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1

Bennett, E. Andrew, Isabelle Crevecoeur, Bence Viola, Anatoly P. Derevianko, Michael V. Shunkov, Thierry Grange, Bruno Maureille, and Eva-Maria Geigl. "Morphology of the Denisovan phalanx closer to modern humans than to Neanderthals." Science Advances 5, no. 9 (September 2019): eaaw3950. http://dx.doi.org/10.1126/sciadv.aaw3950.

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A fully sequenced high-quality genome has revealed in 2010 the existence of a human population in Asia, the Denisovans, related to and contemporaneous with Neanderthals. Only five skeletal remains are known from Denisovans, mostly molars; the proximal fragment of a fifth finger phalanx used to generate the genome, however, was too incomplete to yield useful morphological information. Here, we demonstrate through ancient DNA analysis that a distal fragment of a fifth finger phalanx from the Denisova Cave is the larger, missing part of this phalanx. Our morphometric analysis shows that its dimensions and shape are within the variability of Homo sapiens and distinct from the Neanderthal fifth finger phalanges. Thus, unlike Denisovan molars, which display archaic characteristics not found in modern humans, the only morphologically informative Denisovan postcranial bone identified to date is suggested here to be plesiomorphic and shared between Denisovans and modern humans.
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2

Akkuratov, Evgeny E., Mikhail S. Gelfand, and Ekaterina E. Khrameeva. "Neanderthal and Denisovan ancestry in Papuans: A functional study." Journal of Bioinformatics and Computational Biology 16, no. 02 (April 2018): 1840011. http://dx.doi.org/10.1142/s0219720018400115.

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Sequencing of complete nuclear genomes of Neanderthal and Denisovan stimulated studies about their relationship with modern humans demonstrating, in particular, that DNA alleles from both Neanderthal and Denisovan genomes are present in genomes of modern humans. The Papuan genome is a unique object because it contains both Neanderthal and Denisovan alleles. Here, we have shown that the Papuan genomes contain different gene functional groups inherited from each of the ancient people. The Papuan genomes demonstrate a relative prevalence of Neanderthal alleles in genes responsible for the regulation of transcription and neurogenesis. The enrichment of specific functional groups with Denisovan alleles is less pronounced; these groups are responsible for bone and tissue remodeling. This analysis shows that introgression of alleles from Neanderthals and Denisovans to Papuans occurred independently and retention of these alleles may carry specific adaptive advantages.
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3

Rogers, Alan R., Nathan S. Harris, and Alan A. Achenbach. "Neanderthal-Denisovan ancestors interbred with a distantly related hominin." Science Advances 6, no. 8 (February 2020): eaay5483. http://dx.doi.org/10.1126/sciadv.aay5483.

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Previous research has shown that modern Eurasians interbred with their Neanderthal and Denisovan predecessors. We show here that hundreds of thousands of years earlier, the ancestors of Neanderthals and Denisovans interbred with their own Eurasian predecessors—members of a “superarchaic” population that separated from other humans about 2 million years ago. The superarchaic population was large, with an effective size between 20 and 50 thousand individuals. We confirm previous findings that (i) Denisovans also interbred with superarchaics, (ii) Neanderthals and Denisovans separated early in the middle Pleistocene, (iii) their ancestors endured a bottleneck of population size, and (iv) the Neanderthal population was large at first but then declined in size. We provide qualified support for the view that (v) Neanderthals interbred with the ancestors of modern humans.
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4

Zhang, Dongju, Huan Xia, Fahu Chen, Bo Li, Viviane Slon, Ting Cheng, Ruowei Yang, et al. "Denisovan DNA in Late Pleistocene sediments from Baishiya Karst Cave on the Tibetan Plateau." Science 370, no. 6516 (October 29, 2020): 584–87. http://dx.doi.org/10.1126/science.abb6320.

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A late Middle Pleistocene mandible from Baishiya Karst Cave (BKC) on the Tibetan Plateau has been inferred to be from a Denisovan, an Asian hominin related to Neanderthals, on the basis of an amino acid substitution in its collagen. Here we describe the stratigraphy, chronology, and mitochondrial DNA extracted from the sediments in BKC. We recover Denisovan mitochondrial DNA from sediments deposited ~100 thousand and ~60 thousand years ago (ka) and possibly as recently as ~45 ka. The long-term occupation of BKC by Denisovans suggests that they may have adapted to life at high altitudes and may have contributed such adaptations to modern humans on the Tibetan Plateau.
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5

Schwartz, K., and M. Sorokin. "The human genome reveals the evolution of Homo sapiens." BULLETIN of the L.N. Gumilyov Eurasian National University. BIOSCIENCE Series 134, no. 1 (2021): 38–45. http://dx.doi.org/10.32523/2616-7034-2021-134-1-38-45.

