Archaeological Wool Textiles A Window Into Ancient Sheep Genetics

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Archaeological Wool Textiles: A Window into Ancient Sheep

Genetics?
Chapter · November 2019

DOI: 10.1017/9781108656405.012
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TWELVE
ARCHAEOLOGICAL WOOL TEXTILES:

A WINDOW INTO ANCIENT SHEEP
GENETICS?
Luise Ørsted Brandt and Morten Allentoft
INTRODUCTION
Research in archaeological textiles has traditionally focused on identifying

the raw materials they were made of and understanding the technologies
with which they were produced. However, over the past decade, the number
of studies that discuss social and cultural aspects of archaeological textiles
has increased (Andersson Strand et al. 2010). These include studies on the
organisation of ancient textile production, how textiles reflect ancient trends
and fashion (e.g. Mathiassen et al. 2014), how they relate to the identity of
the individuals who wore them (e.g. Harlow 2012), as well as their use in,
for instance, religious contexts (e.g. Brøns 2017; Brøns and Nosch 2017).
Identifying where the raw material for textile production, the fibres, came
from is key to fully understanding the production line and use of textiles in
ancient communities.
Animal fibres from archaeological textiles have been identified to different
mammalian species. Amongst these are fibres from beaver and rabbit (Rast-
Eicher 2014), horse (Wincott Heckett 2012), goat (Green 1980; Sinding et al.
2015) and sheep (e.g. Barber 1991, 20–30). The type of fibres and their quality
determine the properties of the textile: how it insulates against heat and cold,
its flexibility and ability to absorb water, and how it feels against the skin of the
owner –smooth, soft or itchy.The type and quality of fibres also affect the entire
textile chaîne opératoire from the procurement of the fibres, the techniques used
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I ntroduction 275
to prepare them and the technological possibilities for spinning and weaving
them, to the final visual appearance of the textile (e.g. Andersson Strand 2012).
The refinement of fibres and development of the wool fibre had an enor-
mous impact on ancient societies (e.g. Breniquet and Michel 2014). People
refined fibres to obtain functional or socially desirable properties which
affected the entire chain of processes in the textile production and also cultural
and social aspects of human life.The refined fibres defined the techniques used
for thread production, which textiles they were used for, and the consumers’
perceptions, needs and demands for textiles. Therefore, characterisation of the
raw material in detail has been in sharp focus within archaeological textile
research.
Animal fibres are commonly identified to species by microscopy through
evaluation of morphological traits such as hair diameter, length of the fibre, shape
and distance of the cuticle scales, appearance and dimension of the medulla
and cortex and cross-sectional shape, which differ between species (Hausman
1920; Wildman 1954). These traits are evaluated and identified by comparison
with atlases and reference collections (see e.g. Hausman 1920; Luniak 1953;
Teerink 1991). Despite being widely applied, the reliability of species identi-
fication based on light and electron microscopic observation of skin and hair
morphology has been subject to intense debate for several reasons: (1) the
reproducibility of the method requires extended knowledge and experience;
(2) hair morphology can differ within the same species, between different
parts of the animal’s surface, or according to age, sex, seasonality, nutrition and
health status; (3) these challenges are even more pronounced in archaeological
hairs, which are often poorly preserved and thus complicate the identifications;
and (4) because fibre atlases are based only on modern species, breeds and
populations, and at present no fibre atlas that includes archaeological material
is available (see Brandt et al. 2014). Overall, species identification based on
the microscopic analysis of ancient hairs is not straightforward, which makes
it desirable to develop alternative, ideally more reliable, approaches for the
species identification of archaeological animal fibres.
Within the past two decades, the development of ancient DNA (aDNA)
technologies has resulted in several successful studies of archaeological hair
(see below), suggesting that fibres from archaeological textiles could perhaps
be studied with such aDNA-based approaches. Prior to 2010, to our know-
ledge, wool had not been used for genetic studies, but this changed when
Brandt et al. (2011) demonstrated the potential of analysing DNA from ancient
textiles.
Sheep were the most important source of wool in prehistory (e.g. Barber
1991, 20–30), and apart from species identification, research has often focused
on identification of prehistoric sheep breeds (and wool types) and identifi-
cation of specific traits such as sheep wool quality and colour and how these
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276 Archaeological W ool T extiles and S heep G enetics
changed over time. Identifying different ancient sheep breeds, their develop-
ment and refinement has long been attempted by zooarchaeological analysis
(e.g. Hatting 1991) and wool fibre measurements (e.g. Gleba 2012; Rast-Eicher
and Bender Jørgensen 2013; Ryder 1969, 1983b). For instance, changes in the
wool fibre diameter in textiles from the Apennine peninsula were interpreted
by Gleba (2012) as a development of the sheep fleece from the Bronze Age
to the Iron Age, and distinct fleece types in her study suggest that several
breeds coexisted in the first millennium bc (Gleba 2012). Also, Rast-Eicher
and Bender Jørgensen (2013) speculate from fibre diameter measurements that
different wool types or breeds coexisted in Bronze Age Europe. While the
quality of sheep wool can be estimated by measurements of fibre diameters,
assigning the wool type to a specific prehistoric breed is difficult and unre-
solved. Interestingly, genetic studies of modern sheep (Kijas et al. 2009) and
historical parchment of sheep skin (Teasdale et al. 2015) have demonstrated
genetic differentiation between sheep breeds, which opens the possibility for
also identifying and comparing different lineages or ancestral influences in
prehistoric populations, and also for comparing these with our modern breeds.
Genetic studies have also identified genes in modern sheep involved in wool
quality, such as, e.g., fibre diameter and staple length (Forrest et al. 2009; Gong
et al. 2010, 2011; Itenge-Mweza et al. 2007; Zeng et al. 2011), which could be
used also to study wool fibre traits and development in ancient sheep.
Elucidating species identification, types of breeds, and the genetic link to
specific wool qualities would not only be of interest to the ancient textile
research community, but would also feed into a general archaeological discus-
sion of strategies in prehistoric animal husbandry and the utilisation of pri-
mary and secondary products from different animal species.
Since the first study on archaeological wool textiles (Brandt et al. 2011),
aDNA methodologies have developed rapidly, constantly increasing the possi-
bilities for new research questions and for analysing DNA from many different
archaeological materials (see introduction to DNA in Appendix 12.1). The
potentials and pitfalls of aDNA research on archaeological textiles have not
been appraised in detail despite a great demand for this knowledge, in par-
ticular when considering the new possibilities offered by Next Generation
Sequencing (NGS). In this chapter we review previous research on DNA from
archaeological textiles of sheep wool, present the current state of knowledge,
and discuss the perspectives for future DNA research on archaeological textiles.
PRESERVATION OF ARCHAEOLOGICAL TEXTILES AND THE DNA POTENTIAL
Textiles are rare in archaeological contexts. As with other organic materials, they
often decompose rapidly after disposal. Nevertheless, under extremely advanta-
geous circumstances textiles can be preserved for millennia, thus offering great
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T HE T E X T I L E C HAÎ NE O P É RAT O I RE AN D D N A D E P O S I TI O N 277
potential not only for archaeological research, but also for natural scientific
investigations. Favourable preservation conditions include: environments saturated
with water (waterlogged), as in the case of the north European bogs and Danish
oak coffins (Hald 1980); frozen conditions, as for the Norse burials in Greenland
(Østergård 2004); and the saline or very dry environments such as in the salt
mines in Hallstatt, Austria (Bichler et al. 2005) and the deserts of the Xinjiang
province of China (e.g. Barber 1999). Moreover, textiles can be preserved by
exposure to fire, as observed in the well-known carbonised Iron Age textiles from
Gordion in Turkey (Bellinger 1962), or by contact with metals that can sometimes
mineralise the textiles or prevent the growth of damaging microorganisms. For
example, this is seen in the Danish weapon deposits from Illerup Ådal and Vimose
(Möller-Wiering 2010) and also in inhumation graves, observed in textiles where
dress fasteners and buckles were once attached (Walton-Rogers 2007).
DNA sequences from well-preserved wool textiles could potentially offer
a wealth of information about the animals that provided the raw material.
Biologically, wool is a type of hair. Many studies have documented excellent
DNA preservation in both modern and ancient hair samples (Gilbert et al. 2004,
2007; Rasmussen et al. 2010, 2011; Wilson and Gilbert 2007). These studies have
also demonstrated that hair has many advantages in comparison to bone and
other tissues commonly studied in aDNA contexts. It is not as vulnerable to
modern DNA contamination as bone (Gilbert et al. 2007), and it has proved easy
to decontaminate with a solution of hypochlorite (Gilbert et al. 2004, 463). DNA
extracts from hair seem to contain a high proportion of mitochondrial DNA
compared with extracts from other tissues, which is important if mitochondrial
DNA (mtDNA) is the target, and the survival of mtDNA in hair is better than
in bone (Gilbert et al. 2007). Moreover, DNA in hair has been shown to survive
storage well, even when kept at room temperatures (Gilbert et al. 2007).
Although these previous results of DNA preservation in hair are encour-
aging, they do not imply that DNA is also present in wool. First of all, DNA
survival in hair is known to vary widely with hair type.This is probably a result
of inter-and intra-organismal differences in the hair cell keratinisation process,
which is the formation and hardening of the hair in the follicle where the cells
that form the hair shaft undergo cell death (Bengtsson et al. 2012). For some
individuals, DNA survival even varies between different hair types (Gilbert
et al. 2007;Wilson and Gilbert 2007, 154–155). Second, wool is highly processed
hair, which may have implications for DNA preservation (see below).
THE EFFECTS OF THE TEXTILE CHAÎNE OPÉRATOIRE AND

