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95-607: The Khakhassian people are direct descendants of various ancient cultures that have inhabited southern Siberia, including the Andronovo culture , Samoyedic peoples , the Tagar culture , and the Yenisei Kyrgyz culture, although some populations traditionally called Khakhassian are not related to Khakhassians or any other ethnic group present in the area. The Khakas people were historically known as Kyrgyz , before being labelled as Tatar by

190-537: A Mongolian ruler. The Russians arrived shortly after the Kyrgyz left, and an inflow of Russian agragian settlers began. In the 1820s, gold mines started to be developed around Minusinsk , which became a regional industrial center. The names Khongorai and Khoorai were applied to the Khakas before they became known as the Khakas. Khakas refer to themselves as Tadar. Khoorai (Khorray) has also been in use to refer to them. Now

285-507: A circular mitochondrial genome. Medusozoa and calcarea clades however include species with linear mitochondrial chromosomes. With a few exceptions, animals have 37 genes in their mitochondrial DNA: 13 for proteins , 22 for tRNAs , and 2 for rRNAs . Mitochondrial genomes for animals average about 16,000 base pairs in length. The anemone Isarachnanthus nocturnus has the largest mitochondrial genome of any animal at 80,923 bp. The smallest known mitochondrial genome in animals belongs to

380-585: 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. Entities subject to uniparental inheritance and with little to no recombination may be expected to be subject to Muller's ratchet ,

475-500: 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

570-435: A finding that has been rejected by other scientists. 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 present solely in the midpiece, which is used for propelling the sperm cells, and sometimes the midpiece, along with the tail, is lost during fertilization. In 1999 it

665-434: 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

760-545: 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 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 ,

855-528: A hundred houses have been discovered. Andronovo houses were generally constructed from pine , cedar , or birch , and were usually aligned overlooking the banks of rivers . Larger homes range in the size from 80 to 300 m , and probably belonged to extended families, a typical feature among early Indo-Iranians. Soma may have originated in the Andronovo culture. Andronovo livestock included cattle , horses , sheep , goats and camels . The domestic pig

950-405: A mainstay of phylogenetics and evolutionary biology . It also permits tracing the relationships of populations, and so has become important in anthropology and biogeography . Nuclear and mitochondrial DNA are thought to have separate evolutionary origins, with the mtDNA derived from the circular genomes of bacteria engulfed by the ancestors of modern eukaryotic cells. This theory is called

1045-542: A mutation in mtDNA has been used to help diagnose prostate cancer in patients with negative prostate biopsy . mtDNA alterations can be detected in the bio-fluids of patients with cancer. mtDNA is characterized by the high rate of polymorphisms and mutations. Some of which are increasingly recognized as an important cause of human pathology such as oxidative phosphorylation (OXPHOS) disorders, maternally inherited diabetes and deafness (MIDD), Type 2 diabetes mellitus, Neurodegenerative disease , heart failure and cancer. Though

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1140-626: A mutational (contrary to the selective one) explanation for the observation that long-lived species have GC-rich mtDNA: long-lived species become GC-rich simply because of their biased process of mutagenesis. An association between mtDNA mutational spectrum and species-specific life-history traits in mammals opens a possibility to link these factors together discovering new life-history-specific mutagens in different groups of organisms. Deletion breakpoints frequently occur within or near regions showing non-canonical (non-B) conformations, namely hairpins, cruciforms and cloverleaf-like elements. Moreover, there

1235-514: A plague (although it was not the plague). The three people in that grave all died at the same time, and the researcher believes that this para-plague is what killed them. Most researchers associate the Andronovo horizon with early Indo-Iranian languages . It is credited with the invention of the spoke-wheeled chariot around 2000 BC, if we include the Sintashta culture , where the oldest known chariots have been found. The association between

1330-473: A pottery made by late Bronze Age nomads. "It is likely that militarized elite, whose power was based on the physical control of fellow tribesmen and neighbors with the help of riding and fighting skills, was buried in the Novoilinovsky-2 burial ground. The rider has a significant advantage over the infantryman. There may be another explanation: These elite fulfilled the function of mediating conflicts within

1425-521: A result of mitochondrial donation was a boy born to a Jordanian couple in Mexico on 6 April 2016. 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