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The evolution of modern humans began two and a half million years ago as Homo erectus. Several hundred thousand years ago, Neanderthals, Denisovans, and modern men Homo sapiens have been separated from the Homo erectus branch. Nevertheless, Homo sapiens is the only one that has survived to our days. The complex history of Homo is revealed by genetic research and comparison of the modern human genome with genes of Neanderthals and Denisovans. Svante Pääbo, a professor at the Max Planck Institute for Evolutionary Anthropology, made a significant contribution to these studies and decoded the genome of Neanderthals and Denisovans. Comparison of the genome of modern humans with the genes of Neanderthals and Denisovans made it possible to reveal the size of the population, the paths and times of migrations, interactions of various groups of ancient humans and their biological crossing. It was found that in Eurasia, modern man carries traces of Neanderthal genes, whereas in Asia and Oceania – Denisovan genes. According to anthropological research, the survival of Homo sapiens was driven by the cognitive revolution, which took place about seventy thousand years ago and included the development of language, communication and association in large groups.
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6

Zhang, Xinjun, Kelsey E. Witt, Mayra M. Bañuelos, Amy Ko, Kai Yuan, Shuhua Xu, Rasmus Nielsen, and Emilia Huerta-Sanchez. "The history and evolution of the Denisovan-EPAS1 haplotype in Tibetans." Proceedings of the National Academy of Sciences 118, no. 22 (May 28, 2021): e2020803118. http://dx.doi.org/10.1073/pnas.2020803118.

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Recent studies suggest that admixture with archaic hominins played an important role in facilitating biological adaptations to new environments. For example, interbreeding with Denisovans facilitated the adaptation to high-altitude environments on the Tibetan Plateau. Specifically, the EPAS1 gene, a transcription factor that regulates the response to hypoxia, exhibits strong signatures of both positive selection and introgression from Denisovans in Tibetan individuals. Interestingly, despite being geographically closer to the Denisova Cave, East Asian populations do not harbor as much Denisovan ancestry as populations from Melanesia. Recently, two studies have suggested two independent waves of Denisovan admixture into East Asians, one of which is shared with South Asians and Oceanians. Here, we leverage data from EPAS1 in 78 Tibetan individuals to interrogate which of these two introgression events introduced the EPAS1 beneficial sequence into the ancestral population of Tibetans, and we use the distribution of introgressed segment lengths at this locus to infer the timing of the introgression and selection event. We find that the introgression event unique to East Asians most likely introduced the beneficial haplotype into the ancestral population of Tibetans around 48,700 (16,000–59,500) y ago, and selection started around 9,000 (2,500–42,000) y ago. Our estimates suggest that one of the most convincing examples of adaptive introgression is in fact selection acting on standing archaic variation.
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7

Zavala, Elena I., Zenobia Jacobs, Benjamin Vernot, Michael V. Shunkov, Maxim B. Kozlikin, Anatoly P. Derevianko, Elena Essel, et al. "Pleistocene sediment DNA reveals hominin and faunal turnovers at Denisova Cave." Nature 595, no. 7867 (June 23, 2021): 399–403. http://dx.doi.org/10.1038/s41586-021-03675-0.

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AbstractDenisova Cave in southern Siberia is the type locality of the Denisovans, an archaic hominin group who were related to Neanderthals1–4. The dozen hominin remains recovered from the deposits also include Neanderthals5,6 and the child of a Neanderthal and a Denisovan7, which suggests that Denisova Cave was a contact zone between these archaic hominins. However, uncertainties persist about the order in which these groups appeared at the site, the timing and environmental context of hominin occupation, and the association of particular hominin groups with archaeological assemblages5,8–11. Here we report the analysis of DNA from 728 sediment samples that were collected in a grid-like manner from layers dating to the Pleistocene epoch. We retrieved ancient faunal and hominin mitochondrial (mt)DNA from 685 and 175 samples, respectively. The earliest evidence for hominin mtDNA is of Denisovans, and is associated with early Middle Palaeolithic stone tools that were deposited approximately 250,000 to 170,000 years ago; Neanderthal mtDNA first appears towards the end of this period. We detect a turnover in the mtDNA of Denisovans that coincides with changes in the composition of faunal mtDNA, and evidence that Denisovans and Neanderthals occupied the site repeatedly—possibly until, or after, the onset of the Initial Upper Palaeolithic at least 45,000 years ago, when modern human mtDNA is first recorded in the sediments.
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8

Sawyer, Susanna, Gabriel Renaud, Bence Viola, Jean-Jacques Hublin, Marie-Theres Gansauge, Michael V. Shunkov, Anatoly P. Derevianko, Kay Prüfer, Janet Kelso, and Svante Pääbo. "Nuclear and mitochondrial DNA sequences from two Denisovan individuals." Proceedings of the National Academy of Sciences 112, no. 51 (November 16, 2015): 15696–700. http://dx.doi.org/10.1073/pnas.1519905112.