DEPOSITION ON DNA
The textile chaîne opératoire, the transformation of wool fibres into a textile,
involves several processes, some of which may degrade DNA molecules in
278
the hair. In order to avoid sampling of ancient precious textiles before having
solid knowledge on DNA preservation in wool pre-and post-deposition,
Brandt et al. (2011) set up a pilot-study investigating the DNA content in
modern wool.
Most likely, wool from the entire body of the sheep was used for textile pro-
duction, and both coarser fibres (hairs and kemps) and finer fibres (underwool)
have been found in prehistoric Danish textiles from the Bronze Age and Iron
Age (e.g. Ryder 1983a; Walton 1988). Different natural pigmentations of wool
were also used (e.g. Ryder 1983a). As previously stated, such variety of hair
types could differ in terms of their content of DNA. An initial qPCR assay
(quantitative PCR, see Appendix 12.2 for introduction to PCR) of a range of
samples, including wool from different breeds, body parts, hair types (kemp
and underwool) and pigmentation, demonstrated that amplifiable mtDNA and
nuclear DNA was present in a wide range of modern sheep wool samples, and
that none of the tested variables affected the ability to obtain DNA in modern
samples (Brandt 2010; Brandt et al. 2011).
Preparation of textile fibres by combing, spinning and weaving would prob-
ably not introduce significant damage to the DNA molecules in the fibres.
However, wool treatment processes such as dyeing and mordanting, which
have been documented in archaeological textiles (e.g. Vanden Berghe et al.
2009;Walton 1988), involve heating and chemical treatment of the wool, which
could certainly affect DNA preservation. Brandt et al. (2011) therefore used a
qPCR assay to test how DNA in a range of wool samples survived treatment
with different dyes and mordants. None of the tested dyes (madder, tansy, weld
and woad) seemed to have a significant negative effect on DNA preservation.
This was not, however, the case for the mordants: treatment with 18% alun
caused a 10 to 10,000 times reduction in DNA yield in unburied samples,
while no DNA at all could be amplified from alun mordanted and experimen-
tally buried samples. Also treatment with 5% iron and oak tannin caused an up
to 1,000 times reduction in DNA yield, but could still be PCR-amplified after
burial, perhaps suggesting these treatments are slightly less detrimental than
alun (Brandt 2010; Brandt et al. 2011). It would be highly informative to test
the negative effects of mordants with NGS technology which offers an ability
to analyse millions of highly degraded DNA fragments. This would provide a
much better reflection of the preserved DNA fragment lengths and various
DNA damage parameters than offered by the simple ‘success or failure’ obser-
vation when attempting to amplify DNA with PCR (see Appendix 12.2).
Many of the organic dyes that were used in prehistoric times, for instance
madder and weld, required a mordant to be fixed to wool (Cardon 2007;
Walton Rogers 2007, 37–38). It is therefore highly likely that many dyed pre-
historic textiles have been pre-treated with a mordant. For example, this may
explain the unsuccessful attempts to extract DNA from a textile from the
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T HE T E X T I L E C HAÎ NE O P É RAT O I RE AN D D N A D E P O S I TI O N 279
Iron Age Hammerum grave, which was dyed with madder (Brandt et al. 2011;
Rostholm 2009). Based on the results by Brandt et al. (2011), it is preferable to
avoid analysing mordanted ancient textiles with PCR-based technology, as the
DNA in the wool may have been completely degraded to very short fragments
even before the burial of the textile.
Mordants are hard to detect and thus also hard to avoid during the sampling
process. While tanning agents can be recorded by HPLC (High Performance
Liquid Chromatography –a technique used to separate, identify and quantify
components in a sample), inorganic mordants such as alun and iron can only be
identified through analyses of chemical elements. Ringgaard (2010) used XRF
(X-ray fluorescence) and SEM-EDX (Scanning Electron Microscopy coupled
with Energy Dispersive X-ray Spectroscopy) to detect this. Ringgaard (2010)
states that exchange of ions with adjacent soil during burial can lead to false
positive results, which makes the mordant detection problematic and incon-
clusive (Ringgaard 2010, 256–257). At present, it seems preferable to restrict
genetic analyses to textiles that have either not been dyed at all or, alternatively,
have been dyed with types of dye that do not require a mordant, for example
woad and indigotin (Brandt 2010).
After dyeing, textiles may have been turned into garments, used and re-
used, and perhaps dyed again until finally discarded. The archaeological envir-
onments in which Danish textiles (e.g. Bender Jørgensen 1986; Broholm
and Hald 1940) have survived include different soil types, pH values, and
differences in water and oxygen saturation. For her PhD thesis, conservator
Maj Ringgaard conducted experiments in which she buried samples of wool
textiles for different lengths of time (8, 16 and 24 months) (Ringgaard 2010).
The samples that were buried for 8 months were split into batches with and
without access to oxygen, whereas all the remaining samples were buried
in anoxic environments. These samples were then used to test the effects of
short-term degradation of DNA in wool (Brandt 2010; Brandt et al. 2011).
This experiment demonstrated that mtDNA survives short-term degradation
in an artificial burial environment, and even after 24 months the content of
mtDNA in the samples was unaffected. Nuclear DNA could also be amp-
lified from the samples exposed to short-tem degradation up to 24 months.
Importantly, however, access to oxygen showed a significant decrease in the
amount of amplifiable DNA (Brandt 2010). These tests demonstrated that the
history of every textile, from possible chemical treatment during manufac-
turing to the environment in which it was buried, will affect the potential
for molecular research and should be considered before sampling. It is clear
that mordants and access to oxygen in the burial environment are both highly
negative factors when assessing the potential for aDNA preservation in textiles
(Brandt et al. 2011). The overall conclusion, based on the results from modern
sheep wool samples, is that DNA is certainly present in modern wool, and it
280
should therefore also potentially be possible to recover DNA molecules from

