Barley in Archaeology and Early History

Barley in Archaeology and Early History
Oxford Research Encyclopedia of Environmental

Science
Simone Riehl
Subject: Agriculture and the Environment Online Publication Date: Mar 2019
DOI: 10.1093/acrefore/9780199389414.013.219
Summary and Keywords
In 2018 barley accounts for only 5% of the cereal production worldwide, and regionally
for up to 40% of cereal production. The cereal represents the oldest crop species and is
one of the best adapted crop plants to a broad diversity of climates and environments.
Originating from the wild progenitor species Hordeum vulgare ssp. spontaneum,
biogeographically located in the Fertile Crescent of the Near East, the domesticated form
developed as a founder crop in aceramic Neolithic societies 11,000 years ago, was
cultivated in monocultures in Bronze Age Mesopotamia, entered the New World after
1492 CE, reached a state of global distribution in the 1950s and had reached
approximately 200 accepted botanical varieties by the year 2000.
Its stress tolerance in response to increased aridity and salinity on one hand and
adaptability to cool climates on the other, partially explains its broad range of
applications for subsistence and economy across different cultures, such as for baking,
cooking, beer brewing and as an animal feed.
Although the use of fermented starch for producing alcoholic beverages and foods is
globally documented in archaeological contexts dating from at least the beginning of the
Holocene era, it becomes concrete only in societies with a written culture, such as Bronze
Age Mesopotamia and Egypt, where beer played a considerable role in everyday diet and
its production represented an important sector of productivity.
In 2004 approximately 85% of barley production was destined for feeding animals.
However, as a component of the human diet, studies on the health benefits of the
micronutrients in barley have found that it has a positive effect on blood cholesterol and
glucose levels, and in turn impacts cardiovascular health and diabetes control. The
increasing number of barley-breeding programs worldwide focus on improving the
processing characteristics, nutritional value, and stress tolerance of barley within the
context of global climate change.
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Keywords: Crop plant evolution, cultural history of crop species, domestication, cereal production, beer
production, archaeology of crop species, archaeobotany, agronomy
Systematics and Taxonomy of Wild and

Domesticated Barley Forms
Barley (Hordeum vulgare L.),1 Poaceae family, Triticeae tribe, is a diploid taxon consisting
of two-row forms (Hordeum vulgare convar. distichon (L.) Alef.), with sterile lateral fruits,
and poly-row forms, bearing fertile lateral fruits (Hordeum vulgare ssp. vulgare, in
references before 1990 sometimes referred to as Hordeum vulgare ssp. agriocrithon2)
(Fig. 1). Approximately 200 botanical varieties are included within the domesticated
group (Hanelt et al., 2001). Both the two- and the poly-rowed form produce hulled and
free-threshing varieties.
Domesticated barley
species and landraces, like
their wild progenitor
species Hordeum vulgare
ssp. spontaneum (C. Koch)
Thell., belong to the
primary gene pool of
barley, while Hordeum
bulbosum L. represents
the secondary gene pool,
difficult but not impossible
to cross with Hordeum
vulgare species.
Barley species are

inbreeders and reproduce
Figure 1. Morphology of two-row (a–c) and poly-row
to about 99% by self-
(d–g) barley: two-row barley in frontal view (b) with a
central fertile grain, and two lateral sterile grains, pollination (Von Bothmer,
(c) ear from above; Poly-row barley in frontal view (e) 1992). Outcrossing is
with three fertile grains and ear from above; (f) six-
row variant, (g) four-row variant); rachis internode of promoted by cool and
wild barley with smooth attachment area (h) und moist environmental
rachis internode of domesticated barley with rough
conditions, probably due to
attachment area (i); a–g after Jacomet and Kreuz (19
99), h and i from van Zeist, Smith, Palfenier-Vegter, the increased viability of
Suwijn, and Casparie (1984). pollen under such
conditions.
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Ecology and Biogeography of Barley