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Denisovans, a sister group of Neandertals, have been described on the basis of a nuclear genome sequence from a finger phalanx (Denisova 3) found in Denisova Cave in the Altai Mountains. The only other Denisovan specimen described to date is a molar (Denisova 4) found at the same site. This tooth carries a mtDNA sequence similar to that of Denisova 3. Here we present nuclear DNA sequences from Denisova 4 and a morphological description, as well as mitochondrial and nuclear DNA sequence data, from another molar (Denisova 8) found in Denisova Cave in 2010. This new molar is similar to Denisova 4 in being very large and lacking traits typical of Neandertals and modern humans. Nuclear DNA sequences from the two molars form a clade with Denisova 3. The mtDNA of Denisova 8 is more diverged and has accumulated fewer substitutions than the mtDNAs of the other two specimens, suggesting Denisovans were present in the region over an extended period. The nuclear DNA sequence diversity among the three Denisovans is comparable to that among six Neandertals, but lower than that among present-day humans.
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9

Rogers, Alan R., Ryan J. Bohlender, and Chad D. Huff. "Early history of Neanderthals and Denisovans." Proceedings of the National Academy of Sciences 114, no. 37 (August 7, 2017): 9859–63. http://dx.doi.org/10.1073/pnas.1706426114.

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Extensive DNA sequence data have made it possible to reconstruct human evolutionary history in unprecedented detail. We introduce a method to study the past several hundred thousand years. Our results show that (i) the Neanderthal–Denisovan lineage declined to a small size just after separating from the modern lineage, (ii) Neanderthals and Denisovans separated soon thereafter, and (iii) the subsequent Neanderthal population was large and deeply subdivided. They also (iv) support previous estimates of gene flow from Neanderthals into modern Eurasians. These results suggest an archaic human diaspora early in the Middle Pleistocene.
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10

Lodewijk, Gerrald A., Diana P. Fernandes, Iraklis Vretzakis, Jeanne E. Savage, and Frank M. J. Jacobs. "Evolution of Human Brain Size-Associated NOTCH2NL Genes Proceeds toward Reduced Protein Levels." Molecular Biology and Evolution 37, no. 9 (April 24, 2020): 2531–48. http://dx.doi.org/10.1093/molbev/msaa104.

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Abstract Ever since the availability of genomes from Neanderthals, Denisovans, and ancient humans, the field of evolutionary genomics has been searching for protein-coding variants that may hold clues to how our species evolved over the last ∼600,000 years. In this study, we identify such variants in the human-specific NOTCH2NL gene family, which were recently identified as possible contributors to the evolutionary expansion of the human brain. We find evidence for the existence of unique protein-coding NOTCH2NL variants in Neanderthals and Denisovans which could affect their ability to activate Notch signaling. Furthermore, in the Neanderthal and Denisovan genomes, we find unusual NOTCH2NL configurations, not found in any of the modern human genomes analyzed. Finally, genetic analysis of archaic and modern humans reveals ongoing adaptive evolution of modern human NOTCH2NL genes, identifying three structural variants acting complementary to drive our genome to produce a lower dosage of NOTCH2NL protein. Because copy-number variations of the 1q21.1 locus, encompassing NOTCH2NL genes, are associated with severe neurological disorders, this seemingly contradicting drive toward low levels of NOTCH2NL protein indicates that the optimal dosage of NOTCH2NL may have not yet been settled in the human population.
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11

Derevianko, O. P., M. V. Shunkov, and M. B. Kozlikin. "Who Were the Denisovans?" Archaeology, Ethnology & Anthropology of Eurasia 48, no. 3 (October 4, 2020): 3–32. http://dx.doi.org/10.17746/1563-0110.2020.48.3.003-032.