ancient samples, given the lack of harsh processing and the right preservation
environment.
SAMPLING ARCHAEOLOGICAL TEXTILES FOR GENETIC RESEARCH
When sampling archaeological material for aDNA research it is important

to try to minimise DNA contamination from modern sources (e.g. Allentoft
2013). Ancient textiles and archaeological material in general have often been
handled many times over the decades, and sampling will often have to be
carried out in storage facilities and exhibitions. This will of course limit the
molecular sterility of the sample and the sampling environment, but despite
these circumstances, it is still important to use gloves and sterile tools for sam-
pling, and to thoroughly clean these between each sampling, to avoid cross-
contamination between samples. With NGS approaches (see Appendix 12.2),
contamination with human DNA may not be detrimental, as it can often
be identified bioinformatically and thus excluded from the analyses. Limiting
contamination from external sources (whether human or microbial DNA) is
still extremely important because it will provide more genetic data from the
species of interest in the downstream analyses (Fig. 12.1).
When selecting the exact spot for sampling a textile, the fibres should ideally
come from one thread from one spin direction, which can then be carefully
pulled out. We do not know how common it was to mix wool from different
regions and perhaps breeds, but this was demonstrated to be the case for wool
from the Danish pre-Roman Iron Age large tubular textile from Huldremose
II (Frei et al. 2009). Limiting the sampling to one thread limits the possibility of
discovering whether fibres represent several individuals and breeds potentially
from different geographical origins. If several samples can be spared, it could
be worth sampling in multiple spots to test whether the wool is from different
sheep. Brandt et al. (2011) used 10–70 mg of modern wool and 10–200 mg of
ancient wool, which for most samples equalled a thread of 2–3 cm in length.
DNA EXPERIMENTS ON ARCHAEOLOGICAL TEXTILES: AN OVERVIEW
To date, the number of archaeological sheep wool textiles that have been
analysed for aDNA is limited. Published examples are limited to the textiles
analysed by Brandt (2010, 2015) and Brandt et al. (2011), sheep wool analysed
by Nikulina (2016), Norse textiles analysed by Sinding et al. (2017) and hairs
from the garment worn by the Neolithic Iceman, Ötzi (Olivieri et al. 2012;
O’Sullivan et al. 2016). Table 12.1 provides an overview.
After initial tests on modern samples (see above), Brandt et al. (2011)
attempted to extract and amplify (see PCR in Appendix 12.2) both mtDNA
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DN A E X P E R I M E N T S O N ARC H AE O L O G I C AL TE XTI L E S 281
12.1 Luise Ørsted Brandt analyses DNA from archaeological textiles in a clean laboratory
dedicated to ancient DNA in order to avoid contamination from modern sources of DNA.
(Photo: Morten Allentoft.)
and nuclear DNA from a set of archaeological and historical textiles,