1520-507: A role in the mitochondrial bottleneck, exploiting cell-to-cell variability to ameliorate the inheritance of damaging mutations. According to Justin St. John and colleagues, "At the blastocyst stage, 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." The two strands of

1615-435: A species and also for identifying and quantifying the phylogeny (evolutionary relationships; see phylogenetics ) among different species. 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

1710-623: 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. 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

1805-448: A wide range of mtDNA genomes suggests that both these features may dictate mitochondrial gene retention. Across all organisms, there are six main mitochondrial genome types, classified by 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 heterogeneous molecules. In many unicellular organisms (e.g.,

1900-551: Is a collection of similar local Late Bronze Age cultures that flourished c.  2000–1150 BC, spanning from the southern Urals to the upper Yenisei River in central Siberia and western Xinjiang in the east. In the south, the Andronovo sites reached Tajikistan and Uzbekistan . It is almost universally agreed among scholars that the Andronovo culture was Indo-Iranian . Some researchers have preferred to term it an archaeological complex or archaeological horizon . Andronovo culture's first stage could have begun at

1995-532: Is a linear genome made up of homogeneous DNA molecules (type 5). Great variation in mtDNA gene content and size exists among fungi and plants, although there appears to be a core subset of genes 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

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2090-467: Is a well-established marker of oxidative DNA damage. In persons with amyotrophic lateral sclerosis (ALS), the enzymes that normally repair 8-oxoG DNA damages in the mtDNA of spinal motor neurons are impaired. Thus oxidative damage to mtDNA of motor neurons may be a significant factor in the etiology of ALS. Over the past decade, an Israeli research group led by Professor Vadim Fraifeld has shown that strong and significant correlations exist between

2185-628: Is considered the type site of the Sintashta culture , forming one of the earliest parts of the "Andronovo horizon". It is conjectured that the language spoken was still in the Proto-Indo-Iranian stage. Comparisons between the archaeological evidence of the Andronovo and textual evidence of Indo-Iranians (i. e. the Vedas and the Avesta ) are frequently made to support the Indo-Iranian identity of

2280-487: Is data supporting the involvement of helix-distorting intrinsically curved regions and long G-tetrads in eliciting instability events. In addition, higher breakpoint densities were consistently observed within GC-skewed regions and in the close vicinity of the degenerate sequence motif YMMYMNNMMHM. Unlike nuclear DNA, which is inherited from both parents and in which genes are rearranged in the process of recombination , there

2375-517: Is notably absent, which is typical of a mobile economy. The percentage of cattle among Andronovo remains are significantly higher than among their western Srubna neighbours. The horse was represented on Andronovo sites and was used for both riding and traction. According to the Journal of Archaeological Science , in July 2020, scientists from South Ural State University studied two Late Bronze Age horses with

2470-514: 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 . An IVF technique known as mitochondrial donation or mitochondrial replacement therapy (MRT) results in offspring containing mtDNA from

2565-540: Is severely degraded. Autosomal cells only have two copies of nuclear DNA, but can have hundreds of copies of mtDNA due to the multiple mitochondria present in each cell. This means highly degraded evidence that would not be beneficial for STR analysis could be used in mtDNA analysis. mtDNA may be present in bones, teeth, or hair, which could be the only remains left in the case of severe degradation. In contrast to STR analysis, mtDNA sequencing uses Sanger sequencing . The known sequence and questioned sequence are both compared to

2660-485: Is the first multicellular organism known to have this absence of aerobic respiration and live completely free of oxygen dependency. There are three different mitochondrial genome types in plants and fungi. The first type is a circular genome that has introns (type 2) 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) (type 3). The final genome type found in plants and fungi

2755-452: Is used in an analogous way to determine the patrilineal history.) This is usually accomplished on human mitochondrial DNA by 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

2850-510: 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, 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. mtDNA testing can be used by forensic scientists in cases where nuclear DNA

2945-517: 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

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3040-669: The Abashevo culture . The earliest historical peoples associated with the area are the Cimmerians and Saka / Scythians , appearing in Assyrian records after the decline of the Alekseyevka culture , migrating into Ukraine from ca. the 9th century BC (see also Ukrainian stone stela ), and across the Caucasus into Anatolia and Assyria in the late 8th century BC, and possibly also west into Europe as

