genome contains 37 genes that encode 13 proteins, 22 tRNAs, and 2 rRNAs. • The 13 mitochondrial gene-encoded proteins all instruct cells to produce protein subunits of the enzyme complexes of the oxidative phosphorylation system, which enables mitochondria to act as the powerhouses of our cells. • The small mitochondrial genome is not able to independently produce all of the proteins needed for functionality; thus, mitochondria rely heavily on imported nuclear gene products. • One mitochondrion contains dozens of copies of its mitochondrial genome. In addition, each cell contains numerous mitochondria. Therefore, a
  19. 19. 15 given cell can contain several thousand copies of its mitochondrial genome, but only one copy of its nuclear genome. • The mitochondrial genome is not enveloped, and is it not packaged into chromatin. • The mitochondrial genome contains few, if any, noncoding DNA sequences. (Three percent of the mitochondrial genome is noncoding DNA, whereas 93% of the nuclear genome is noncoding DNA). • Some mitochondrial coding sequences (triplet codons) do not follow the universal codon usage rules when they are translated into proteins. • Some mitochondrial nucleotide bases exhibit functional overlap between two genes; in other words, the same nucleotide can sometimes function as both the last base of one gene and the first base of the next gene. • The mitochondrial mode of inheritance is strictly maternal, whereas nuclear genomes are inherited equally from both parents. Therefore, mitochondria-associated disease mutations are also always inherited maternally. • Mitochondrial genes on both DNA strands are transcribed in a polycistronic manner: Large mitochondrial mRNAs contain the instructions to build many different proteins, which are encoded one after the next along the mRNA. In contrast, nuclear genes are usually transcribed one at a time from their own mRNA.
  20. 20. 16 17.Reference 1. Siekevitz P (1957). "Powerhouse of the cell". Scientific American. 197 (1): 131–40.Bibcode:1957SciAm.197a.131S. 2. Iborra FJ, Kimura H, Cook PR (May 2004). "The functional organization of mitochondrial genomes in human cells". BMC Biology. 3. Sykes B (10 September 2003). "Mitochondrial DNA and human history". The Human Genome. Wellcome Trust. Archived from the original on 7 September 2015. Retrieved 5 February 2012. 4. Anderson S, Bankier AT, Barrell BG, de Bruijn MH, Coulson AR, Drouin J, Eperon IC, Nierlich DP, Roe BA, Sanger F, Schreier PH, Smith AJ, Staden R, Young IG. (1981). "Sequence and organization of the human mitochondrial genome". Nature. 290 (5806): 457–65. Bibcode:1981Natur.290..457A. doi:10.1038/290457a0. PMID 7219534. S2CID 4355527. 5. Boursot P, Bonhomme F (1 January 1986). "[Not Available]". Génétique, Sélection, Évolution. 18 (1): 73–98. doi:10.1186/1297-9686-18-1-73. PMC 2713894. PMID 22879234. 6. Delsuc F, Stanhope MJ, Douzery EJ (August 2003). "Molecular systematics of armadillos (Xenarthra, Dasypodidae): contribution of maximum likelihood and Bayesian analyses of mitochondrial and nuclear genes". Molecular Phylogenetics and Evolution. 28 (2): 261–75. doi:10.1016/s1055- 7903(03)00111-8. PMID 12878463. Archivedfrom the original on 4 November 2018. Retrieved 4 November 2018. 7. Hassanin A, An J, Ropiquet A, Nguyen TT, Couloux A (March 2013). "Combining multiple autosomal introns for studying shallow phylogeny and taxonomy of Laurasiatherian mammals: Application to the tribe Bovini
  21. 21. 17 (Cetartiodactyla, Bovidae)". Molecular Phylogenetics and Evolution. 66 (3): 766–75. doi:10.1016/j.ympev.2012.11.003. PMID 23159894. 8. a b c Johnston IG, Williams BP (February 2016). "Evolutionary Inference across Eukaryotes Identifies Specific Pressures Favoring Mitochondrial Gene Retention". Cell Systems. 2 (2): 101–11. doi:10.1016/j.cels.2016.01.013. PMID 27135164. 9. van der Giezen M, Tovar J, Clark CG (2005). "Mitochondrion‐Derived Organelles in Protists and Fungi". A Survey of Cell Biology. International Review of Cytology. Vol. 244. pp. 175–225. doi:10.1016/S0074- 7696(05)44005-X. ISBN 978-0-12-364648-4. PMID 16157181. 10.Adams KL, Palmer JD (December 2003). "Evolution of mitochondrial gene content: gene loss and transfer to the nucleus". Molecular Phylogenetics and Evolution. 29 (3): 380–95. doi:10.1016/S1055-7903(03)00194-5. PMID 14615181. 11.Björkholm P, Harish A, Hagström E, Ernst AM, Andersson SG (August 2015). "Mitochondrial genomes are retained by selective constraints on protein targeting". Proceedings of the National Academy of Sciences of the United States of America. 112 (33): 10154–61. Bibcode:2015PNAS..11210154B. doi:10.1073/pnas.1421372112. PMC 4547212. PMID 26195779. 12.Manicom B. Q., Bar-Joseph M., Rosner A., Vigodsky-Haas H., Kotze J. M. (1987). Potential applications of random dna probes and restriction-fragment- length-polymorphisms in the taxonomy of the fusaria. Phytopathology 77, 669–672. 10.1094/Phyto-77-669

×