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Cross-linked DNA extracted from the 4,000-year-old liver of the ancient Egyptian priest Nekht-Ankh.

Ancient DNA (aDNA) is DNA isolated from ancient specimens.[1][2] Due to degradation processes (including cross-linking, deamination and fragmentation) ancient DNA is of lower quality in comparison with modern genetic material.[3] Even under the best preservation conditions, there is an upper boundary of 0.4-1.5 million years for a sample at around to contain sufficient DNA for contemporary sequencing technologies.[4] Genetic material has been recovered from archaeological and historical skeletal material, mummified tissues, archival collections of non-frozen medical specimens, preserved plant remains, ice and permafrost cores as well as marine and lake sediments.

Contents

History of ancient DNA studiesEdit

1980sEdit

The first study of what would come to be called aDNA was conducted in 1984, when Russ Higuchi and colleagues at the University of California, Berkeley reported that traces of DNA from a museum specimen of the Quagga not only remained in the specimen over 150 years after the death of the individual, but could be extracted and sequenced.[5] Over the next two years, through investigations into natural and artificially mummified specimens, Svante Pääbo confirmed that this phenomenon was not limited to relatively recent museum specimens but could apparently be replicated in a range of mummified human samples that dated as far back as several thousand years.[6]

The laborious processes that were required at that time to sequence such DNA (through bacterial cloning) were an effective brake on the development of the field of ancient DNA (aDNA). However, with the development of the Polymerase Chain Reaction (PCR) in the late 1980s, the field began to progress rapidly.[7][8] Double primer PCR amplification of aDNA (jumping-PCR) can produce highly skewed and non-authentic sequence artifacts. Multiple primer, nested PCR strategy was used to overcome those shortcomings.

1990sEdit

The post-PCR era heralded a wave of publications as numerous research groups tried their hands at aDNA. Soon a series of incredible findings had been published, claiming authentic DNA could be extracted from specimens that were millions of years old, into the realms of what Lindahl (1993b) has labelled Antediluvian DNA.[9] The majority of such claims were based on the retrieval of DNA from organisms preserved in amber. Insects such as stingless bees,[10] termites,[11] and wood gnats,[12] as well as plant[13] and bacterial[14] sequences were extracted from Dominican amber dating to the Oligocene epoch. Still older sources of Lebanese amber-encased weevils, dating to within the Cretaceous epoch, reportedly also yielded authentic DNA.[15] DNA retrieval was not limited to amber.

Several sediment-preserved plant remains dating to the Miocene were successfully investigated.[16] Then, in 1994 and to international acclaim, Woodward et al. reported the most exciting results to date[17] — mitochondrial cytochrome b sequences that had apparently been extracted from dinosaur bones dating to more than 80 million years ago. When in 1995 two further studies reported dinosaur DNA sequences extracted from a Cretaceous egg,[18] it seemed that the field would revolutionize knowledge of the Earth's evolutionary past. Even these extraordinary ages were topped by the claimed retrieval of 250-million-year-old halobacterial sequences from halite.[19][20]

Whole genome sequencing started to yield results in 1995.

2000sEdit

Single primer extension (abr. SPEX) amplification was introduced in 2007 to address postmortem DNA modification damage.[21]

Ancient DNA revolutionEdit

Since 2009 the field of aDNA-studies has been revolutionzed, with the introduction of much cheaper research-techniques, leading to new insights in human migrations.[22]

Problems and errorsEdit

Degradation processesEdit

Due to degradation processes (including cross-linking, deamination and fragmentation) ancient DNA is of lower quality in comparison with modern genetic material.[3] The damage characteristics and ability of aDNA to survive through time restricts possible analyses and places an upper limit on the age of successful samples Allentoft et al. (2012). There is a theoretical relationship between time and DNA degradation,[23] although differences in environmental conditions complicates things. Samples subjected to different conditions are unlikely to predictably align to a uniform age-degradation relationship.[24] The environmental effects may even matter after excavation, as DNA decay rates may increase,[25] particularly under fluctuating storage conditions.[26] Even under the best preservation conditions, there is an upper boundary of 0.4-1.5 million years for a sample at around to contain sufficient DNA for contemporary sequencing technologies.[4]

Research into the decay of mitochondrial and nuclear DNA in Moa bones has modelled mitochondrial DNA degradation to an average length of 1 base pair after 6,830,000 years at −5 °C.[27] The decay kinetics have been measured by accelerated aging experiments further displaying the strong influence of storage temperature and humidity on DNA decay.[28] Nuclear DNA degrades at least twice as fast as mtDNA. As such, early studies that reported recovery of much older DNA, for example from Cretaceous dinosaur remains, may have stemmed from contamination of the sample.