3135-773: The Indo-Iranian to Central Asia, but not as far as the seats of the Medes , Persians or Indo-Aryans". He has developed the Kulturkugel model that has the Indo-Iranians taking over Bactria-Margiana cultural traits but preserving their language and religion while moving into Iran and India. Based on its use by Indo-Aryans in Mitanni and Vedic India, its prior absence in the Near East and Harappan India, and its 17th–16th century BC attestation at

3230-648: The Minusinsk depression, with some sites as far west as the southern Ural Mountains , overlapping with the area of the earlier Afanasevo culture . Additional sites are scattered as far south as the Kopet Dag ( Turkmenistan ), the Pamir ( Tajikistan ) and the Tian Shan ( Kyrgyzstan ). The northern boundary vaguely corresponds to the beginning of the Taiga . More recently, evidence for

3325-551: The Revolution of 1905 , a movement towards autonomy developed. When Soviets came to power in 1923, the Khakas National District was established, and various ethnic groups (Beltir, Sagai, Kachin, Koibal , and Kyzyl) were artificially "combined" into one—the Khakas. The National District was reorganized into Khakas Autonomous Oblast , a part of Krasnoyarsk Krai , in 1930. The Republic of Khakassia in its present form

3420-867: The Thracians (see Thraco-Cimmerian ), and the Sigynnae , located by Herodotus beyond the Danube, north of the Thracians, and by Strabo near the Caspian Sea . Both Herodotus and Strabo identify them as Iranian. The Andronovo culture comprised both highly mobile communities and settled villages, with a notable concentration of settlements in its Central Asian regions. Fortifications include ditches, earthen banks as well as timber palisades, of which an estimated twenty have been discovered. Andronovo villages typically contain around two to twenty houses, but settlements containing as many as

3515-518: The ciliate Tetrahymena and the green alga Chlamydomonas reinhardtii ), and in rare cases also in multicellular organisms (e.g. in some species of Cnidaria ), the mtDNA is linear 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 . Most ( bilaterian ) animals have

3610-499: The endosymbiotic theory . In the cells of extant organisms, the vast majority of the proteins in the mitochondria (numbering approximately 1500 different types in mammals ) are coded by nuclear DNA , but the genes for some, if not most, of them are thought to be of bacterial origin, having 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

3705-510: The Andronovo area, i. e., Uralic and Yeniseian. MtDNA Mitochondrial DNA ( mtDNA and mDNA ) is the DNA located in the mitochondria organelles in a eukaryotic cell that converts chemical energy from food into adenosine triphosphate (ATP). Mitochondrial DNA is a small portion of the DNA contained in a eukaryotic cell; most of the DNA is in the cell nucleus , and, in plants and algae,

3800-504: The Andronovo culture and the Indo-Iranians is corroborated by the distribution of Iranian place-names across the Andronovo horizon and by the historical evidence of dominance by various Iranian-speaking peoples, including the Saka ( Scythians ), Sarmatians and Alans , throughout the Andronovo horizon during the 1st millennium BC. The Sintashta site on the upper Ural River , noted for its chariot burials and kurgans containing horse burials,

3895-683: The Andronovo people spoke a separate branch of Indo-Iranian. He claims that borrowings in the Finno-Ugric languages support this view. Vladimir Napolskikh has proposed that borrowings in Finno-Ugric indicate that the language was specifically of the Indo-Aryan type. Since older forms of Indo-Iranian words have been taken over in Uralic and Proto-Yeniseian , occupation by some other languages (also lost ones) cannot be ruled out altogether, at least for part of

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3990-510: The Andronovo site of Sintashta , Kuzmina (1994) argues that the chariot corroborates the identification of Andronovo as Indo-Iranian. Klejn (1974) and Brentjes (1981) found the Andronovo culture much too late for an Indo-Iranian identification since chariot-using Aryans appear in Mitanni by the 15th century BC. However, Anthony & Vinogradov (1995) dated a chariot burial at Krivoye Lake to around 2000 BC. Eugene Helimski has suggested that