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We provide a comprehensive summary of data relating to the origin, chronology, and culture of the Denisovans— a separate hominin population, fi rst described in 2010 on the basis of aDNA extracted from fossils found in Denisova Cave, in the northwestern part of the Russian Altai. We cite the results of morphological and genomic studies of the teeth and postcranial bones of those hominins. On the basis of a large series of optical and radiocarbon dates of the Pleistocene strata of Denisova Cave, the timeline for the hominin evolution in that region is reconstructed. The chronology of the evolutionary events based on aDNA is discussed. We provide a detailed description of stone and bone tools, and ornaments made of various materials, from Denisova habitation horizons. It is demonstrated that the Paleolithic cultural sequence in that cave is the most complete in North and Central Asia, spanning the principal stages of human evolutionary history over the last 300 thousand years. Denisovan origins and their role in the emergence of anatomically modern humans are reconstructed on the basis of a large body of archaeological, skeletal, and genetic data relating to Africa and Eurasia. It is concluded that the Neanderthal and Denisovan genetic legacy in the modern human gene pool indicates the existence of several zones in Africa and Eurasia where H. erectus evolution proceeded independently. The same applies to the evolution of lithic technologies.
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12

Bailey, Shara E., Jean-Jacques Hublin, and Susan C. Antón. "Rare dental trait provides morphological evidence of archaic introgression in Asian fossil record." Proceedings of the National Academy of Sciences 116, no. 30 (July 8, 2019): 14806–7. http://dx.doi.org/10.1073/pnas.1907557116.

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The recently described Denisovan hemimandible from Xiahe, China [F. Chen et al., (2019) Nature 569, 409–412], possesses an unusual dental feature: a 3-rooted lower second molar. A survey of the clinical and bioarchaeological literature demonstrates that the 3-rooted lower molar is rare (less than 3.5% occurrence) in non-Asian Homo sapiens. In contrast, its presence in Asian-derived populations can exceed 40% in China and the New World. It has long been thought that the prevalence of 3-rooted lower molars in Asia is a relatively late acquisition occurring well after the origin and dispersal of H. sapiens. However, the presence of a 3-rooted lower second molar in this 160,000-y-old fossil hominin suggests greater antiquity for the trait. Importantly, it also provides morphological evidence of a strong link between archaic and recent Asian H. sapiens populations. This link provides compelling evidence that modern Asian lineages acquired the 3-rooted lower molar via introgression from Denisovans.
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13

Wilson, Clare. "Denisovans in Tibet." New Scientist 242, no. 3228 (May 2019): 8. http://dx.doi.org/10.1016/s0262-4079(19)30755-9.

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14

Stringer, Chris B., and Ian Barnes. "Deciphering the Denisovans." Proceedings of the National Academy of Sciences 112, no. 51 (December 14, 2015): 15542–43. http://dx.doi.org/10.1073/pnas.1522477112.

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15

George, Alison. "Denisovan art uncovered?" New Scientist 243, no. 3240 (July 2019): 13. http://dx.doi.org/10.1016/s0262-4079(19)31358-2.

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16

Barras, Colin. "Denisovan tools unearthed." New Scientist 245, no. 3272 (March 2020): 14. http://dx.doi.org/10.1016/s0262-4079(20)30482-6.

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17

Villanea, Fernando A., Emilia Huerta-Sanchez, and Keolu Fox. "ABO Genetic Variation in Neanderthals and Denisovans." Molecular Biology and Evolution 38, no. 8 (April 23, 2021): 3373–82. http://dx.doi.org/10.1093/molbev/msab109.

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Abstract Variation at the ABO locus was one of the earliest sources of data in the study of human population identity and history, and to this day remains widely genotyped due to its importance in blood and tissue transfusions. Here, we look at ABO blood type variants in our archaic relatives: Neanderthals and Denisovans. Our goal is to understand the genetic landscape of the ABO gene in archaic humans, and how it relates to modern human ABO variation. We found two Neanderthal variants of the O allele in the Siberian Neanderthals (O1 and O2), one of these variants is shared with an European Neanderthal, who is a heterozygote for this O1 variant and a rare cis-AB variant. The Denisovan individual is heterozygous for two variants of the O1 allele, functionally similar to variants found widely in modern humans. Perhaps more surprisingly, the O2 allele variant found in Siberian Neanderthals can be found at low frequencies in modern Europeans and Southeast Asians, and the O1 allele variant found in Siberian and European Neanderthal is also found at very low frequency in modern East Asians. Our genetic distance analyses suggest both alleles survive in modern humans due to inbreeding with Neanderthals. We find that the sequence backgrounds of the surviving Neanderthal-like O alleles in modern humans retain a higher sequence divergence than other surviving Neanderthal genome fragments, supporting a view of balancing selection operating in the Neanderthal ABO alleles by retaining highly diverse haplotypes compared with portions of the genome evolving neutrally.
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18

Derevianko, A. P., M. V. Shunkov, and M. B. Kozlikin. "Who Were the Denisovans?" Archaeology, Ethnology and Anthropology of Eurasia (Russian-language) 48, no. 3 (2020): 3–32. http://dx.doi.org/10.17746/1563-0102.2020.48.3.003-032.