spanning different contexts and dating from the Bronze Age to histor-
ical times (see Table 12.1). They obtained a 121 bp long region from the
mtDNA 16s gene, which enabled them to identify the wool as deriving
from domestic sheep (Ovis aries), although their nuclear DNA assay failed
to yield DNA on all samples. Using an improved extraction method (see
Appendix 12.3), Brandt (2015) revisited two textile samples, one from the
Bronze Age Borum Eshøj burial mound and the other from the Iron Age
Hammerum inhumation grave, which were originally analysed by Brandt
et al. (2011), as well as additional Iron Age textiles. The DNA extracts were
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282
Table 12.1 Summary of aDNA research on ancient wool textiles and sheep hair.
Site Specimen and Date Dye Extraction Target DNA Result Reference
context method
Borum Textile from 1350 bc Not analysed Gilbert et al. mtDNA 16S no DNA Brandt et al.
Eshøj, oak coffin 2004 121bp, 2011
Denmark nuDNA 66bp
Voldtofte, Urn wrapping 900–700 bc Not analysed Gilbert et al. mtDNA 16S no DNA Brandt et al.
Denmark in burial 2004 121bp, nuDNA 2011
mound 66bp
Krogens Textile from 400–200 bc Not analysed Gilbert et al. mtDNA 16S mtDNA –species ID Brandt et al.
Mølle peat bog 2004 121bp, nuDNA Ovis aries 2011
Mose, 66bp
Denmark
Hammerum, Textile from 0–130 ad Not detected but Gilbert et al. mtDNA 16S no DNA Brandt et al.
Denmark inhumation some reddish 2004 121bp, nuDNA 2011
grave parts 66bp
Vrangstrup, Textile from 250–320 ad No dye Gilbert et al. mtDNA 16S no DNA Brandt et al.
Denmark inhumation 2004 121bp, nuDNA 2011
grave 66bp
Mammen, Textile from 970–971 ad No dye, but Gilbert et al. mtDNA 16S mtDNA –species ID Brandt et al.
Denmark chamber reddish of 2004 121bp, nuDNA Ovis aries 2011
burial colour 66bp
GUS, Textile from 13th No dye, natural Gilbert et al. mtDNA 16S mtDNA –species ID Brandt et al.
Greenland culture century pigmentation 2004 121bp, nuDNA Ovis aries 2011
layer 66bp
GUS, Textile from 13th No dye, natural Gilbert et al. mtDNA 16S mtDNA –species ID Brandt et al.
Greenland culture century pigmentation 2004 121bp, nuDNA Ovis aries 2011
layer 66bp
283
Borreby, Gobelin 17th White/undyed Gilbert et al. mtDNA 16S mtDNA –species ID Brandt et al.
Denmark tapestry century 2004 121bp, nuDNA Ovis aries 2011
66bp
Borreby, Gobelin 17th Beige Gilbert et al. mtDNA 16S mtDNA –species ID Brandt et al.
66bp
Borreby, Gobelin 17th Green Gilbert et al. mtDNA 16S mtDNA –species ID Brandt et al.
66bp
Borreby, Gobelin 17th Red Gilbert et al. mtDNA 16S mtDNA –species ID Brandt et al.
66bp
Borreby, Gobelin 17th Blue Gilbert et al. mtDNA 16S mtDNA –species ID Brandt et al.
66bp
Borreby, Gobelin 17th Dark red Gilbert et al. mtDNA 16S mtDNA –species ID Brandt et al.
66bp
Borreby, Gobelin 17th Brown Gilbert et al. mtDNA 16S no DNA Brandt et al.
Denmark tapestry century 2004 121bp, nuDNA 2011
66bp
Borum Eshøj, Textile from 1350 bc Not analysed Orlando et al. mtDNA 97 bp no DNA Brandt 2015
Denmark oak coffin 2011
Hammerum, Textile from 0–130 ad Not detected but Orlando et al. mtDNA 97 bp no DNA Brandt 2015
Denmark inhumation some reddish 2011
grave parts
Hammerum, Textile from 0–130 ad Not detected but Orlando et al. mtDNA 97 bp no DNA Brandt 2015
Denmark inhumation some reddish 2011
grave parts
(continued)
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284
284
Table 12.1 (Cont.)
Site Specimen and Date Dye Extraction Target DNA Result Reference
context method
Lønne Hede, Textile from 40–70 ad No information Orlando et al. mtDNA 97 bp no DNA Brandt 2015
Denmark inhumation 2011
grave
(1969)
Lønne Hede, Textile from 1–70 ad Red Orlando et al. mtDNA 97 bp no DNA Brandt 2015
grave 2
Lønne Hede, Textile from 1–70 ad Red/brown Orlando et al. mtDNA 97 bp no DNA Brandt 2015
grave 1
Feddersen Tuft of wool Roman No information Unknown mtDNA, not Not reproducible Nikulina 2016
Wierde, from period specified
Germany culture
layer
Germany culture
layer
Germany culture
layer
Germany culture
layer
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Ötzal Alps, Sheep hair 3359–3105 Not analysed Gilbert et al. mtDNA, several mtDNA, 2429 bp Olivieri et al.
Italy from Ötzi’s bc 2007 regions 2012
clothing,
glacier
Ötzal Alps, Sheep hair 3359–3105 Not analysed Gamba et al. Full mitogenome 3,6X mitogenome O´Sullivan
Italy from Ötzi’s bc 2014 et al. 2016
clothing,
glacier
Ötzal Alps, Sheep hair 3359–3105 Not analysed Gamba et al. Full mitogenome 15,5X mitogenome O´Sullivan
Italy from Ötzi’s bc 2014 et al. 2016
clothing,
glacier
GUS, Textile from 13th Black, natural Orlando et al. Full 20X mitogenome Sinding
Greenland culture century pigmentation 2011 mitogenome et al. 2017
layer
285
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tested with a PCR assay targeting a 97 bp fragment of mtDNA, but these

tests yielded no amplifiable mtDNA.
DNA AND TEXTILES FROM BOGS AND BURIALS
The waterlogged environments of the Danish oak coffins and bogs have
preserved one of the world’s finest collections of archaeological textiles (e.g.
Broholm and Hald 1940; Hald 1980). Despite their great physical preservation,
both of these environments have been demonstrated to be harsh on DNA
(Hughes et al. 1986; Brandt et al. 2011).The observation that oak coffin material
is difficult for aDNA analysis was also demonstrated by negative results based
on human hair analysed with NGS technology from oak coffins from Egtved
(Frei et al. 2015) and Borum Eshøj.1 The negative results are most likely an
effect of the acidic environment inside Bronze Age burial mounds (Breuning-
Madsen et al. 2001, 692; 2003, 343), causing the DNA to fragment at a high rate.
An early and failed attempt to analyse DNA from the Lindow Man found
in a British bog (Hughes et al. 1986) lowered the expectations for preser-
vation of DNA in acidic bog environments. This finding was supported by
negative DNA results using NGS on the Danish bog body (bone and skin) of
the famous Tollund Man.2 However, Brandt et al. (2011) successfully extracted
mtDNA from a pre-Roman Iron Age textile found in the Danish bog Krogens
Mølle Mose. This particular bog may be an exceptional case, as it is situated
on limestone, which creates a more alkaline environment (Karin M. Frei,
National Museum, Copenhagen, Denmark, pers. comm.), and combined with
the anoxic environments of bogs, this has probably provided highly favour-
able preservation conditions. In general, textiles found in bogs are not ideal
candidates for molecular research. Moreover, the sphagnum in bogs naturally
tans organic material (van der Sanden 1996, 18), which could also have a nega-
tive impact, similar to the tannin tested in Brandt et al. (2011). This is unfor-
tunate, given the relatively larger number of well-preserved textiles from the
Danish bogs.
Samples from less acidic soils have offered better possibilities for aDNA
retrieval, such as those from Gården under Sandet (‘Farm beneath the Sand’),
Greenland, and Mammen, Denmark (Brandt et al. 2011). The textiles from
Mammen were recovered from a Viking Age chamber in the Bjerringhøj
mound near Viborg in Denmark (Brandt et al. 2011). These textiles are
amongst the best-preserved archaeological textiles from Danish inhumation
graves (Østergård 1991) (Fig. 12.2). Most probably, protection by the wooden
chamber has favoured morphological and molecular preservation, which is
why the sample yielded a 121 bp sequence of mtDNA. The textiles from the
Norse site Gården under Sandet in Greenland were found in a permafrozen
layer and analysed by Brandt et al. (2011) and Sinding et al. (2017). The two
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D N A A N D TE XTI L E S F R O M B O G S AN D B U RI AL S 287
12.2 Embroidery from the garment belonging to a male found in Bjerringhøj, Mammen,
close to Viborg in Denmark. The burial is dated to ad 970–971. Through the analysis of mito-
chondrial DNA, the wool in the textile was species-identified as deriving from domestic sheep
(Ovis aries).
(Photo: Lennart Larsen.)
samples analysed by Brandt et al. (2011) yielded 121 bp sequences of mtDNA,