4085-680: The Andronovo. The modern explanations for the Indo-Iranianization of Greater Iran and the Indian subcontinent rely heavily on the supposition that the Andronovo expanded southwards into Central Asia or at least achieved linguistic dominance across the Bronze Age urban centres of the region, such as the Bactria–Margiana Archaeological Complex . While the earlier phases of the Andronovo culture are regarded as co-ordinate with

4180-438: The DNA also is found in plastids , such as chloroplasts . Human mitochondrial DNA was the first significant part of the human genome to be sequenced. This sequencing revealed that human mtDNA has 16,569 base pairs and encodes 13 proteins . As in other vertebrates, the human mitochondrial genetic code differs slightly from nuclear DNA. Since animal mtDNA evolves faster than nuclear genetic markers, it represents

4275-782: The Dzungars and Russians in 1635. The Dzungar Oirat Kalmyks coerced the Yenisei Kyrgyz into submission. Some of the Yenisei Kyrgyz were relocated into the Dzungar Khanate by the Dzungars , and then the Qing moved them from Dzungaria to northeastern China in 1761, where they became known as the Fuyu Kyrgyz . Sibe Bannermen were stationed in Dzungaria while Northeastern China (Manchuria) was where some of

4370-641: The Imperial Russians following the conquest of Siberia . The name Tatar then became the autonym used by the Khakas to refer to themselves, in the form Tadar . Following the Russian Revolution , the Soviet authorities changed the name of the group to Khakas , a newly-formed name based on the Chinese name for the Kyrgyz people, Xiaqiasi . The Yenisei Kyrgyz were made to pay tribute in a treaty concluded between

4465-648: The Khakas call themselves Tadar and do not use Khakas to describe themselves in their own language. They are also called Abaka Tatars. During the 19th century, many Khakas accepted the Russian ways of life, and most were converted en masse to Russian Orthodox Christianity . Shamanism , however, is still common;. Many Christians practice shamanism with Christianity. In Imperial Russia, the Khakas used to be known under other names, used mostly in historic contexts: Minusinsk Tatars ( Russian : минуси́нские тата́ры ), Abakan Tatars (абака́нские тата́ры), and Yenisei Turks . During

4560-632: The Khakas predominantly are Orthodox Christians ( Russian Orthodox Church ). Also there is traditional shamanism ( Tengrism ), including following movements: Andronovo culture Pontic Steppe Caucasus East Asia Eastern Europe Northern Europe Pontic Steppe Northern/Eastern Steppe Europe South Asia Steppe Europe Caucasus India Indo-Aryans Iranians East Asia Europe East Asia Europe Indo-Aryan Iranian Indo-Aryan Iranian Others European The Andronovo culture

4655-642: The Khakhas population. Other paternal haplogroups in Khakassians include Haplogroup Q , which is probably the "original" Siberian lineage in Khakassians. It has a frequency of approximately 4.8% in the Khakassian population. Minor frequencies of haplogroups R1b , C3 , and E1 were also reported. Over 80% of Khakassian mtDNA lineages belong to East Eurasian lineages, although a significant percentage (18.9%) belong to various West Eurasian mtDNA lineages. At present,

4750-558: The Revised Cambridge Reference Sequence to generate their respective haplotypes. If the known sample sequence and questioned sequence originated from the same matriline, one would expect to see identical sequences and identical differences from the rCRS. Cases arise where there are no known samples to collect and the unknown sequence can be searched in a database such as EMPOP. The Scientific Working Group on DNA Analysis Methods recommends three conclusions for describing

4845-543: 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

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4940-419: The accumulation of mtDNA damage in several organs of rats. For example, dietary restriction prevented age-related accumulation of mtDNA damage in the cortex and decreased it in the lung and testis. Increased mt DNA damage is a feature of several neurodegenerative diseases . The brains of individuals with Alzheimer's disease have elevated levels of oxidative DNA damage in both nuclear DNA and mtDNA, but

5035-655: The aid of radiocarbon dating from Kurgan 5 of the Novoilinovsky 2 cemetery in the Lisakovsk city in the Kostanay region . Researcher Igor Chechushkov, indicated that the Andronovites had an ability on horse riding several centuries earlier than many researchers had previously expected. Among the horses investigated, the stallion was nearly 20 years old and the mare was 18 years old. According to scientists, animals were buried with