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19

Gibbons, A. "Who Were the Denisovans?" Science 333, no. 6046 (August 25, 2011): 1084–87. http://dx.doi.org/10.1126/science.333.6046.1084.

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20

Chen, Fahu, Huan Xia, and Dongju Zhang. "A review of Denisovans." Chinese Science Bulletin 65, no. 25 (April 14, 2020): 2763–74. http://dx.doi.org/10.1360/tb-2020-0280.

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21

Otto, Grant. "Archaic admixture with Denisovans." Nature Reviews Genetics 19, no. 5 (April 4, 2018): 251. http://dx.doi.org/10.1038/nrg.2018.18.

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22

Massilani, Diyendo, Laurits Skov, Mateja Hajdinjak, Byambaa Gunchinsuren, Damdinsuren Tseveendorj, Seonbok Yi, Jungeun Lee, et al. "Denisovan ancestry and population history of early East Asians." Science 370, no. 6516 (October 29, 2020): 579–83. http://dx.doi.org/10.1126/science.abc1166.

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We present analyses of the genome of a ~34,000-year-old hominin skull cap discovered in the Salkhit Valley in northeastern Mongolia. We show that this individual was a female member of a modern human population that, following the split between East and West Eurasians, experienced substantial gene flow from West Eurasians. Both she and a 40,000-year-old individual from Tianyuan outside Beijing carried genomic segments of Denisovan ancestry. These segments derive from the same Denisovan admixture event(s) that contributed to present-day mainland Asians but are distinct from the Denisovan DNA segments in present-day Papuans and Aboriginal Australians.
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23

Slon, Viviane, Bence Viola, Gabriel Renaud, Marie-Theres Gansauge, Stefano Benazzi, Susanna Sawyer, Jean-Jacques Hublin, et al. "A fourth Denisovan individual." Science Advances 3, no. 7 (July 2017): e1700186. http://dx.doi.org/10.1126/sciadv.1700186.

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24

Agoni, Lorenzo, Aaron Golden, Chandan Guha, and Jack Lenz. "Neandertal and Denisovan retroviruses." Current Biology 22, no. 11 (June 2012): R437—R438. http://dx.doi.org/10.1016/j.cub.2012.04.049.

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25

Tashi, Tsewang, N. Scott Reading, Anna Shestakova, Tatiana Burjanivova, Tanna Uran, Felipe R. Lorenzo V, Hao Hu, et al. "Tibetan Gain-of-Function Variant of Prolyl Hydroxylase 2 (EGLN1) and Selected SNPs of HIF-2-Alpha (EPAS1) Are Associated with Lower Hemoglobin Values in Tibetans." Blood 126, no. 23 (December 3, 2015): 3332. http://dx.doi.org/10.1182/blood.v126.23.3332.3332.