which allowed for a species identification to Ovis aries (sheep). Sinding et al.
(2017) generated a full mitochondrial genome from a textile sample from the
same site using NGS technology. However, textile samples from five other
Norse sites failed to provide enough DNA sequences from any species to be
taken into account (Sinding et al. 2017). It is unknown whether any nuclear
DNA is present in these samples, but this would be interesting to test in future
research. In general, samples from permafrozen layers seem to represent the
most promising preservation context, as also demonstrated by positive DNA
288
results from sheep hair samples from the clothing of the Iceman, Ötzi (see
later; Olivieri et al. 2012; O’Sullivan et al. 2016). Temperature is clearly one of
the most crucial factors for aDNA preservation (see Appendix 12.4), which
also explains why positive results were obtained from hair of snow hare and
goat from the ‘Farm beneath the Sand’ (Sinding et al. 2015), and why a com-
plete ancient human genome could be bioinformatically reconstructed based
on DNA in hair from permafrozen layers near Qeqertasussuk, Greenland
(Rasmussen et al. 2010). This is particularly promising for studying the textiles
of the Norse, which could ideally provide us with information on whether
these were made of local sheep wool or imported. Other factors like museum
storage, time and conditions are also involved in determining the limit of long-
term DNA preservation (e.g. Pruvost et al. 2007), but the effects of these are
not as well understood.
Nikulina (2016) analysed four tufts of sheep wool from the Iron Age site
of Feddersen Wierde in northern Germany. The protocol is unpublished and
no information is available on the exact DNA target region, but the experi-
ment demonstrated high fragmentation of the DNA in the samples, low
reproducibility and signs of contamination, which leaves the results incon-
clusive. Nikulina’s results confirm that successful aDNA amplification is more
unlikely on samples from temperate environments than on those from cold
contexts.
Studies of the mtDNA Control Region and/or the Cytochrome b gene of
modern domesticated sheep breeds from a wide geographical range have iden-
tified five haplogroups (i.e. overall genetic variants, see Appendix 12.1): A, B,
C, D and E (Guo et al. 2005; Hiendleder et al. 1998, 2002; Meadows et al. 2005,
2007, 2011; Pedrosa et al. 2005; Wood and Phua 1996). The haplogroups have
been tentatively proposed to each represent a separate domestication event
(Meadows et al. 2011; Pedrosa et al. 2005). Haplogroups A and B are the most
frequent, and have been identified in modern sheep from all sampled geo-
graphical regions. Haplotype A occurs particularly frequently in Asian sheep,
whereas haplogroup B dominates in European sheep (Hiendleder et al. 1998;
Wood and Phua 1996). Haplogroup C is less prevalent but has been located
within both Asia, the Fertile Crescent and Europe (Guo et al. 2005; Pedrosa
et al. 2005; Pereira et al. 2006; Tapio et al. 2006). Haplogroup D and E are the
least frequent and have only been identified in samples from Turkey and the
Caucasus (Meadows et al. 2007;Tapio et al. 2006). Sheep haplogroups have also
been identified in a number of studies on ancient teeth (Cai et al. 2007) and
bone samples (Brandt 2015; Cai et al. 2011; Horsburgh and Rhines 2010; Niemi
et al. 2013; Rannamäe et al. 2016). Both Olivieri et al. (2012) and O’Sullivan
et al. (2016) analysed sheep hair from the garment of the Copper Age Iceman,
Ötzi (around 5,350–5,100 years bp), who was found extremely well-preserved
in a glacier on the border between Austria and Italy in 1991. Olivieri et al.
289
D N A and Textiles from B ogs and B urials 289
(2012) analysed an unspecified part of the garment of ‘black hair shafts’, which
they species-identified to sheep and from which they gained a total of 2,429 bp
of mtDNA using several sets of PCR primers (Olivieri et al. 2012). Also here,
the permafrozen layer in which the Iceman was preserved has provided highly
favourable conditions for DNA preservation. The analysed DNA sequences
enabled the identification of the wool-providing sheep as belonging to the
mitochondrial haplogroup B (for explanation, see Appendix 12.1 and below).
O’Sullivan et al. (2016) applied NGS technology to sequence the DNA in
hairs from both the light coat and dark coat sheep of Ötzi’s garment (Niall
J. O’Sullivan, Institute for Mummy Studies, EURAC Research, Bolzano, Italy,
pers. comm.) and managed to assemble complete mitochondrial genomes
from both (O’Sullivan et al. 2016). Both individuals were again placed within
the sheep mitochondrial haplogroup B. Likewise, the DNA in the sheep wool
textile sample from the ‘Farm beneath the Sand’ from Sinding et al. (2017)
could also be assigned to haplogroup B.
Today, haplogroup B is the dominating haplogroup within modern
European sheep. In previous analyses it has been identified in Finnish Iron
Age, medieval, and post-medieval sheep bones, whereas haplogroup A was
observed in the medieval and post-medieval samples (Niemi et al. 2013). Both
haplogroups A and B have been observed in Estonian sheep bones from the
Bronze Age and Middle Ages (Rannamäe et al. 2016), and in sheep from the
Százhalombatta-Földvár Bronze Age settlement in Hungary (Sabatini et al.
2019), whereas haplogroup B was found in two Danish samples from the
Neolithic period and the Roman Iron Age (Brandt 2015). Thus, the oldest
European sheep samples analysed to date belong to haplogroup B, and it will
be interesting to see if future research can confirm haplogroup B as the oldest
European sheep lineage. The four sheep haplotypes obtained by O’Sullivan
et al. (2016) cluster closely together in the phylogenetic tree –potentially
representing sheep from the same population. If indeed the garments were
produced in the same place, this could suggest a genetic structure defined by
geography in Copper Age sheep populations, but more data from many more
sites is needed to confirm this.
The mtDNA haplogroups can provide insights into the genetic diversity
of past sheep populations and possibly uncover geographically defined gen-
etic structures and also movement of sheep between geographical regions.
Currently, the number of ancient sheep mtDNA sequences is limited and
much of the comparative data is from modern individuals. DNA studies of
modern populations are unable to account for lost genetic variation (e.g. now
extinct haplogroups) and cannot be used to directly document genetic changes
through time. Therefore, analyses and comparisons with more ancient sheep
mtDNA sequences in the future could potentially help clarify prehistoric
movements of sheep and the emergence of new sheep breeds at certain points
290
in time and space. If this could in time be extended into nuclear DNA analyses,
it could provide even higher resolution in addressing these questions.
MITOCHONDRIAL AND NUCLEAR DNA: PROS AND CONS
In addition to species identification, mtDNA data can sometimes be used