5130-476: The characteristics of Andronovo culture is its pottery, especially in campsites located in Central Asia, some of them very close to settlements of Bactria–Margiana Archaeological Complex in the south. This pottery is called Incised Coarse Ware (ICW), which is handmade and grey to brown in color, as well as incised with geometrical decoration, spread over much of Eurasian region, from Southern Urals to Kashgar ,

5225-458: The chronology and model of eastward spread due to increasing evidence for the earlier presence of these cultural features in parts of east Central Asia. The geographical extent of the culture is vast and difficult to delineate exactly. On its western fringes, it overlaps with the approximately contemporaneous, but distinct, Srubna culture in the Volga - Ural interfluvial. To the east, it reaches into

5320-456: The coding instructions for some proteins, which may have an effect on organism metabolism and/or fitness. 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 heart, eye, and muscle movements. Some evidence suggests that they might be major contributors to the aging process and age-associated pathologies . Particularly in

5415-410: The collective, and therefore had power and high social status. Metaphorically, this kind of elite can be called Sheriffs of the Bronze Age" said Igor Chechushkov. The Andronovo dead were buried in timber or stone chambers under both round and rectangular kurgans ( tumuli ). Burials were accompanied by livestock, wheeled vehicles, cheek-pieces for horses, and weapons, ceramics and ornaments. Among

5510-434: The comb jelly Vallicula multiformis , which consist of 9,961 bp. In February 2020, a jellyfish-related parasite – Henneguya salminicola – was discovered that lacks a 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

5605-714: 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 disease and 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. Recently

5700-403: The differences between a known mtDNA sequence and a questioned mtDNA sequence: exclusion for two or more differences between the sequences, inconclusive if there is one nucleotide difference, or cannot exclude if there are no nucleotide differences between the two sequences. The rapid mutation rate (in animals) makes mtDNA useful for assessing genetic relationships of individuals or groups within

5795-626: The differences in animal species maximum life spans in a multiplicative manner (i.e., species maximum life span = their mtDNA GC% * metabolic rate). To support the scientific community in carrying out comparative analyses between mtDNA features and longevity across animals, a dedicated database was built named MitoAge . De novo mutations arise either due to mistakes during DNA replication or due to unrepaired damage caused in turn by endogenous and exogenous mutagens. It has been long believed that mtDNA can be particularly sensitive to damage caused by reactive oxygen species (ROS), however G>T substitutions,

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5890-407: The end of the 3rd millennium BC, with cattle grazing, as natural fodder was by no means difficult to find in the pastures close to dwellings. The slightly older Sintashta culture ( c. 2200–1900 BC), formerly included within the Andronovo culture, is now considered separately to Early Andronovo cultures. Allentoft et al. (2015) concluded from their genetic studies that the Andronovo culture and

5985-577: The final Alekseyevka Phase (1400–1000 BC), the Andronovo cultures move intensively eastwards, expanding as far east as the Upper Yenisei River , succeeding the non-Indo-European Okunev culture . In southern Siberia and Kazakhstan, the Andronovo culture was succeeded by the Karasuk culture (1500–800 BC). On its western border, it is roughly contemporaneous with the Srubna culture , which partly derives from

6080-598: The former (or Srubna), alternatively the result of an amalgamation of steppe cultures and the Central Asian oasis cultures ( Bishkent culture and Vakhsh culture ). In the initial Sintashta- Petrovka phase, the Andronovo culture is limited to the northern and western steppes in the southern Urals -Kazakhstan. Since then, at the 2nd millennium, in the Alakul Phase (2000–1700 BC), the Fedorovo Phase (1850–1450 BC) and

6175-403: The genetic distances of distantly related species. Statistical models that treat substitution rates among codon positions separately, can thus be used to simultaneously estimate phylogenies that contain both closely and distantly related species Mitochondrial DNA was admitted into evidence for the first time ever in a United States courtroom in 1996 during State of Tennessee v. Paul Ware . In

6270-477: The hallmark of the oxidative damage in the nuclear genome, are very rare in mtDNA and do not increase with age. Comparing the mtDNA mutational spectra of hundreds of mammalian species, it has been recently demonstrated that species with extended lifespans have an increased rate of A>G substitutions on single-stranded heavy chain. This discovery led to the hypothesis that A>G is a mitochondria-specific marker of age-associated oxidative damage. This finding provides