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Abstract Hypoxia is the principal driver of erythropoiesis. Tibetan highlanders are protected from polycythemia despite living at high altitude. Whole genome sequencing studies narrowed evolutionary selected Tibetan haplotypes to two loci, EGLN1 and EPAS1 (Huerta-Sanchez et al, Nature, 2014, 512:194). Our previous work identified a functional variant of the principal negative regulator of hypoxia inducible transcription factors (HIFs), prolyl hydroxylase 2 (PHD2D4E/C127S, encoded by EGLN112C>G,380G>C), that has a lower Km for oxygen, thereby explaining the lack of polycythemia, and is present in ~88% of Tibetans (Lorenzo FR et al, Nat Genet. 2014, 46:951). Additionally, a highly selected Tibetan EPAS1 (encoding HIF-2a) haplotype has introgressed from prehistoric Denisovans (Huerta-Sanchez et al, Nature, 2014, 512:194). This Tibetan EPAS1 haplotype has no exonic variants but a unique epigenetic profile when analyzed by ENCODE software. We analyzed genomes of 260 healthy adult Tibetans residing in different altitudes in China, India and the US with appropriate IRB approvals and informed consents. Since EGLN112C>G,380G>C is located in a GC rich region that is not easily detectable by NGS and Sanger sequencing, we developed a high resolution melting assay for its detection. For the EPAS1 haplotype we analyzed 10 SNPs (5 Denisovan and 5 non-Denisovan, each with unique linkage disequilibrium (LD) value) in its evolutionary selected Tibetan region and correlated them with hemoglobin levels. We found that EGLN112C>G (D4E) is always in cis (i.e complete LD) with the EGLN1380G>C (C127S) missense variant, constituting a unique Tibetan-specific positively selected haplotype. The prevalence of PHD2D4E/C127S variants increased with altitude, seen in 52 out of 55 individuals (94.5%) at altitudes above 4000 m, consistent with positive selection of PHD2D4E/C127S at high altitude. Across all altitudes, after adjusting for age, gender and MCHC, the mean hemoglobin level was lower in PHD2D4E/C127S heterozygotes (14.4 g/dl; p=0.009), and tended to be lower in homozygotes (14.7 g/dl; p=0.10) than in PHD2 wild-type subjects (15.2 g/dl). Combining the homozygotes and heterozygotes, the hemoglobin was lower (14.6 g/dl; p=0.015) compared to PHD2 wild-type subjects. This lower hemoglobin level in those with PHD2D4E/C127S variants also extended to the subset of individuals residing at low altitudes (200 m). Thus, blunting of the HIF pathway by PHD2D4E/C127S persists even at the ambient oxygen of lower altitudes, although none these low-altitude residents were found to be anemic. We then evaluated contributions of EPAS1 SNPs to hemoglobin as the response variable using linear regression controlled for age, sex, altitude and EGLN1 genotype. Five of the 10 selected EPAS1 SNPs remained significant after multiple testing corrections (FDR <0.05). At low altitude these SNPs are associated with lower hemoglobin concentration, but the, number of Tibetans with wild-type EGLN1 and EPAS1 haplotypes at higher altitude (>3,500 m; ~11,500 feet) was not sufficient for analysis. In conclusion, we show that the gain-of-function PHD2D4E/C127S is a Tibetan specific variant that forms a part of the genetic basis of high altitude adaption by lowering hemoglobin at all altitudes. Tibetans have a unique EPAS1 haplotype introgressed from prehistoric Denisovans, but no coding mutations in this EPAS1 haplotype were found. However, selected SNPs in this Tibetan EPAS1 haplotypes also contribute to their decreased hemoglobin. At the time of writing of this report we have obtained samples from additional 110 Tibetan subjects from Yushu area of Tibetan Plateau residing at altitude 3500 meters and higher (average 3700m). This will permit more rigorous analyses of combined effect of these two EGLN1 and EPAS1 positively selected Tibetan variants on hemoglobin levels at various altitudes. Our investigation of epigenetic alterations of the DNase hypersensitive regions and methylated regions of the Tibetan selected EPAS1 haplotype and the effect of this EPAS1 haplotype on transcriptome and determination of its functional relevance is in progress. Disclosures No relevant conflicts of interest to declare.
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26

Petr, Martin, Mateja Hajdinjak, Qiaomei Fu, Elena Essel, Hélène Rougier, Isabelle Crevecoeur, Patrick Semal, et al. "The evolutionary history of Neanderthal and Denisovan Y chromosomes." Science 369, no. 6511 (September 24, 2020): 1653–56. http://dx.doi.org/10.1126/science.abb6460.

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Ancient DNA has provided new insights into many aspects of human history. However, we lack comprehensive studies of the Y chromosomes of Denisovans and Neanderthals because the majority of specimens that have been sequenced to sufficient coverage are female. Sequencing Y chromosomes from two Denisovans and three Neanderthals shows that the Y chromosomes of Denisovans split around 700 thousand years ago from a lineage shared by Neanderthals and modern human Y chromosomes, which diverged from each other around 370 thousand years ago. The phylogenetic relationships of archaic and modern human Y chromosomes differ from the population relationships inferred from the autosomal genomes and mirror mitochondrial DNA phylogenies, indicating replacement of both the mitochondrial and Y chromosomal gene pools in late Neanderthals. This replacement is plausible if the low effective population size of Neanderthals resulted in an increased genetic load in Neanderthals relative to modern humans.
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27

Zhang, Dongju, Huan Xia, Ting Cheng, and Fahu Chen. "New portraits of the Denisovans." Science Bulletin 65, no. 1 (January 2020): 1–3. http://dx.doi.org/10.1016/j.scib.2019.10.013.

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28

Mao, Steve. "Denisovans shaped our genomes, twice." Science 360, no. 6385 (April 12, 2018): 167.4–168. http://dx.doi.org/10.1126/science.360.6385.167-d.

