Ancient DNA(aDNA) isDNAisolated from ancient sources (typicallyspecimens,but alsoenvironmental DNA).[1][2]Due to degradation processes (includingcross-linking,deaminationandfragmentation)[3]ancient DNA is more degraded in comparison with contemporary genetic material.[4]Genetic material has been recovered from paleo/archaeological and historical skeletal material,mummifiedtissues, archival collections of non-frozen medical specimens, preserved plant remains, ice and frompermafrostcores, marine and lake sediments andexcavationdirt.

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

Even under the best preservation conditions, there is an upper boundary of 0.4–1.5 million years for a sample to contain sufficient DNA for sequencing technologies.[5]The oldest DNA sequenced from physical specimens are frommammothmolars in Siberia over 1 million years old.[6]In 2022, two-million year old genetic material was recovered from sediments inGreenland,and is currently considered the oldest DNA discovered so far.[7][8]

History of ancient DNA studies

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1980s

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Quagga (Equus quagga quagga), an extinct sub-species of zebra.

The first study of what would come to be called aDNA was conducted in 1984, when Russ Higuchi and colleagues at theUniversity of California, Berkeleyreported that traces of DNA from a museum specimen of theQuagganot only remained in the specimen over 150 years after the death of the individual, but could be extracted and sequenced.[9]Over the next two years, through investigations into natural and artificially mummified specimens,Svante Pääboconfirmed that this phenomenon was not limited to relatively recent museum specimens but could apparently be replicated in a range ofmummifiedhuman samples that dated as far back as several thousand years.[10][11][12]

The laborious processes that were required at that time to sequence such DNA (throughbacterial cloning) were an effective brake on the study of ancient DNA (aDNA) and the field ofmuseomics.However, with the development of thePolymerase Chain Reaction(PCR) in the late 1980s, the field began to progress rapidly.[13][14][15]Double primer PCR amplification of aDNA (jumping-PCR) can produce highly skewed and non-authentic sequence artifacts. Multiple primer,nested PCRstrategy was used to overcome those shortcomings.

1990s

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A diptera (Mycetophilidae) from the Eocene (40-50 million years ago) in a piece of transparent Baltic amber along with other smaller inclusions. Shown under daylight (big photograph) and under UV light (small photograph).

The post-PCR era heralded a wave of publications as numerous research groups claimed success in isolating 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 labelledAntediluvianDNA.[16]The majority of such claims were based on the retrieval of DNA from organisms preserved inamber.Insects such as stingless bees,[17][18]termites,[19]and wood gnats,[20]as well as plant[21]and bacterial[22]sequences were said to have been extracted fromDominicanamber dating to theOligoceneepoch. Still older sources of Lebanese amber-encasedweevils,dating to within theCretaceousepoch, reportedly also yielded authentic DNA.[23]Claims of DNA retrieval were not limited to amber.

Reports of several sediment-preserved plant remains dating to theMiocenewere published.[24][25]Then in 1994, Woodwardet al.reported what at the time was called the most exciting results to date[26]— 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,[27][28]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 fromhalite.[29][30]

The development of a better understanding of the kinetics of DNA preservation, the risks of sample contamination and other complicating factors led the field to view these results more skeptically. Numerous careful attempts failed to replicate many of the findings, and all of the decade's claims of multi-million year old aDNA would come to be dismissed as inauthentic.[31]

2000s

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Single primer extension amplification was introduced in 2007 to address postmortem DNA modification damage.[32]Since 2009 the field of aDNA studies has been revolutionized with the introduction of much cheaper research techniques.[33]The use of high-throughputNext Generation Sequencing(NGS) techniques in the field of ancient DNA research has been essential for reconstructing the genomes of ancient or extinct organisms. A single-stranded DNA (ssDNA) library preparation method has sparked great interest among ancient DNA (aDNA) researchers.[34][35]

Svante Pääbo (left) with his medal for the Nobel Prize on Physiology or Medicine.

