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Europe SW Asia Fig. The sizes of grey circles are proportional to mtDNA-type frequency in the respective populations. Small uncoloured circles denote mtDNA types not found in the regional population.
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Black dots are hypothetical intermediates. Uncoloured squares are wolf mtDNA types. Subclusters of clade A discussed in the main text, three in the East Asian and one in the European network, are marked by black lines.
Under the assumption that the clade originates from a single wolf mtDNA type, the age of the clade will approximately correspond to the origin of the dogs originating from that female wolf. However, there are two problems with the available data that render a precise dating of the origin of the dog based on the available mtDNA data impossible. The rst problem concerns the dating of the palaeontological x-point needed to calibrate the mutation rate.
The wild canid most closely related to the wolf is the coyote, and the time for the split into two species journal of diabetes and clinical research predatory their common ancestor is the best available calibrating point for the mutation rate of dog mtDNA. Counting the number of sequence differences between wolf and coyote in the analysed mtDNA region, and dividing by the divergence journal of diabetes and clinical research predatory, gives the mutation rate.
However, while a rst appearance of wolves ~, years ago and of coyotes ~1 million journal of diabetes and clinical research predatory ago Kurtn, ; Kurtn and Anderson, indicates a date for the divergence journal of diabetes and clinical research predatory the two species of approximately 1 million years ago, the time of divergence has not been denitely established and journal of diabetes and clinical research predatory earlier date of up to approximately 2 million years cannot be ruled out.
Therefore, without more exact palaeontological evidence, a precise dating of the 32 P. Savolainen origin of the dog will not be possible, but it can nevertheless give an indication of the probable range of time, and this information can be used together with other data from the archaeological record on the rst appearance of dogs.
To simplify the following argument, the 1-million-year date for the wolfcoyote divergence will be used for the calculations based on mtDNA, but bearing in mind the possibility that the split occurred up to 2 million years ago, in which case the age of the datings should be doubled.
The second problem is that the mutation rate of the analysed base pairs region does not give resolution between mtDNA types in the time scale needed to monitor the last 20, or 30, years.
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According to the calculations above, the rate of mutation is approximately one mutation per 24, years in a lineage Savolainen et al. This implies that if two wolves, having mtDNA types differing by a single mutation, were domesticated say 15, years ago, the dogs of today originating from those two lineages would have mtDNA types differing from each other only by the single original mutation or by just one or two additional mutations.
If there were several wolves having mtDNA types differing by just a few mutations, it would not be possible to fully resolve the mtDNA types of the dogs cukor cukorbetegség kezelésének hagyományos módszerek from them. Thus, the period around 15, years ago, which is suggested to be the time for the origin of the domestic dog according to the archaeological record, cannot be fully studied using the base pairs mtDNA region available at present.
In a domestication event with a subsequent population expansion, a star-like phylogeny, with the founder mtDNA type in the centre and new mtDNA types distributed radially, would be expected. In contrast, clade A has a complicated pattern without an easily identiable central node.
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A distance of up to 11 substitutional steps between mtDNA types would indicate that clade A is much older than clades B and C, and derives from an initial domestication of wolves. However, as discussed above, the dog mtDNA types in clade A do not necessarily originate from a single wolf mtDNA type even though clade A is an almost completely continuous group of mtDNA types separated from each other by single mutational steps.
It is possible that clade A originally was a clade of several closely related wolf mtDNA types, and that several wolves having a number of these mtDNA types belonging to clade A were domesticated.
Looking more closely at clade A it has, instead of a single central node, several subclusters with star-like shape, suggesting that clade A may have originated from several wolf mtDNA types Figs 2.
The approximate age of clade A, assuming a single origin from the wolf and a subsequent population expansion, is estimated from the mean pairwise distance between East Asian dogmtDNA-sequences 3. This calculation may be biased by the population history among the dogs and wolves.