Age limitEdit

A critical review of ancient DNA literature through the development of the field highlights that few studies after about 2002 have succeeded in amplifying DNA from remains older than several hundred thousand years.[29] A greater appreciation for the risks of environmental contamination and studies on the chemical stability of DNA have resulted in concerns being raised over previously reported results. The dinosaur DNA was later revealed to be human Y-chromosome,[30] while the DNA reported from encapsulated halobacteria has been criticized based on its similarity to modern bacteria, which hints at contamination.[31] A 2007 study also suggests that these bacterial DNA samples may not have survived from ancient times, but may instead be the product of long-term, low-level metabolic activity.[32]

aDNA may contain a large number of postmortem mutations, increasing with time. Some regions of polynucleotite are more susceptible to this degradation, so sequence data can bypass statistical filters used to check the validity of data.[33] Due to sequencing errors, great caution should be applied to interpretation of population size.[34] Substitutions resulting from deamination cytosine residues are vastly over-represented in the ancient DNA sequences. Miscoding of C to T and G to A accounts for the majority of errors.[35]

ContaminationEdit

Another problem with ancient DNA samples is contamination by modern human DNA and by microbial DNA (most of which is also ancient).[36][37] New methods have emerged in recent years to prevent possible contamination of aDNA samples, including conducting extractions under extreme sterile conditions, using special adapters to identify endogenous molecules of the sample (over ones that may have been introduced during analysis), and applying bioinformatics to resulting sequences based on known reads in order approximate rates of contamination.[38]

Non-human aDNAEdit

Despite the problems associated with 'antediluvian' DNA, a wide and ever-increasing range of aDNA sequences have now been published from a range of animal and plant taxa. Tissues examined include artificially or naturally mummified animal remains,[5][39] bone (c.f. Hagelberg et al. 1989; Cooper et al. 1992; Hagelberg et al. 1994),[40] paleofaeces,[41][42] alcohol preserved specimens (Junqueira et al. 2002), rodent middens,[43] dried plant remains (Goloubinoff et al. 1993; Dumolin-Lapegue et al. 1999) and recently, extractions of animal and plant DNA directly from soil samples.[44]

In June 2013, a group of researchers announced that they had sequenced the DNA of a 560–780 thousand year old horse, using material extracted from a leg bone found buried in permafrost in Canada's Yukon territory.[45]

In 2013, a German team reconstructed the mitochondrial genome of an Ursus deningeri more than 300,000 years old, proving that authentic ancient DNA can be preserved for hundreds of thousand years outside of permafrost.[46]

Human aDNAEdit

Due to the considerable anthropological, archaeological, and public interest directed toward human remains, they have received considerable attention from the DNA community.

SourcesEdit

Due to the morphological preservation in mummies, many studies from the 1990s and 2000s used mummified tissue as a source of ancient human DNA. Examples include both naturally preserved specimens, for example, those preserved in ice, such as the Ötzi the Iceman (Handt et al. 1994), or through rapid desiccation, such as high-altitude mummies from the Andes (c.f. Pääbo 1986; Montiel et al. 2001), as well as various sources of artificially preserved tissue (such as the chemically treated mummies of ancient Egypt).[47] However, mummified remains are a limited resource. The majority of human aDNA studies have focused on extracting DNA from two sources that are much more common in the archaeological recordbone and teeth. Several other sources have also yielded DNA, including paleofaeces (Poinar et al. 2001) and hair (Baker et al. 2001, Gilbert et al. 2004). Contamination remains a major problem when working on ancient human material.

Ancient pathogen DNA has been successfully retrieved from samples dating to more than 5,000 years old in humans and as long as 17,000 years ago in other species. In addition to the usual sources of mummified tissue, bones and teeth, such studies have also examined a range of other tissue samples, including calcified pleura (Donoghue et al. 1998), tissue embedded in paraffin,[48][49] and formalin-fixed tissue.[50] Efficient computational tools have been developed for pathogen and microorganism aDNA analyses in a small (QIIME) and large scale (FALCON [51]).