In addition to these technical innovations, the start of the decade saw the field begin to develop better standards and criteria for evaluating DNA results, as well as a better understanding of the potential pitfalls.[31][36]

2020s

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Autumn of 2022, the Nobel Prize of Physiology or Medicine was awarded to Svante Pääbo "for his discoveries concerning the genomes of extinct hominins and human evolution".[37]A few days later, on the 7th of December 2022, a study inNaturereported that two-million year old genetic material was found in Greenland, and is currently considered the oldest DNA discovered so far.[7][8]

Problems and errors

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Degradation processes

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Due to degradation processes (including cross-linking, deamination and fragmentation),[3]ancient DNA is of lower quality than modern genetic material.[4]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.[4]There is a theoretical correlation between time and DNA degradation,[38]although differences in environmental conditions complicate matters. Samples subjected to different conditions are unlikely to predictably align to a uniform age-degradation relationship.[39]The environmental effects may even matter after excavation, as DNA decay-rates may increase,[40]particularly under fluctuating storage conditions.[41]Even under the best preservation conditions, there is an upper boundary of 0.4 to 1.5 million years for a sample to contain sufficient DNA for contemporary sequencing technologies.[5]

Research into the decay ofmitochondrialandnuclear DNAinmoabones has modelled mitochondrial DNA degradation to an average length of 1base pairafter 6,830,000 years at −5 °C.[4]The decay kinetics have been measured by accelerated aging experiments, further displaying the strong influence of storage temperature and humidity on DNA decay.[42]Nuclear DNA degrades at least twice as fast as mtDNA. Early studies that reported recovery of much older DNA, for example fromCretaceousdinosaurremains, may have stemmed from contamination of the sample.

Age limit

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A critical review of ancient DNA literature through the development of the field highlights that few studies have succeeded in amplifying DNA from remains older than several hundred thousand years.[43]A greater appreciation for the risks of environmental contamination and studies on thechemical stabilityof DNA have raised concerns over previously reported results. The alleged dinosaur DNA was later revealed to be humanY-chromosome.[44]The DNA reported from encapsulatedhalobacteriahas been criticized based on its similarity to modern bacteria, which hints at contamination,[36]or they may be the product of long-term, low-levelmetabolicactivity.[45]

aDNA may contain a large number of postmortemmutations,increasing with time. Some regions of polynucleotide are more susceptible to this degradation, allowing erroneous sequence data to bypass statistical filters used to check the validity of data.[31]Due to sequencing errors, great caution should be applied to interpretation of population size.[46]Substitutions resulting fromdeaminationofcytosineresidues are vastly over-represented in the ancient DNA sequences. Miscoding ofCtoTandGtoAaccounts for the majority of errors.[47]

Contamination

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Another problem with ancient DNA samples is contamination by modern human DNA and by microbial DNA (most of which is also ancient).[48][49]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 (distinguished from those introduced during analysis), and applying bioinformatics to resulting sequences based on known reads in order to approximate rates of contamination.[50][51]

Authentication of aDNA

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Development in the aDNA field in the 2000s increased the importance of authenticating recovered DNA to confirm that it is indeed ancient and not the result of recent contamination. As DNA degrades over time, the nucleotides that make up the DNA may change, especially at the ends of the DNA molecules. The deamination of cytosine to uracil at the ends of DNA molecules has become a way of authentication. During DNA sequencing, the DNA polymerases will incorporate an adenine (A) across from the uracil (U), leading to cytosine (C) to thymine (T) substitutions in the aDNA data.[52]These substitutions increase in frequency as the sample gets older. Frequency measurement of the C-T level, ancient DNA damage, can be made using various software such as mapDamage2.0 or PMDtools[53][54]and interactively on metaDMG.[55]Due to hydrolytic depurination, DNA fragments into smaller pieces, leading to single-stranded breaks. Combined with the damage pattern, this short fragment length can also help differentiate between modern and ancient DNA.[56][57]