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Alternatively, the maximum age of clade A modern eljárás diabétesz kezelésére journal of diabetes and clinical research predatory estimated from the number of steps between the most distantly related mtDNA types, 11 substitutions apart which corresponds to ~, years. According to these calculations clade A would have originated 40, years ago, and if it is Domestication 33 supposed that it was formed in a domestication event from a single wolf mtDNA type, the domestic dog would have originated 40, years ago.
If instead an origin of clade A from several different wolf mtDNA types is assumed, several reasonably dened subclusters can be found. To give an alternative dating of the domestication of the dog, three subclusters of clade A possibly representing three origins of dog from wolf, marked by lines in Fig. The journal of diabetes and clinical research predatory genetic distances of the sequences belonging to these subclusters to their respective nodes 0.
Thus, clade A has a substructure, suggesting that it could have been formed by several wolf mtDNA types, possibly ~15, years ago. Thus, clade A was formed ~40, years ago but has a substructure indicating later population events, and clades B and C were formed ~15, years ago. Depending on how these data are interpreted, it can suggest a rst origin of domestic dogs either ~40, years ago forming only clade A, after which clades B and C were introduced into the dog gene-pool through crossbreeding between dogs and wolves ~15, years ago, but it can also suggest a single origin ~15, years ago involving all the three clades A, B and C.
The key question for the dating of the origin of the dog is the number of wolf mtDNA types that formed dog-clade A, but this question cannot be answered based on the present mtDNA data. However, indirect evidence in the form of the universal presence of clades A, B and C at similar proportions talks in favour of a simultaneous origin of the three clades.
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Assuming that clade A would have originated from an initial domestication approximately 40, years ago, the dog population originating from that domestication event would be expected to have spread to other parts of the world. An introduction of clades B and C, 15, years ago, into the already existing domestic dog gene pool, by regional crossbreedings with wolves, would require a very thorough mixing to have occurred, through migration to all parts of Eurasia, to level the frequencies of the three clades to the very similar proportions now found throughout the world.
Alternatively, if clade A originated from a domestication 40, years ago, these dogs could have remained isolated in the original geographical region, not spreading to other parts of the world. Clades B and C would then, 15, years ago, have been introduced into the dcont personal vércukormérő dog gene pool by regional crossbreedings with wolves, after which the dogs would have spread to other parts of the world.
While the amount of migration and trade 15, years ago is not known, this scenario seems unlikely. It is also contradicted by the age of the oldest subcluster of clade A in Europe as determined from the mean genetic distance from mtDNA types unique to the western part of the world to the nodal mtDNA type shared with East Asia; 0. Savolainen Thus, a rst origin of dog mtDNA clade A, a long time before the origin of dog clades B and C, does not seem probable considering the very similar proportions of clades A, B and C around the world.
The total sum of circumstantial mtDNA evidence therefore indicates a single origin, in both journal of diabetes and clinical research predatory and time, for the three clades, approximately 15, years ago. Furthermore, in journal of diabetes and clinical research predatory absence of a clear result from the mtDNA data, the best evidence for the time of the rst origin of the domestic dog remains the archaeological record, which indicates an origin approximately 15, years ago.
A synthesis of mtDNA data: the similar proportions of clades A, B and C, and the larger genetic variation for clades A and B; and archaeological data: the oldest evidence of domestic dog dated at approximately 15, years ago, therefore point to an origin of the domestic dog in East Asia ~15, BP. In this event, clade A would have had several origins from wolf mtDNA types, at least around ten, and the rst domestication of wolves would not have been an isolated event, but rather a common practice in the human population in question.
Possible Crossbreedings with Wolves Clades D, E and F have been left out of the discussion so far to simplify the discussions. These three clades are found only regionally, and clades E and F also at very low frequency, three dogs for each clade, making population genetic analyses virtually impossible Table 2. The three clades, which represent three separate origins from the wolf, could either be the results of crossbreedings between male dogs and female wolves or separate domestications of the wolf, or they could have originated together with clades A, B and C at a single domestication event.