to assign individuals to specific populations, when the inter- population
haplogroup frequencies are highly differentiated. For example, this is the case
for the European Mesolithic hunter-gatherers and the Neolithic peoples that
replaced them (e.g. Brandt et al. 2013). This, however, requires that mtDNA
haplogroup frequencies have already been mapped out in the ancient
populations, which is currently not the case for sheep. Though mtDNA has
proved valuable for identifying species and studying basic genetic diversity,
nuclear genetic markers can provide much deeper insights into population
histories, and provide higher resolution to identify and quantify genetic popu-
lation structure in sheep (Kijas et al. 2012; Tapio et al. 2010). Moreover, nuclear
genome-scale analyses can be used to characterise genes that were artificially
selected for during the domestication process, for example to increase wool
production. This would allow us to discuss animal husbandry, development of
specialised breeds and resource exploitation in prehistoric times.
Despite successes in extracting and amplifying nuclear DNA from arch-
aeological human hair (Rasmussen et al. 2010, 2011) and modern sheep
wool (Brandt et al. 2011), nuclear DNA sequences from archaeological or
historical sheep wool textiles have not been published. There are likely sev-
eral explanations for this. First, nuclear DNA degrades faster than mtDNA,
possibly because mtDNA is protected behind a double membrane and by
having a circular structure, in contrast to nuclear DNA (Allentoft et al. 2012).
Bengtsson et al. (2012) confirmed that this difference in preservation is also
the case for hair. Thus, NGS-based technology must be attempted in this con-
text since it offers much better possibilities for sequencing the short nuclear
DNA compared with PCR-based approaches (see Appendix 12.2). Second,
each cell contains only two nuclear copies (in diploid organisms), but hundreds
to thousands of mitochondria. Because of rapid hair growth, this numerical
difference may be even more elevated in hair where metabolically active cells
have many mitochondria to meet the high energy demands of the cells. Third,
although several studies have shown that hair can yield well-preserved DNA,
the actual amount of DNA in hair is relatively low compared with that in
other tissues (Bengtsson et al. 2012; Gilbert et al. 2004, 2007).
Although hair samples have provided excellent results of nuclear DNA from
prehistoric and historic individuals, these examples come mainly from very cold
environments, such as from the exceptional environment of the permafrozen
hair tuft from Qeqertasussuk, Greenland (Rasmussen et al. 2010), mammoth
hair samples from permafrozen deposits in Siberia (Gilbert et al. 2007) or a
291
C onclusion 291
lock of hair sampled (and never buried) from an Aboriginal Australian in 1923
(Ramussen et al. 2011), which is not old enough to have experienced severe
biomolecule degradation.
Based on the above observations, one cannot rule out the possibility of
obtaining nuclear DNA in ancient untreated textiles, in particular if they
have been preserved in a cold environment and if they are analysed with
NGS technology. Despite this possibility, genetic knowledge on breeds,
population structure and genetic traits is perhaps most likely to come from
other sources, such as parchment, bones or teeth. The downside of using
other materials is that they cannot be concluded to come from animals that
delivered wool for textile production. A benefit is that bones and teeth are
not limited to times of wool production (from the Bronze Age onwards),
but can provide evidence of the full domestication history of sheep. Most of
the studies of such archaeological sheep samples have focused on mtDNA,
whereas a few have demonstrated the possibility of also studying nuclear
markers (Niemi et al. 2013; Rannamäe et al. 2016). Recently, Teasdale et al.
(2015) managed to sequence 7–9 per cent of the sheep genome based on
historical parchment samples, which allowed the mapping of genetic affinity
to modern British sheep breeds.
CONCLUSION
The feasibility of recovering mtDNA from modern sheep wool and histor-
ical and archaeological sheep wool textiles from Denmark and Greenland was
demonstrated by Brandt et al. (2011) and Sinding et al. (2017). Subsequently,
Olivieri et al. (2012) and O’Sullivan et al. (2016) have succesfully confirmed
the potentials by analysing mtDNA from Copper Age sheep hair from the skin
garment worn by Ötzi. Other studies have been less successful in recovering
DNA from ancient wool (Brandt 2015; Nikulina 2016).Two factors seem to be
key to the success or failure to PCR-amplify mtDNA from archaeological wool
textiles and sheep hair: wool treatment processes and the burial environment.
The mordanting process and oxic, acidic and non-cold preservation environ-
ments seem to have a negative effect on the ability to recover aDNA from
wool when using traditional PCR-based methods. This is very likely because
these factors cause a faster fragmentation of the DNA strands. Therefore, each
textile’s chaîne opératoire, including treatment processes, context, storage history
and possible conservation, should be considered before sampling.
Textile samples from cold, anoxic environments with a relatively neutral
pH, and which have not undergone harsh treatment processes, seem to pre-
sent the best opportunities for aDNA research. Nuclear DNA has not yet
been successfully retrieved from ancient textiles. This may be due to a low
amount of DNA in hair combined with a more rapid degradation compared
with mtDNA. It is hoped that NGS technology will change this in the future.
292
The reviewed studies have demonstrated that aDNA in textiles is highly

degraded, emphasising the importance of using NGS, which does not require
target molecules of specific lengths in contrast to a PCR-based strategy. We
therefore recommend that future experiments on aDNA from textiles are
carried out with NGS technology in line with the recent study by Sinding
et al. (2017). In addition to all the genomic information in the sequence
data, NGS technology also offers much deeper insights into DNA preserva-
tion (average fragment length, C-T damage rates, contamination, etc.) than
can ever be obtained with the simple positive/negative signal from a PCR
reaction. In time, genome-scale analyses of ancient sheep facilitated by NGS
will potentially provide the opportunity to study selection and domestica-
tion processes, including the introduction of various genetic traits known
from modern sheep –for example, relating to wool type and quality. In that
context, however, it may be more feasible to use teeth or petrous bones (see
Appendix 12.4), which are arguably less valuable than ancient textiles and are
likely to provide a better quality of DNA.
Appendix 12.1. The DNA Code and the Genome

Deoxyribonucleic acid (DNA) is the hereditary material of all known living
organisms (apart from RNA vira). The DNA molecule is a double-stranded
DNA helix that stores the genetic information in a ‘code’ that is a sequence of
four chemical bases: adenine (A), thymine (T), cytosine (C) and guanine (G).
These bases pair with each other, so that A on one strand always pairs with T
on the opposite strand, and C on one strand always pairs with G on the other
(Fig. 12.3). The pairs are often referred to as base pairs (bp). Nucleotides are
arranged in two long strands that are reverse copies of each other.This code or
sequence of A, T, C, and G is translated by the cell to produce proteins that are
used in a wide array of functions in the body.
The complete DNA sequence is known as the genome and it is subdivided
into chromosomes. The chromosomes are located in the cell nucleus (nuclear
DNA) where they exist in two variants (in diploid organisms), one from each
parent. A small part of the genome is found outside the nucleus in the cell’s
energy- producing organelles, the mitochondria. This mitochondrial DNA
(mtDNA) is inherited only through the maternal lineage. Given the massive
difference in size between the nuclear and the mitochondrial genomes (c. 3
billion bp vs 16,569 bp in humans), nuclear DNA holds by far the most infor-
mation with regards to functionally important genetic traits and also provides
much higher resolution in population genetic studies. However, nuclear DNA
is only present in two non-identical copies per cell, whereas one cell can con-
tain thousands of mitochondrial genomes. Because of this numeric difference,
293
AP P E N D I X 1 2 . 2 : P C R AN D N G S 293
12.3 Representation of the DNA code.