ResultsEdit

Taking preventative measures in their procedure against such contamination though, a 2012 study analyzed bone samples of a Neanderthal group in the El Sidrón cave, finding new insights on potential kinship and genetic diversity from the aDNA.[52] In November 2015, scientists reported finding a 110,000-year-old tooth containing DNA from the Denisovan hominin, an extinct species of human in the genus Homo.[53][54]

The research has added new complexity to the peopling of Eurasia. It has also revealed new information about links between the ancestors of Central Asians and the indigenous peoples of the Americas. In Africa, older DNA degrades quickly due to the warmer tropical climate, although, in September 2017, ancient DNA samples, as old as 8,100 years old, have been reported.[55]

Researchers specializing in ancient DNAEdit

See alsoEdit

Further readingEdit

ReferencesEdit

  1. ^ Pevsner 2015.
  2. ^ Jones 2016.
  3. ^ a b Allentoft, M.E. et al., 2012. The half-life of DNA in bone: measuring decay kinetics in 158 dated fossils. Proceedings of the Royal Society of London B: Biological Sciences, 279, pp.4724–4733.
  4. ^ a b Willerslev, E. et al., 2004. Long-term persistence of bacterial DNA. Current biology: CB, 14(1), pp.9–10.
  5. ^ a b Higuchi R; Bowman B; Freiberger M; Ryder OA; Wilson AC (1984). "DNA sequences from the quagga, an extinct member of the horse family". Nature. 312 (5991): 282–4. Bibcode:1984Natur.312..282H. doi:10.1038/312282a0. PMID 6504142. 
  6. ^ Pääbo 1985a; Pääbo 1985b; Pääbo 1986
  7. ^ Mullis KB; Faloona FA (1987). "Specific synthesis of DNA in vitro via a polymerase-catalyzed chain reaction". Meth. Enzymol. Methods in Enzymology. 155: 335–50. doi:10.1016/0076-6879(87)55023-6. ISBN 978-0-12-182056-5. PMID 3431465. 
  8. ^ Saiki RK, Gelfand DH, Stoffel S, et al. (January 1988). "Primer-directed enzymatic amplification of DNA with a thermostable DNA polymerase". Science. 239 (4839): 487–91. Bibcode:1988Sci...239..487S. doi:10.1126/science.2448875. PMID 2448875. 
  9. ^ Lindahl 1993b
  10. ^ Cano et al. 1992a; Cano et al. 1992b
  11. ^ De Salle et al. 1992; De Salle et al. 1993
  12. ^ De Salle and Grimaldi 1994
  13. ^ Poinar et al. 1993
  14. ^ Cano et al. 1994
  15. ^ Cano et al. 1993
  16. ^ Golenberg et al. 1990; Golenberg 1991
  17. ^ Woodward SR; Weyand NJ; Bunnell M (November 1994). "DNA sequence from Cretaceous period bone fragments". Science. 266 (5188): 1229–32. Bibcode:1994Sci...266.1229W. doi:10.1126/science.7973705. PMID 7973705. 
  18. ^ An et al. 1995; Li et al. 1995
  19. ^ Vreeland RH; Rosenzweig WD; Powers DW (October 2000). "Isolation of a 250 million-year-old halotolerant bacterium from a primary salt crystal". Nature. 407 (6806): 897–900. Bibcode:2000Natur.407..897V. doi:10.1038/35038060. PMID 11057666. 
  20. ^ Fish SA; Shepherd TJ; McGenity TJ; Grant WD (May 2002). "Recovery of 16S ribosomal RNA gene fragments from ancient halite". Nature. 417 (6887): 432–6. Bibcode:2002Natur.417..432F. doi:10.1038/417432a. PMID 12024211. 
  21. ^ Brotherton, P; Endicott, P; Sanchez, Jj; Beaumont, M; Barnett, R; Austin, J; Cooper, A (2007). "Novel high-resolution characterization of ancient DNA reveals C > U-type base modification events as the sole cause of post-mortem miscoding lesions" (Free full text). Nucleic Acids Research. 35 (17): 5717–28. doi:10.1093/nar/gkm588. ISSN 0305-1048. PMC 2034480 . PMID 17715147. 
  22. ^ Reich 2018.
  23. ^ Hebsgaard, M.B., Phillips, M.J. & Willerslev, E., 2005. Geologically ancient DNA: fact or artefact? Trends in microbiology, 13(5), pp.212–220.
  24. ^ Hansen, A.J. et al., 2006. Crosslinks rather than strand breaks determine access to ancient DNA sequences from frozen sediments. Genetics, 173(2), pp.1175–1179.
  25. ^ Pruvost, M. et al., 2007. Freshly excavated fossil bones are best for amplification of ancient DNA. Proceedings of the National Academy of Sciences of the United States of America, 104(3), pp.739–744.