6365-421: 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

6460-524: The idea is controversial, some evidence suggests a link between aging and mitochondrial genome dysfunction. In essence, mutations in mtDNA upset a careful balance of reactive oxygen species (ROS) production and enzymatic ROS scavenging (by enzymes like superoxide dismutase , catalase , glutathione peroxidase and others). However, some mutations that increase ROS production (e.g., by reducing antioxidant defenses) in worms increase, rather than decrease, their longevity. Also, naked mole rats , rodents about

6555-462: 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

6650-538: The introduction of Indo-Iranian speakers to Iran and South Asia", despite the absence of the characteristic timber graves of the steppe in the Near East, or south of the region between Kopet Dag and Pamir - Karakorum . Mallory acknowledges the difficulties of making a case for expansions from Andronovo to northern India, and that attempts to link the Indo-Aryans to such sites as the Beshkent and Vakhsh cultures "only gets

6745-654: The late period of Indo-Iranian linguistic unity, it is likely that in the later period they constituted a branch of the Iranians. According to Narasimhan et al. (2019), the expansion of the Andronovo culture towards the BMAC took place via the Inner Asia Mountain Corridor . According to Hiebert, an expansion of the BMAC into Iran and the margin of the Indus Valley is "the best candidate for an archaeological correlate of

6840-415: The levels of the mtDNA-encoded RNAs in bovine tissues has 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

6935-443: The longevity of species. The application of a mitochondrial-specific ROS scavenger, which lead to a significant longevity of the mice studied, suggests that mitochondria may still be well-implicated in ageing. Extensive research is being conducted to further investigate this link and methods to combat ageing. Presently, gene therapy and nutraceutical supplementation are popular areas of ongoing research. Bjelakovic et al. analyzed

7030-424: 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

7125-525: 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. 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

7220-469: The most notable remains are the burials of chariots, dating from around 2000 BC and possibly earlier. The chariots are found with paired horse-teams, and the ritual burial of the horse in a "head and hooves" cult has also been found. Some Andronovo dead were buried in pairs, of adults or adult and child. At Kytmanovo in Russia between Mongolia and Kazakhstan, dated 1746–1626 BC, a strain of Yersinia pestis

7315-480: The mtDNA base composition and animal species-specific maximum life spans. As demonstrated in their work, higher mtDNA guanine + cytosine content ( GC% ) strongly associates with longer maximum life spans across animal species. An additional observation is that the mtDNA GC% correlation with the maximum life spans is independent of the well-known correlation between animal species metabolic rate and maximum life spans. The mtDNA GC% and resting metabolic rate explain

7410-833: The mtDNA has approximately 10-fold higher levels than nuclear DNA. It has been proposed that aged mitochondria is the critical factor in the origin of neurodegeneration in Alzheimer's disease. Analysis of the brains of AD patients suggested an impaired function of the DNA repair pathway, which would cause reduce the overall quality of mtDNA. In Huntington's disease , mutant huntingtin protein causes mitochondrial dysfunction involving inhibition of mitochondrial electron transport , higher levels of reactive oxygen species and increased oxidative stress . Mutant huntingtin protein promotes oxidative damage to mtDNA, as well as nuclear DNA, that may contribute to Huntington's disease pathology . The DNA oxidation product 8-oxoguanine (8-oxoG)

7505-558: The mtDNAs were taken. mtDNA can be used to estimate the relationship between both closely related and distantly related species. Due to the high mutation rate of mtDNA in animals, the 3rd positions of the codons change relatively rapidly, and thus provide information about the genetic distances among closely related individuals or species. On the other hand, the substitution rate of mt-proteins is very low, thus amino acid changes accumulate slowly (with corresponding slow changes at 1st and 2nd codon positions) and thus they provide information about

7600-472: 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 the nucleus and is enabled by multiple copies of mtDNA present in mitochondria. The outcome of mutation in mtDNA may be an alteration in

7695-487: 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. The promoters for