(Graphics: Sidsel Frisch.)
mtDNA is generally easier to obtain from degraded material than nuclear

DNA and has thus traditionally been the focus of aDNA studies, before the
advent of Next Generation Sequencing (NGS, see Appendix 12.2).
Based on sequence variation in the mtDNA, an individual can be placed
into groups termed ‘haplogroups’ based on their shared variable sites
(polymorphisms). These mtDNA haplogroups can be used to explore the
ancestral relationships of individuals or populations based on their shared or
differing variable sites. However, with the invention of powerful sequencing
technologies, most genetic studies of modern populations are now based on
genome-wide analyses, where the information in millions of variable gen-
etic markers across the nuclear genome is explored, instead of restricting ana-
lyses to a few hundred variable sites in the mitochondria. This switch has also
happened in the field of ancient DNA.
Appendix 12.2. Polymerase Chain Reaction (PCR ) and Next

Generation Sequencing (NGS )
When Polymerase Chain Reaction (PCR) was introduced in the 1980s, it
revolutionised molecular biology. It allowed the production of millions of
294
294 ARCHAEO L O G I CA L W O O L T E X T I L E S AN D S H E E P G E N E TI C S
12.4 Schematic representation of the Polymerase Chain Reaction. (Graphics: Sidsel Frisch.)
copies of DNA from a single starting molecule of DNA, allowing the applica-
tion of Sanger sequencing without the tedious exercise of first having to clone
the DNA fragment into bacterial vectors. Briefly, PCR works by using cycles
of three thermal shifts which results in first opening the DNA double helix,
then binding small complementary DNA strands of generally 20 bp (primers)
to pre-selected and known target regions of DNA (such as a well-characterised
gene, or a haplogroup-defining segment of mitochondrial DNA). Lastly, the
enzyme Taq polymerase extends the strand from the primer along the target
template molecule which results in the formation of two identical copies of
the starting strand (Fig. 12.4). By repeating this cycle, millions of identical
copies of the target DNA (typically lengths of 200–1,000 bp) can be generated
in a short amount of time, allowing for DNA sequencing.This technology was
also widely useful in aDNA studies where the number of starting molecules is
often very low. However, because DNA degrades through time (e.g. Allentoft
et al. 2012), archaeological DNA is often very fragmented and rarely exceeds
200 bp (see Appendix 12.4). This is unfortunate as the DNA targets recovered
by PCR have to be long enough to bind two primers of about 20 bp each
as well as the sequence of interest (min. 30 bp). In archaeological samples
from harsh environments, DNA fragments of this length may simply not be
preserved, making PCR a very inefficient tool. Also, because of the extreme
295
AP P E N D I X 1 2 . 2 : P C R AN D N G S 295
12S rDNA
Control region Phe
Thr Val
16S rDNA
Cyt b
Pro Leu
ND1
Glu
ND6 IIe
Gln Met
ND5
Ala ND2
Asn
Cys
Leu Trp
Ser Tyr
His
Ser
ND4 CO I
ND4L Asp
Arg
ND3 CO III Lys CO II
Gly ATpase
6 and 8
12.5 Representation of the location of mitochondrial and nuclear DNA, respectively, in

the cell.
(Graphics: Sidsel Frisch.)
sensitivity of PCR, contamination from a modern DNA source is a major

challenge because in the absence of any authentic DNA, even a few contam-
inating molecules can be copied into millions during the amplification process,
yielding a false positive result for the sample in question.
The introduction of Next Generation Sequencing (NGS) in 2005 offered
the opportunity to generate billions of different target DNA sequences in a
short amount of time (e.g. Metzker 2010). In contrast to one or a few pre-
selected target genes in PCR-based research, this allows for the sequencing of
complete genomes of any organism (Fig. 12.5) – including ancient ones (e.g.
Rasmussen et al. 2010). Moreover, this technology can sequence much shorter
DNA sequences than offered by PCR, which enables working on samples with
a higher fragmentation of DNA. When combined with new DNA extraction
296
methods retaining shorter fragments (e.g. Allentoft et al. 2015), this is an
extremely powerful and useful technology for aDNA research. Lastly, NGS data
also allows for a thorough investigation of DNA authenticity, partly by assessing
DNA damage patterns (e.g. Briggs et al. 2007; Ginolhac et al. 2011), and partly
by testing for the presence of DNA from more than one source (Renaud et al.
2015). This does not imply that DNA contamination is no longer a problem in
aDNA research, but it does mean that our ability to identify DNA contamin-
ation has much improved with the shift to NGS-based experiments.
Appendix 12.3. Extraction and Amplification of DNA from

Archaeological Textiles Made of Animal Fibres
Prior to DNA extraction, archaeological and historical textile samples (gen-
erally between 10 and 200 mg) are preferably washed briefly in a 10% bleach
solution to decontaminate them from external contaminant DNA.The crucial
step with hair extraction is the ‘digestion’ step in which the hair is dissolved
to get the DNA molecules into solution. This is challenging, as hair can be
very hard to dissolve. In Brandt et al. (2011), the wool samples were incubated
at 56°C overnight in a solution (buffer) containing the enzyme proteinase-K,
Tris, NaCl, SDS, EDTA, CaCl2, H2O and DTT developed specifically for hair
by Gilbert et al. (2004). The next step is the purification of DNA from the
digested solution, which can be carried out in a variety of ways. Brandt et al.
(2011) used the Qiagen ‘QiaQuick’ kit, applying the PCR Purification Spin
Protocol, which captures DNA molecules on silica filters from where they are
washed and then released again in EB buffer (QIAgen). Brandt (2015) applied a
refined extraction protocol, digesting the wool samples overnight at only 37°C
(to avoid further fragmentation of the DNA) and using a silica-in-solution
approach, where DNA binds to silica particles in the presence of guanidium
thiocyanate Tris, NaCl and EDTA (e.g. Orlando et al. 2011), followed by puri-
fication with 80% ethanol, and then resuspended in EB buffer (Qiagen).
In case of a PCR amplification assay (see Appendix 12.2), the target gene will
often be project-specific and depends on the aim. Brandt et al. (2011) aimed
at demonstrating the potentials of DNA from wool and used a PCR assay
targeting a 121 bp region of the mtDNA 16s gene using primers 16sMam3
and 16sMam4 (Haile et al. 2009) and a 66 bp nuclear DNA segment of exon
10 in the sheep prolactin receptor gene (PRLR). Successful PCR products
were purified, cloned, and sequenced (8 clones per PCR). Species identity
was confirmed as 100% identity matches, against a database of 16s mammalian
sequences. In contrast, Brandt (2015) specifically targeted ancient sheep DNA
with a PCR assay designed to target a 97 bp amplicon of the sheep D-loop,
using primers Ovis-1f and Ovis-1r (Brandt et al. 2011). Unfortunately, none
of these samples proved successful.
297
AP P E N D I X 1 2 . 4 : D N A D AMAG E 297
Appendix 12.4. DNA Damage