The wild progenitor of domesticated barley, Hordeum vulgare ssp. spontaneum (C. Koch)
Thell., exhibits a broad ecological amplitude and a considerably broader geographic
distribution than wild wheats. Harlan and Zohary (1966) observed that the primary
habitats for barley differ from those of the wheats, with wild barley appearing in drier
habitats, extending into the steppes and deserts along the wadis. Outside its distribution
center in the Fertile Crescent, wild barley grows mostly as a segetal or ruderal plant.
Wild barley frequently occurs in Near Eastern open, oak-dominated woodland down to
200–250 mm of mean annual precipitation. Michael Zohary (1973) mentions the species
as part of several plant associations in Palestine (Cis- and Transjordan). These are open,
pine-dominated or semi-steppe batha associations (e.g., Ziziphetum loti pastorale) of
habitats formerly dominated by oak park forest (Quercus ithaburensis), but are
exterminated if extensive grazing occurs.
Domesticated barley has a shorter vegetative cycle than wheat, and is more stress-
tolerant in response to increased salinity and aridity. The crop is heat-tolerant in a dry
climate and moisture-tolerant in a cool one, but performs poorly under hot and humid
conditions (Poehlman, 1985). As noted by Braun (1995), referring to Garnsey (1988),
statistical data on traditional farming in Greece for the period between 1931 and 1960
indicated that wheat failure occurred over five times more frequently than barley failures.
The cultural effects of these properties may be seen in the fact that barley is used for
bread-baking only in geographic regions with extreme climate conditions that limit
cultivation of wheat, rye, or oats (Körber-Grohne, 1987). Furthermore, barley has also
been found to be more competitive with weeds than wheat species are, due to a greater
tillering ability and below-ground root system.
It has been shown that the genome combinations in modern barley landraces descend
from different regions of the Fertile Crescent, which has been interpreted as supporting
their relatively late evolution (Poets, Fang, Clegg, & Morrell, 2015). Genomic analysis of
6,000-year-old barley from the southern Levant, however, has indicated that ancient and
modern landraces are genetically similar (Mascher et al., 2016), thus suggesting an early
evolution of landraces. Permanent mutations and segregation products of crosses
probably complicate the long-term reconstruction of barley evolution, indicated in
morphological overlap, which often hinders the differentiation between true, wild
spontaneum, and primitive forms and landraces of cultivated barley (Backes, Orabi,
Fischbeck, & Jahoor, 2006). Furthermore, genetic research on wild barley occupying a
multi-slope transect at Lower Nahal Oren, Mount Carmel, Israel, found evidence for twin
evolutionary processes of adaptation and speciation (Yang, Zhang, Bolshoy, Beharav, &
Nevo, 2009). The wild barley samples displayed dramatic interslope adaptive and
speciational divergence on an “African” dry slope and a “European” humid one, which
cannot be explained by mutation, gene flow, or chance effects, but supports adaptive
natural microclimatic selection as a major evolutionary divergent driving force. These
results are crucial for understanding the role of gene and genome biodiversity on locally
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diverging microclimatic units and would at least partially explain the interregional
diversity in archaeobotanical assemblages at the onset of barley cultivation.
Evolutionary History of Wild and Domesticated

Barley
Barley (Hordeum vulgare ssp. vulgare) alongside the hulled-wheat species, pulses, and
flaxseed, are among the first domesticated species, i.e. the “Founder Crops,” of the
Fertile Crescent.
Traits of domestication, which after Hammer (1984) can be summarized under the term
“domestication syndrome,” comprise changes in reproduction features (i.e. brittle vs.
tough rachis internodes), in seed size, the kernel-row type, some of which are related to
different geographic regions, hulled vs. naked kernels, loss of photoperiod sensitivity and
of seed dormancy, and the growth habit of the plants (Backes et al., 2006). The first three
characteristics can be morphologically recognized in archaeobotanical material, whereas
the latter three cannot.
While wild cereals reproduce spontaneously upon ripeness through disarticulation of the
ear into the smaller units (triplets) at predefined abscission areas, these areas have lost
their function in domesticated species and usually break above the attachment area only
during the threshing process (Fig. 1). Such differences in the reproduction behavior of
the wild and the domesticated forms result in different morphological outlines of the
attachment areas in the rachises (disarticulation zone; Jacomet & Kreuz, 1999; Zohary,
Hopf, & Weiss, 2012), thus the phenotypic morphology of the barley rachis serves
archaeobotanists to determine the transition from wild progenitors with a brittle floral
axis (rachis) to domesticated forms with a tough rachis (e.g., Weide, Riehl, Zeidi, &
Conard, 2015). The genetic background of the differences in the disarticulation zone
derives from independent recessive mutations that cause cell-wall thickening, converting
the wild type into a tough, non-brittle form that promotes grain retention
(Pourkheirandish et al., 2015).
Ancient DNA studies have contributed to an assessment of selection during different