7790-520: The person they accompanied throughout their lives, and they were used not only for food, but also for harnessing to vehicles and riding. Agriculture did not play an important role in the Andronovo economy. The Andronovo culture is notable for regional advances in metallurgy . They mined deposits of copper ore in the Altai Mountains from around the 14th century BC. Bronze objects were numerous, and workshops existed for working copper. One of

7885-466: 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 (type 4) while the other is a heterogeneous collection of linear molecules (type 6). Genome types 4 and 6 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,

7980-744: The preceding Sintashta culture were derived from an eastern migration of the Corded Ware culture , given the higher proportion of ancestry matching the earlier farmers of Europe, similar to the admixture found in the genomes of the Corded Ware population. The name derives from the village of Andronovo in the Uzhursky District of Kranoyarsk Krai, Siberia, where the Russian zoologist Arkadi Tugarinov discovered its first remains in 1914. Several graves were discovered, with skeletons in crouched positions, buried with richly decorated pottery. The Andronovo culture

8075-622: The presence of the culture in Xinjiang in far-western China has also been found, mainly concentrated in the area comprising Tashkurgan , Ili , Bortala , and Tacheng area. In the Volga basin, interaction with the Srubna culture was the most intense and prolonged, and Federovo style pottery is found as far west as Volgograd . Mallory notes that the Tazabagyab culture south of Andronovo could be an offshoot of

8170-495: 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. 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

8265-481: The remaining Öelet Oirats were deported to. The Nonni basin was where Oirat Öelet deportees were settled. The Yenisei Kyrgyz were deported along with the Öelet. Chinese and Oirat replaced Oirat and Kyrgyz during Manchukuo as the dual languages of the Nonni-based Yenisei Kyrgyz. In the 17th century, the Khakas formed Khakassia in the middle of the lands of Yenisei Kyrgyz, who at the time were vassals of

8360-409: The results of 78 studies between 1977 and 2012, involving a total of 296,707 participants, and concluded that antioxidant supplements do not reduce all-cause mortality nor extend lifespan, while some of them, such as beta carotene, vitamin E, and higher doses of vitamin A, may actually increase mortality. In a recent study, it was shown that dietary restriction can reverse ageing alterations by affecting

8455-442: 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 circular chromosomes (lengths 1556, 84 and 45 kilobases), which are entirely or largely autonomous with regard to their replication . Protists contain the most diverse mitochondrial genomes, with five different types found in this kingdom. Type 2, type 3 and type 5 of

8550-500: The size of mice , live about eight times longer than mice despite having reduced, compared to mice, antioxidant defenses and increased oxidative damage to biomolecules. Once, there was thought to be a positive feedback loop at work (a 'Vicious Cycle'); as mitochondrial DNA accumulates genetic damage caused by free radicals, the mitochondria lose function and leak free radicals into the cytosol . A decrease in mitochondrial function reduces overall metabolic efficiency. However, this concept

8645-410: Was conclusively disproved when it was demonstrated that mice, which were genetically altered to accumulate mtDNA mutations at accelerated rate do age prematurely, but their tissues do not produce more ROS as predicted by the 'Vicious Cycle' hypothesis. Supporting a link between longevity and mitochondrial DNA, some studies have found correlations between biochemical properties of the mitochondrial DNA and

8740-434: Was established in 1992. Khakas account for only about 12% of the total population of the republic (78,500 as of 1989 Census). Khakas traditionally practiced nomadic herding, agriculture, hunting, and fishing. The Beltir people specialized in handicraft as well. Herding sheep and cattle is still common, although the republic became more industrialized over time. Genetic research has identified 4 primary paternal lineages in

8835-423: Was extracted from a dead woman's tooth in a grave common to her and to two children. This strain's genes express flagellin , which triggers the human immune response. However, by contrast with other prehistoric Yersinia pestis bacteria, the strain does so weakly; later, historic plague does not express flagellin at all, accounting for its virulence. The Kytmanovo strain was therefore under selection toward becoming

8930-488: Was first identified by the Russian archaeologist Sergei Teploukhov in the 1920s. The culture of Sarazm (4th–3rd millennium BC) precedes the arrival of the Andronovo steppe culture in South Central Asia in the 2nd millennium BC. Currently only two sub-cultures are considered as part of Andronovo culture: Other authors identify the following sub-culture also as part of Andronovo: Some authors have challenged

9025-460: 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,

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