  26. ^ Burger, J. et al., 1999. DNA preservation: a microsatellite-DNA study on ancient skeletal remains. Electrophoresis, 20(8), pp.1722–1728.
  27. ^ Allentoft ME; Collins M; Harker D; Haile J; Oskam CL; Hale ML; Campos PF; Samaniego JA; Gilbert MTP; Willerslev E; Zhang G; Scofield RP; Holdaway RN; Bunce M (2012). "The half-life of DNA in bone: measuring decay kinetics in 158 dated fossils". Proceedings of the Royal Society B. 279 (1748): 4724–33. doi:10.1098/rspb.2012.1745. PMC 3497090 . PMID 23055061. 
  28. ^ Grass, R. N.; Heckel, R.; Puddu, M.; Paunescu, D.; Stark, W. J. (2015). "Robust Chemical Preservation of Digital Information on DNA in Silica with Error-Correcting Codes". Angewandte Chemie International Edition. 54 (8): 2552–2555. doi:10.1002/anie.201411378. PMID 25650567. 
  29. ^ Willerslev E, Hansen AJ, Binladen J, et al. (May 2003). "Diverse plant and animal genetic records from Holocene and Pleistocene sediments". Science. 300 (5620): 791–5. Bibcode:2003Sci...300..791W. doi:10.1126/science.1084114. PMID 12702808. 
  30. ^ Zischler H; Höss M; Handt O; von Haeseler A; van der Kuyl AC; Goudsmit J (May 1995). "Detecting dinosaur DNA". Science. 268 (5214): 1192–3; author reply 1194. doi:10.1126/science.7605504. PMID 7605504. 
  31. ^ Nicholls H (February 2005). "Ancient DNA Comes of Age". PLOS Biology. 3 (2): e56. doi:10.1371/journal.pbio.0030056. PMC 548952 . PMID 15719062. 
  32. ^ Johnson SS; Hebsgaard MB; Christensen TR; Mastepanov M; Nielsen R; Munch K; Brand T; Gilbert MT; Zuber MT; Bunce M; Rønn R; Gilichinsky D; Froese D; Willerslev E (September 2007). "Ancient bacteria show evidence of DNA repair". PNAS. 104 (36): 14401–5. Bibcode:2007PNAS..10414401J. doi:10.1073/pnas.0706787104. PMC 1958816 . PMID 17728401. 
  33. ^ Pääbo, Svante; Poinar, Hendrik; Serre, David; Jaenicke-Després, Viviane; Hebler, Juliane; Rohland, Nadin; Kuch, Melanie; Krause, Johannes; Vigilant, Linda; Hofreiter, Michael (2004). "Genetic Analyses from Ancient DNA" (PDF). Annual Review of Genetics. 38 (1): 645–679. doi:10.1146/annurev.genet.37.110801.143214. ISSN 0066-4197. PMID 15568989. Archived from the original (PDF) on December 17, 2008. Retrieved December 1, 2008. 
  34. ^ Johnson, Pl; Slatkin, M (Jan 2008). "Accounting for bias from sequencing error in population genetic estimates" (Free full text). Molecular Biology and Evolution. 25 (1): 199–206. doi:10.1093/molbev/msm239. ISSN 0737-4038. PMID 17981928. 
  35. ^ Briggs, Aw; Stenzel, U; Johnson, Pl; Green, Re; Kelso, J; Prüfer, K; Meyer, M; Krause, J; Ronan, Mt; et al. (Sep 2007). "Patterns of damage in genomic DNA sequences from a Neandertal" (Free full text). Proceedings of the National Academy of Sciences of the United States of America. 104 (37): 14616–21. Bibcode:2007PNAS..10414616B. doi:10.1073/pnas.0704665104. ISSN 0027-8424. PMC 1976210 . PMID 17715061. 
  36. ^ Gansauge, Marie-Theres; Meyer, Matthias (2014). "Selective enrichment of damaged DNA molecules for ancient genome sequencing". Genome Research. 24 (9): 1543–9. doi:10.1101/gr.174201.114. PMC 4158764 . PMID 25081630. 
  37. ^ Pratas, Diogo; Hosseini, Morteza; Grilo, Goncalo; Pinho, Armando; Silva, Raquel; Caetano, Tania; Carneiro, Joao; Pereira, Filipe (2018). "Metagenomic Composition Analysis of an Ancient Sequenced Polar Bear Jawbone from Svalbard". Genes. 9 (9): 445. doi:10.3390/genes9090445. 
  38. ^ Slatkin, Montgomery; Racimo, Fernando (2016-06-07). "Ancient DNA and human history". Proceedings of the National Academy of Sciences. 113 (23): 6380–6387. doi:10.1073/pnas.1524306113. ISSN 0027-8424. PMC 4988579 . PMID 27274045. 
  39. ^ Thomas RH; Schaffner W; Wilson AC; Pääbo S (August 1989). "DNA phylogeny of the extinct marsupial wolf". Nature. 340 (6233): 465–7. Bibcode:1989Natur.340..465T. doi:10.1038/340465a0. PMID 2755507. 
  40. ^ Hänni C; Laudet V; Stehelin D; Taberlet P (December 1994). "Tracking the origins of the cave bear (Ursus spelaeus) by mitochondrial DNA sequencing". Proceedings of the National Academy of Sciences of the United States of America. 91 (25): 12336–40. Bibcode:1994PNAS...9112336H. doi:10.1073/pnas.91.25.12336. PMC 45432 . PMID 7991628. 