DNA damage, such as that caused by hydrolytic and oxidative processes
(reviewed in Lindahl 1993; Willerslev and Cooper 2005), occurs rapidly even
in living tissue. There damage is repaired by enzymatic mechanisms in the cell.
This is not the case in dead organisms (Lindahl 1993), where DNA damage
will accumulate over time. Such steady accumulation of damage implies that
the DNA in ancient samples is often highly fragmented (i.e. the average lengths
are most often below 100 bp) and thus beyond access for PCR-based research
(see Appendix 12.2). Moreover, because DNA fragmentation follows an expo-
nential decline model, it is possible to calculate an average fragmentation rate
when the age of the sample is known (Allentoft et al. 2012).
Because the authentic endogenous DNA degrades to small fragments
beyond analytical recovery (i.e. <30 bp), the bulk of DNA analysed from an
ancient sample will often be microbial DNA, derived from the preservation
environment (i.e. soil bacteria). Indeed, the endogenous DNA content (DNA
from the target organism) is below 1 per cent in most ancient human samples
(e.g. Allentoft et al. 2015), making genome-scale sequencing of ancient DNA
samples a very costly exercise. In skeletal material, this can be partly mitigated
by carefully sampling either the cementum layer in teeth roots (Damgaard et al.
2015) or the otic capsule inside the petrous bone (Pinhasi et al. 2015), which
have both been demonstrated to yield a high endogenous DNA content.
In addition to age, temperature is another key factor for biomolecule pres-
ervation since the rate of both the spontaneous and enzymatic degradation
processes decreases at lower temperatures (Lindahl 1993; Lindahl and Nyberg
1972; Smith et al. 2001). This explains why the first complete ancient genome
was recovered from a Greenlandic Palaeo-Eskimo (Rasmussen et al. 2010), and
why the current well-accepted aDNA age-record is ascribed to a c. 700,000-
year-old horse bone recovered from permafrost in Alaska (Orlando et al. 2013).
Other factors like soil pH, oxygen presence, water saturation, storage time and
conditions are also involved in determining the limit of long-term DNA pres-
ervation, but the effects of these are less well understood.
Another defining damage feature of aDNA is introduced by post-
mortem modifications of the nucleotides close to the strand breaks, causing
misinterpretations by the polymerase enzyme in the PCR reaction. The result
when the DNA is sequenced is an apparent accumulation of C to T nucleotide
changes in the 5’ends of the DNA molecules and G to A changes in the 3’ends
(e.g. Briggs et al. 2007).This damage pattern expected for aDNA can therefore
be used to support the authenticity of the DNA molecules, for example using
the software package MapDamage (Ginolhac et al. 2011).
Despite many poorly understood factors, the level and character of DNA
damage is largely determined by age and by the preservation environment.
298
Therefore, knowledge of the preservation and burial environment is of great

importance when evaluating the potentials of aDNA analysis. Also, in many
cases, a direct link between the visual and molecular preservation state exists
(e.g. Hansen et al. 2017). If a sample appears very poorly preserved, for example
brittle and ‘chalky’ bone material from a temperate or warm preservation
environment, it will very rarely produce good DNA results.
NOTES
1 Unpublished data analysed by M. Allentoft.

2 Unpublished data analysed by M. Allentoft.
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Archaeological Wool Textiles: A Window into Ancient Sheep

Chapter · November 2019

Luise Ørsted Brandt

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ARCHAEOLOGICAL WOOL TEXTILES:

Luise Ørsted Brandt and Morten Allentoft

Research in archaeological textiles has traditionally focused on identifying

276 Archaeological W ool T extiles and S heep G enetics

PRESERVATION OF ARCHAEOLOGICAL TEXTILES AND THE DNA POTENTIAL

T HE T E X T I L E C HAÎ NE O P É RAT O I RE AN D D N A D E P O S I TI O N 277

THE EFFECTS OF THE TEXTILE CHAÎNE OPÉRATOIRE AND

278 Archaeological W ool T extiles and S heep G enetics

T HE T E X T I L E C HAÎ NE O P É RAT O I RE AN D D N A D E P O S I TI O N 279

280 Archaeological W ool T extiles and S heep G enetics

should therefore also potentially be possible to recover DNA molecules from

SAMPLING ARCHAEOLOGICAL TEXTILES FOR GENETIC RESEARCH

When sampling archaeological material for aDNA research it is important

DNA EXPERIMENTS ON ARCHAEOLOGICAL TEXTILES: AN OVERVIEW

DN A E X P E R I M E N T S O N ARC H AE O L O G I C AL TE XTI L E S 281

and nuclear DNA from a set of archaeological and historical textiles,

286 Archaeological W ool T extiles and S heep G enetics

tested with a PCR assay targeting a 97 bp fragment of mtDNA, but these

DNA AND TEXTILES FROM BOGS AND BURIALS

samples analysed by Brandt et al. (2011) yielded 121 bp sequences of mtDNA,

288 Archaeological W ool T extiles and S heep G enetics

D N A and Textiles from B ogs and B urials 289

290 Archaeological W ool T extiles and S heep G enetics

MITOCHONDRIAL AND NUCLEAR DNA: PROS AND CONS

In addition to species identification, mtDNA data can sometimes be used

292 Archaeological W ool T extiles and S heep G enetics

The reviewed studies have demonstrated that aDNA in textiles is highly

Appendix 12.1. The DNA Code and the Genome

12.3 Representation of the DNA code.

mtDNA is generally easier to obtain from degraded material than nuclear

Appendix 12.2. Polymerase Chain Reaction (PCR ) and Next

12.4 Schematic representation of the Polymerase Chain Reaction. (Graphics: Sidsel Frisch.)

12.5 Representation of the location of mitochondrial and nuclear DNA, respectively, in

sensitivity of PCR, contamination from a modern DNA source is a major

296 Archaeological W ool T extiles and S heep G enetics

Appendix 12.3. Extraction and Amplification of DNA from

Appendix 12.4. DNA Damage

298 Archaeological W ool T extiles and S heep G enetics

Therefore, knowledge of the preservation and burial environment is of great

1 Unpublished data analysed by M. Allentoft.

300 Archaeological W ool T extiles and S heep G enetics

302 Archaeological W ool T extiles and S heep G enetics

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Luise Ørsted Brandt and Morten Allentoft

MITOCHONDRIAL AND NUCLEAR DNA: PROS AND CONS

Appendix 12.1. The DNA Code and the Genome