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29

SONG, Jianlan. "Denisovan from Qinghai-Tibet Plateau." Bulletin of the Chinese Academy of Sciences 34, no. 1 (January 1, 2020): 14–15. http://dx.doi.org/10.3724/sp.j.7101866520.

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30

Barras, Colin. "Milk tooth reveals oldest Denisovan." New Scientist 235, no. 3134 (July 2017): 16. http://dx.doi.org/10.1016/s0262-4079(17)31359-3.

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31

Zahn, L. M. "Denisovan DNA retained in Melanesians." Science 352, no. 6282 (April 7, 2016): 183. http://dx.doi.org/10.1126/science.352.6282.183-c.

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32

Lee, A., D. Huntley, P. Aiewsakun, R. K. Kanda, C. Lynn, and M. Tristem. "Novel Denisovan and Neanderthal Retroviruses." Journal of Virology 88, no. 21 (August 20, 2014): 12907–9. http://dx.doi.org/10.1128/jvi.01825-14.

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33

Gibbons, A. "Cave was lasting home to Denisovans." Science 349, no. 6254 (September 17, 2015): 1270–71. http://dx.doi.org/10.1126/science.349.6254.1270-b.

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34

Cooper, A., and C. B. Stringer. "Did the Denisovans Cross Wallace's Line?" Science 342, no. 6156 (October 17, 2013): 321–23. http://dx.doi.org/10.1126/science.1244869.

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35

Hawks, John. "Neanderthals and Denisovans as biological invaders." Proceedings of the National Academy of Sciences 114, no. 37 (August 31, 2017): 9761–63. http://dx.doi.org/10.1073/pnas.1713163114.

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36

Price, Michael. "Face of the mysterious Denisovans emerges." Science 365, no. 6459 (September 19, 2019): 1232. http://dx.doi.org/10.1126/science.365.6459.1232.

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37

Guimarães, Santiago Wolnei Ferreira, and Hilton P. Silva. "What have the revelations about Neanderthal DNA revealed about Homo sapiens?" Anthropological Review 83, no. 1 (March 1, 2020): 93–107. http://dx.doi.org/10.2478/anre-2020-0008.

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Abstract:
AbstractGenetic studies have presented increasing indications about the complexity of the interactions between Homo sapiens, Neanderthals and Denisovans, during Pleistocene. The results indicate potential replacement or admixture of the groups of hominins that lived in the same region at different times. Recently, the time of separation among these hominins in relation to the Last Common Ancestor – LCA has been reasonably well established. Events of mixing with emphasis on the Neanderthal gene flow into H. sapiens outside Africa, Denisovans into H. sapiens ancestors in Oceania and continental Asia, Neanderthals into Denisovans, as well as the origin of some phenotypic features in specific populations such as the color of the skin, eyes, hair and predisposition to develop certain kinds of diseases have also been found. The current information supports the existence of both replacement and interbreeding events, and indicates the need to revise the two main explanatory models, the Multiregional and the Out-of-Africa hypotheses, about the origin and evolution of H. sapiens and its co-relatives. There is definitely no longer the possibility of justifying only one model over the other. This paper aims to provide a brief review and update on the debate around this issue, considering the advances brought about by the recent genetic as well as morphological traits analyses.
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38

Wilson, Clare. "DNA reveals our recent Denisovan ancestry." New Scientist 242, no. 3224 (April 2019): 9. http://dx.doi.org/10.1016/s0262-4079(19)30580-9.

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39

Ferguson, Kennan. "What Was Politics to the Denisovan?" Political Theory 42, no. 2 (November 5, 2013): 167–87. http://dx.doi.org/10.1177/0090591713506714.

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40

Neubauer, Fernanda. "UMA BREVE VISÃO GERAL DOS ÚLTIMOS 10 ANOS DAS PRINCIPAIS DESCOBERTAS DO PLEISTOCENO SUPERIOR NO VELHO MUNDO: HOMO FLORESIENSIS, NEANDERTAL, DENISOVAN." Cadernos do LEPAARQ (UFPEL) 16, no. 32 (December 15, 2019): 204. http://dx.doi.org/10.15210/lepaarq.v16i32.14039.