  41. ^ Poinar HN, Hofreiter M, Spaulding WG, et al. (July 1998). "Molecular coproscopy: dung and diet of the extinct ground sloth Nothrotheriops shastensis". Science. 281 (5375): 402–6. Bibcode:1998Sci...281..402P. doi:10.1126/science.281.5375.402. PMID 9665881. 
  42. ^ Hofreiter M, Poinar HN, Spaulding WG, et al. (December 2000). "A molecular analysis of ground sloth diet through the last glaciation". Mol. Ecol. 9 (12): 1975–84. doi:10.1046/j.1365-294X.2000.01106.x. PMID 11123610. 
  43. ^ Kuch M; Rohland N; Betancourt JL; Latorre C; Steppan S; Poinar HN (May 2002). "Molecular analysis of an 11,700-year-old rodent midden from the Atacama Desert, Chile". Mol. Ecol. 11 (5): 913–24. doi:10.1046/j.1365-294X.2002.01492.x. PMID 11975707. 
  44. ^ Willerslev E; Cooper A (January 2005). "Ancient DNA". Proceedings of the Royal Society B. 272 (1558): 3–16. doi:10.1098/rspb.2004.2813. PMC 1634942 . PMID 15875564. 
  45. ^ Erika Check Hayden (26 June 2013). "First horses arose 4 million years ago". Nature. doi:10.1038/nature.2013.13261. 
  46. ^ Dabney; et al. (2013). "Complete mitochondrial genome sequence of a Middle Pleistocene cave bear reconstructed from ultrashort DNA fragments". PNAS. 110 (39): 15758–15763. Bibcode:2013PNAS..11015758D. doi:10.1073/pnas.1314445110. PMC 3785785 . Retrieved 18 January 2014. 
  47. ^ Hänni C; Laudet V; Coll J; Stehelin D (July 1994). "An unusual mitochondrial DNA sequence variant from an Egyptian mummy". Genomics. 22 (2): 487–9. doi:10.1006/geno.1994.1417. PMID 7806242. 
  48. ^ Jackson PJ, Hugh-Jones ME, Adair DM, et al. (February 1998). "PCR analysis of tissue samples from the 1979 Sverdlovsk anthrax victims: The presence of multiple Bacillus anthracis strains in different victims". Proceedings of the National Academy of Sciences of the United States of America. 95 (3): 1224–9. Bibcode:1998PNAS...95.1224J. doi:10.1073/pnas.95.3.1224. PMC 18726 . PMID 9448313. 
  49. ^ Basler CF, Reid AH, Dybing JK, et al. (February 2001). "Sequence of the 1918 pandemic influenza virus nonstructural gene (NS) segment and characterization of recombinant viruses bearing the 1918 NS genes". Proceedings of the National Academy of Sciences of the United States of America. 98 (5): 2746–51. Bibcode:2001PNAS...98.2746B. doi:10.1073/pnas.031575198. PMC 30210 . PMID 11226311. 
  50. ^ Taubenberger JK; Reid AH; Krafft AE; Bijwaard KE; Fanning TG (March 1997). "Initial genetic characterization of the 1918 "Spanish" influenza virus". Science. 275 (5307): 1793–6. doi:10.1126/science.275.5307.1793. PMID 9065404. 
  51. ^ Pratas D; Pinho AJ; Silva RM; Rodrigues JMOS; Hosseini M; Caetano T; Ferreira PJSG (February 2018). "FALCON: a method to infer metagenomic composition of ancient DNA". bioRxiv. doi:10.1101/267179. 
  52. ^ Lalueza-Fox, Carles; Rosas, Antonio; Rasilla, Marco de la (2012-01-20). "Palaeogenetic research at the El Sidrón Neanderthal site". Annals of Anatomy - Anatomischer Anzeiger. Special Issue: Ancient DNA. 194 (1): 133–137. doi:10.1016/j.aanat.2011.01.014. 
  53. ^ Zimmer, Carl (16 November 2015). "In a Tooth, DNA From Some Very Old Cousins, the Denisovans". The New York Times. Retrieved 16 November 2015. 
  54. ^ Sawyer, Susanna; Renaud, Gabriel; Viola, Bence; Hublin, Jean-Jacques; Gansauge, Marie-Theres; Shunkov, Michael V.; Derevianko, Anatoly P.; Prüfer, Kay; Kelso, Janet; Pääbo, Svante (11 November 2015). "Nuclear and mitochondrial DNA sequences from two Denisovan individuals". PNAS. 112: 201519905. Bibcode:2015PNAS..11215696S. doi:10.1073/pnas.1519905112. Retrieved 16 November 2015. 
  55. ^ Zimmer, Carl (21 September 2017). "Clues to Africa's Mysterious Past Found in Ancient Skeletons". The New York Times. Retrieved 21 September 2017. 
  56. ^ Diamond, Jared (April 20, 2018). A Brand-New Version of Our Origin Story. The New York Times. Retrieved April 23, 2018. 