stages of domestication (Gutaker & Burbano, 2017). These authors state that limiting
domestication studies to the phenotypic and/or molecular characterization of
contemporary cultivated plants followed by comparison to their extant wild relatives only
allows us to identify the final effect of thousands of years of crop evolution, while
evolutionary processes during different stages of domestication (i.e., pre-domestication,
fixation of domestication syndrome traits, geographic expansion, and nutritional
improvement) can only be recognized through retrospective approaches, i.e. additional
consideration of historical samples, including archaeobotanical remains. This also applies
to the supposed multiple geographic origins of domesticated barley forms.
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As early as 1936, Vavilov (cited by Schiemann, 1948) noted the existence of 2–3 centers of
diversity for barley in the geographic region of the northern Fertile Crescent (i.e., the
historical Eastern Anatolia, extending to Syria-Palestine, Armenia, and Western Persia)
and the mountain areas of Eastern Asia (i.e. historical eastern Tibet, the Eastern
Himalayas as far as Japan). More recent genetic studies of modern barley suggest at least
two different regions in which domestication events took place, one in the Levant and a
second in Iran or further east, which evolved approximately 2000 years later (Morrell &
Clegg, 2007; Fig. 2). The fact previously noted by Zohary (1973), that wild barley is
extremely polymorphic on the morphological and ecological level, provides the logical
explanation to later genetic findings of multiple centers of origin of domesticated barley
(cf. Yang et al., 2009).
Pourkheirandish et al. (2015) also suggest two regions of anthropogenic selection for the
non-brittle phenotype, i.e. the Southern and Northern Levant; btr1-type barleys emerged
in the Southern Levant prior to the appearance of btr2-types in the Northern Levant.
They also analyzed Central Asian accessions and suggest that these were derived from
material spreading by humans from the Levant.
The inclusion of
quantitative data on
archaeobotanical findings
(e.g., seed or chaff counts)
allows us to estimate the
changing selection
coefficients for the
evolution of non-shattering
barley, as suggested by
Allaby, Stevens, Lucas,
Maeda, and Fuller (2017),
who found that their data
support two discernible
Figure 2. Genetically investigated accessions of wild
barley with sequencing of 18 loci (asterisks— barley domestications in
accessions with sequencing of four loci): Red— the Southern Levant.
Samples with majority assignment to the eastern
cluster, Blue—Samples with majority assignment to
the western cluster; Black triangles – Neolithic sites
with early evidence of domesticated barley (Morrell
& Clegg, 2007).
Barley in Human Subsistence (Pleistocene Through the Transition to

the Holocene Era)
Particular characteristics of these cereals, such as the combination of high calorific value
with a high proportion of carbohydrates, considerable seed production that takes place in
short reproduction cycles of few weeks or months, and their suitability for long-term
storage may have represented attractive advantages at the very beginnings of
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agriculture, and probably determined their dominance in archaeobotanical assemblages

from the mid-late Neolithic onwards and in human nutrition up to the 21st century.
Early records of wild-barley consumption derive from Middle Paleolithic Kebara Cave in
Israel and Shanidar III in northern Iraq, dating to at least 60,000 and 44,000 years BP
respectively (Lev, Kislev, & Bar-Yosef, 2005; Henry, Brooks, & Piperno, 2011). Only a
limited number of Epipaleolithic sites have been studied for their plant remains. At
Epipaleolithic Ohalo II in Israel (approx. 23000 BP) wild barley is also very frequent and
has been systematically gathered (Kislev, Nadel, & Carmi, 1992), if not cultivated jointly
with other wild progenitor cereals and pulses (Snir et al., 2015). Iraq ed-Dubb (Colledge,
2001), Wadi Hammeh 27, and Ha-Yonim Cave (Hopf & Bar-Yosef, 1987), all geographically
located in the Southern Levant, also contain deposits that have been dated to the
Epipaleolithic. These are generally dominated by wild barley seeds, while other wild
progenitor species of modern crops are rare. However, in Northern Mesopotamian sites,
such as Abu Hureyra (Hillman, Hedges, Moore, Colledge, & Pettitt, 2001) and Tell
Mureybit (van Zeist & Bakker-Heeres, 1984), wild einkorn finds are dominant, although
these have no center of domestication in the Southern Levant (Heun, 1997; Heun,
Haldorsen, & Vollan, 2008; Kilian et al., 2007). Although the low number of sites for this
period prohibits any statistical assessment, the geographic differences of dominant wild
cereal species may relate to their ancient biogeography and/or reflect human choice as
part of an opportunistic behavior. As at Tell Abu Hureyra, rye (Secale sp.) is also
recorded, which may be indicative of cooler and moister conditions, this supports the
argument of biogeographic differences occurring due to different regional climatic
conditions.
For the pre-pottery Neolithic A (PPNA, approx. 9500–9000 BCE) there are considerably
fewer archaeological sites than for the Pre-Pottery Neolithic B (PPNB). Conventional
thinking still holds that PPNA represents the beginnings of cultivating wild cereals in a
small number of sites in the Fertile Crescent, which are otherwise dominated by non-
progenitor species. Many of the early sites with indications of wild cereal cultivation show
archaeological evidence of changing occupational structures and communal, monolithic
architecture, accompanied by storage installations, a large-sized, often immobile grinding
stone, and occasionally also sickle blades (Willcox & Stordeur, 2012; Riehl, Zeidi, &
Conard, 2013), in sum indicating changes in population and social organization (Watkins,
2008, 2010, 2013). Similarly, as during the Epipaleolithic era, wild barley is the taxon that
is represented best at the majority of PPNA sites, while at singular locations other wild
progenitor species of modern crops are dominant, for example, wild einkorn at Tell
Mureybit, a wild vetch form at M’lefaat (Savard, Nesbitt, & Gale, 2003). For Tell Aswad
the dominant taxa have been identified as domesticated emmer and einkorn (van Zeist &
Bakker-Heeres, 1982, but see also Willcox & Stordeur, 2012).
The evidence of crop-cultivating communities changes dramatically during the pre-