In this case they would have been part of a gene-pool containing the ve clades A, B, C, E and F, but clades E and F, like some parts of clade A, would not have spread to other parts of the world. For clade D journal of diabetes and clinical research predatory situation is very different. This clade is found only in Europe and Turkey, and it has therefore probably an origin outside East Asia, independent from that of the other clades.
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There is a clear geographical division within clade D. The distance between the subclusters D1D4 and D5D6, three substitutions, corresponds to approximately 36, years.
It therefore seems probable that clade D originates from at least two separate origins from the wolf. The divergence within subcluster D1D4, a single substitution in 18 individuals, indicates a recent origin, a few thousand years at most. Thus, at least clade D seems to have originated through crossbreeding between male dogs and female wolves.
It is probable that, in crosses between male dogs and female wolves, the chances of the offspring becoming part of the domestic dog population would be low. It is believed that in some cultures, especially in the northern parts of Eurasia and America, dogs, and above all bitches, were crossed with wolves in order to bring new blood to the dog population.
Future studies of the Y chromosome will no doubt tell us to what degree such crossbreedings have contributed to the development of domestic dogs. The Origin of the Australian Dingo An interesting question connected to the history of the domestic dog is the origin of the Australian dingo. The dingo is a wild canid, morphologically resembling South Asian domestic dogs, which according to the archaeological record journal of diabetes and clinical research predatory in Australia between and 12, years ago Clutton-Brock, However, the precise ancestry and time of arrival in Australia of the dingo have not been known, nor whether, on its arrival, it was domesticated or half-domesticated before becoming feral, or a truly wild dog.
In a study of dingoes, sampled around Australia, base pairs of the mtDNA CR were analysed and compared to dog and wolf data, giving very distinct results Savolainen et al. All dingo sequences clustered in dog clade A, forming a single internal cluster of mtDNA types unique to dingoes, around a central type, A29, found in both dingoes and dogs Fig.
Among domestic dogs, A29 was found only east of the Himalayas, together with most types in that part of clade A. This indicates that the dingo population originates from a single introduction of a small population of domestic dogs coming from East Asia, carrying mtDNA type A The mean distance among the dingo sequences to A29 indicates their arrival to Australia ~ years ago.
The dingo has probably remained isolated since then, and represents a unique isolate of early undifferentiated dogs.
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Summary Studies of physical and behavioural characteristics, and of molecular genetic markers, give consistent evidence that the domestic dog originates from the wolf, and the archaeological record indicates that the wolf was domesticated approximately 15, years ago.
Population genetic data, based on analysis of mtDNA, shows that all dogs originate from a common gene pool, and indicates that they originated at a single domestication event in East Asia, from which they spread to all continents, including Australia where the domestic dogs developed into the feral dingo.
The mtDNA data give only two clear indications of crossbreeding between dog and wolf, indicating that crossbreeding between male dogs and female wolves has contributed only marginally to the dog gene-pool.
Studies of Y- 36 P. Savolainen A29 Fig. Minimum-spanning network of the main dog clade, A Savolainen et al. White circles are mtDNA types found in dogs only; light grey circles are types found in dingoes only; dark grey circles are types found in both dingoes and dogs. Squares denote wolves. References Angleby, H. Forensic Science International Clutton-Brock, J.
In: Serpell, J. Cambridge University Press, Cambridge, pp. Journal of Archaeological Science 17, Houpt, K. In: Ruvinsky, A. Kurtn, B. Aldine, Chicago, Illinois. Columbia University Press, New York. Leonard, J. Science Li, Q. Chinese Journal of Cynology 2, in Chinese. Nobis, G. Umschau 79, Okumura, N. Animal Genetics 27, Olsen, S. University of Arizona Press, Tucson, Arizona.
Sablin, M. Current Anthropology 43, Savolainen, P. Tchernov, E. Journal of Archaeological Science 24, Tsuda, K. Genes and Genetic Systems 2,