BibliographyEdit

ConferencesEdit

  • The 1st International Ancient DNA Conference was held at St John's College, Nottingham, England, from July 8 to July 10, 1991.
  • The 2nd International Ancient DNA Conference was held at the Smithsonian Institution, Washington D.C., USA, from October 7 to October 9, 1993.
  • The 3rd International Ancient DNA Conference was held at Oxford University, Oxford, England, from July 21 to July 22, 1995.
  • The 4th International Ancient DNA Conference was held at the Georg-August University, Göttingen, Germany, from June 5 to June 7, 1997.
  • [1] – The 5th International Ancient DNA Conference was held at the University of Manchester, Manchester, England, from July 12 to July 14, 2000.
  • The 6th International Conference on Ancient DNA and Associated Biomolecules was held at the Hebrew University, Jerusalem, Tel-Aviv and Rehovot, Israel, from July 21 to July 25, 2002.
  • The 7th International Conference on Ancient DNA & Associated Biomolecules was held at the University of Queensland, Brisbane, Australia, from July 10 to July 17, 2004.
  • [2] – The 8th International Conference on Ancient DNA and Associated Biomolecules was held at the Medical University of Łódź, Łódź, Poland, from October 16 to October 19, 2006.
  • The 9th International Conference on Ancient DNA and Associated Biomolecules was held at the University of Naples, Naples & Pompeii, Italy, from October 19 to October 22, 2008.
  • [3] – The 10th International Conference on Ancient DNA and Associated Biomolecules was held at the Ludwig-Maximilians-University, Munich, Germany, from October 10 to October 13, 2010.
  • [4] A day with ancient DNA held at the University of Florence, Florence, Italy, 22 March 2012.
  • The 1st International Symposium of Biomolecular Archaeology was held at the Vrij Universiteit, Amsterdam, Netherlands.
  • [5] The 2nd International Symposium of Biomolecular Archaeology was held at the University of Stockholm, Sweden, from 7–9 September 2006
  • [6] The 3rd International Symposium of Biomolecular Archaeology was held at the University of York, UK, from 14–16 September 2008
  • [7] The 4th International Symposium of Biomolecular Archaeology was held at the University of Copenhagen, Denmark, from 8–10 September 2010
  • [8] The 5th International Symposium of Biomolecular Archaeology was held at Beijing, China from 15–1 August 2012

See alsoEdit

Read moreEdit

External linksEdit