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Nos últimos dez anos, novos dados fósseis, arqueológicos e genéticos alteraram significativamente nossa compreensão sobre o povoamento do Velho Mundo no Pleistoceno Superior. Os pesquisadores há muito têm sido desafiados a definir o lugar da humanidade na evolução e a rastrear nossa filogenia. Diferenças na morfologia esquelética de fósseis de hominídeos muitas vezes levaram à nomeação de novas espécies distintas, mas descobertas genéticas recentes desafiaram a perspectiva tradicional, demonstrando que o DNA humano moderno contém genes herdados dos Neandertais e Denisovans, questionando assim seu status como uma espécie separada. A recente descoberta do Homo floresiensis da Ilha de Flores também levantou questões interessantes sobre a quantidade de diversidade genética e morfológica que estava presente durante o Pleistoceno Superior. Este artigo discute a natureza e as implicações da evidência em relação ao Homo floresiensis, Neandertais e Denisovans, e analisa brevemente as principais descobertas do Pleistoceno Superior nos últimos dez anos de pesquisa no Velho Mundo e sua importância para o estudo da evolução humana.Abstract: In the last ten years, new fossil, archaeological, and genetic data have significantly altered our understanding of the peopling of the Old World in the Late Pleistocene. Scholars have long been challenged to define humanity’s place in evolution and to trace our phylogeny. Differences in the skeletal morphology of hominin fossils have often led to the naming of distinct new species, but recent genetic findings have challenged the traditional perspective by demonstrating that modern human DNA contains genes inherited from Neandertals and Denisovans, thus questioning their status as separate species. The recent discovery of Homo floresiensis from Flores Island has also raised interesting queries about how much genetic and morphological diversity was present during the Late Pleistocene. This article discusses the nature and implications of the evidence with respect to Homo floresiensis. Neandertals and Denisovans, and briefly reviews major Late Pleistocene discoveries from the last ten years of research in the Old World and their significance to the study of human evolution.
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Gibbons, Ann. "Ancient jaw gives elusive Denisovans a face." Science 364, no. 6439 (May 2, 2019): 418–19. http://dx.doi.org/10.1126/science.364.6439.418.

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42

Gibbons, A. "Elusive Denisovans Sighted in Oldest Human DNA." Science 342, no. 6163 (December 5, 2013): 1156. http://dx.doi.org/10.1126/science.342.6163.1156.

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Povysil, Gundula, and Sepp Hochreiter. "IBD Sharing between Africans, Neandertals, and Denisovans." Genome Biology and Evolution 8, no. 12 (December 2016): 3406–16. http://dx.doi.org/10.1093/gbe/evw234.

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44

Marshall, Michael. "Did humans meet mysterious Denisovans in Indonesia?" New Scientist 251, no. 3350 (September 2021): 15. http://dx.doi.org/10.1016/s0262-4079(21)01552-9.

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Gibbons, A. "A Denisovan Legacy in the Immune System?" Science 333, no. 6046 (August 25, 2011): 1086. http://dx.doi.org/10.1126/science.333.6046.1086.

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46

Clyde, Dorothy. "The girl with Neanderthal and Denisovan parents." Nature Reviews Genetics 19, no. 11 (September 12, 2018): 668–69. http://dx.doi.org/10.1038/s41576-018-0054-6.

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47

Slon, Viviane, Charlotte Hopfe, Clemens L. Weiß, Fabrizio Mafessoni, Marco de la Rasilla, Carles Lalueza-Fox, Antonio Rosas, et al. "Neandertal and Denisovan DNA from Pleistocene sediments." Science 356, no. 6338 (April 27, 2017): 605–8. http://dx.doi.org/10.1126/science.aam9695.

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48

Marchi, Emanuele, Alex Kanapin, Matthew Byott, Gkikas Magiorkinis, and Robert Belshaw. "Neanderthal and Denisovan retroviruses in modern humans." Current Biology 23, no. 22 (November 2013): R994—R995. http://dx.doi.org/10.1016/j.cub.2013.10.028.

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Gokhman, David, Nadav Mishol, Marc de Manuel, David de Juan, Jonathan Shuqrun, Eran Meshorer, Tomas Marques-Bonet, Yoel Rak, and Liran Carmel. "Reconstructing Denisovan Anatomy Using DNA Methylation Maps." Cell 180, no. 3 (February 2020): 601. http://dx.doi.org/10.1016/j.cell.2020.01.020.

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Jacobs, Guy S., Georgi Hudjashov, Lauri Saag, Pradiptajati Kusuma, Chelzie C. Darusallam, Daniel J. Lawson, Mayukh Mondal, et al. "Multiple Deeply Divergent Denisovan Ancestries in Papuans." Cell 177, no. 4 (May 2019): 1010–21. http://dx.doi.org/10.1016/j.cell.2019.02.035.

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