Non-human aDNA

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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 planttaxa.Tissues examined include artificially or naturally mummified animal remains,[9][58]bone,[59][60][61][62]shells,[63]paleofaeces,[64][65]alcohol preserved specimens,[66]rodent middens,[67]dried plant remains,[68][69]and recently, extractions of animal and plant DNA directly fromsoilsamples.[70]

In June 2013, a group of researchers includingEske Willerslev,Marcus Thomas Pius Gilbertand Orlando Ludovic of theCentre for Geogenetics,Natural History Museum of Denmarkat theUniversity of Copenhagen,announced that they had sequenced the DNA of a 560–780 thousand year old horse, using material extracted from a leg bone found buried inpermafrostin Canada'sYukonterritory.[71][72][73]A German team also reported in 2013 the reconstructedmitochondrial genomeof a bear,Ursus deningeri,more than 300,000 years old, proving that authentic ancient DNA can be preserved for hundreds of thousand years outside of permafrost.[74]The DNA sequence of even older nuclear DNA was reported in 2021 from the permafrost-preserved teeth of two Siberianmammoths,both over a million years old.[6][75]

Researchers in 2016 measured chloroplast DNA in marine sediment cores, and found diatom DNA dating back to 1.4 million years.[76]This DNA had a half-life significantly longer than previous research, of up to 15,000 years. Kirkpatrick's team also found that DNA only decayed along a half-life rate until about 100 thousand years, at which point it followed a slower, power-law decay rate.[76]

Human aDNA

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Map of human fossils with an age of at least ~40,000 years that yielded genome-wide data[77]

Due to the considerableanthropological,archaeological,andpublic interestdirected toward human remains, they have received considerable attention from the DNA community. There are also more profound contamination issues, since the specimens belong to the same species as the researchers collecting and evaluating the samples.

Sources

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Due to themorphologicalpreservation 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, such as theÖtzi the Icemanfrozen in a glacier[78]and bodies preserved through rapiddesiccationat high altitude in the Andes,[12][79]as well as various chemically treated preserved tissue such as the mummies of ancient Egypt.[80]However, mummified remains are a limited resource. The majority of human aDNA studies have focused on extracting DNA from two sources much more common in thearchaeological record:bonesandteeth.The bone that is most often used for DNA extraction is thepetrousear bone, since its dense structure provides good conditions for DNA preservation.[81]Several other sources have also yielded DNA, includingpaleofaeces,[82]andhair.[83][84]Contamination remains a major problem when working on ancient human material.

AncientpathogenDNA 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 calcifiedpleura,[85]tissue embedded inparaffin,[86][87]andformalin-fixed tissue.[88]Efficient computational tools have been developed for pathogen and microorganism aDNA analyses in a small (QIIME[89]) and large scale (FALCON[90]).

Results

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Taking preventative measures in their procedure against such contamination though, a 2012 study analyzed bone samples of aNeanderthalgroup in the El Sidrón cave, finding new insights on potential kinship and genetic diversity from the aDNA.[91]In November 2015, scientists reported finding a 110,000-year-old tooth containing DNA from theDenisovan hominin,anextinctspeciesofhumanin the genusHomo.[92][93]

The research has added new complexity to the peopling of Eurasia. A study from 2018[94]showed that aBronze Agemass migration had greatly impacted the genetic makeup of the British Isles, bringing with it theBell Beakerculture from mainland Europe.

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.[95]

Moreover, ancient DNA has helped researchers to estimate modern human divergence.[96]By sequencing African genomes from three Stone Age hunter gatherers (2000 years old) and four Iron Age farmers (300 to 500 years old), Schlebusch and colleagues were able to push back the date of the earliest divergence between human populations to 350,000 to 260,000 years ago.

As of 2021, the oldest completely reconstructed human genomes are~45,000 years old.[97][77]Such genetic data provides insights into the migration and genetic history – e.g.of Europe– including aboutinterbreeding between archaic and modern humanslike a common admixture between initial European modern humans and Neanderthals.[98][77][99]

Researchers specializing in ancient DNA

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See also

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References

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