pottery Neolithic B, when phenotypically domesticated cereals become more frequent,
and fully domesticated crops were implemented during the following phases. Apart from
a massive increase of sites during this period, barley no longer represents the dominant
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species in most of the locations, but the hulled-wheat species, as well as lentils become
more prominent in the majority of the settlements. Both domesticated and wild barley
appear in higher counts, mainly in the southeastern regions of the Fertile Crescent.
The process from incipient cultivation of wild progenitors of modern cereals to fully
domesticated crop species is conventionally assumed to cover a range of several hundred
to approximately 2000 years, a model that has been described as “protracted
domestication” (Allaby, Fuller, & Brown, 2008), in contrast to a “rapid domestication
event” (Heun, Abbo, Lev-Yadun, & Gopher, 2012), according to which traditional model, a
high level of artificial selection gives rise to the rapid and geographically definable origin
of crop domestication (Allaby et al., 2015). In fact, this only seems plausible for einkorn
wheat. Starting dispersion of domesticated species also becomes visible during the PPNB,
including at Ҫatal Höyük, Can Hasan III, and Aşikli Höyük in Turkey, as well as sites on
Cyprus (Willcox, 2001). The evidence for Iran is still limited, but emmer appears to be the
first domesticated species, although wild barley has been used in subsistence for several
hundreds of years (Riehl et al., 2013). Overall, the archaeobotanical evidence supports
the model of a protracted domestication process. Computational models applied to
phylogeographic data revealed that genetic data diversity is also compatible with the
notion of a protracted origin (Allaby et al., 2008).
The Evolution and Dispersal of Domesticated Barley Species From

the Fertile Crescent Into Other Geographic Regions (Holocene Era)
Considering barley cultivation from the historical perspective, the beginning of its
domestication is probably, by a considerable distance, the topic on which the most has
been published.
It is commonly accepted that while the domesticated two-row barley form occurs at the
very beginning of agriculture, poly-row forms appear somewhat later in the PPNB,
including six-row barley bearing three fertile grains per triplet instead of one as in the
two-row form. The later appearance of six-row barley, only in relation with already
cultivated assemblages or as a weed species, has been described by Zohary in 1963 as a
phenomenon of environmentally adaptive specialization. According to him the arrow-like
shape of the triplet in two-row forms ensures that the seeds will bypass impenetrable
surfaces (e.g., rocks) and reach soil when they fall to the ground. The different shape of
the six-rowed mutants excludes such an environmental advantage, thus selection in the
natural original habitats of wild barley acts against the survival of six-rowed mutants, and
would explain their relatively late appearance in crop assemblages, as well as their
repeated origin at different times and in different regions (Komatsuda et al., 2007).
The spread of agriculture from the Fertile Crescent into macro- and micro-climatically
different regions with altered abiotic and biotic conditions far away from the
biogeographic distribution of the wild progenitor species has been described as one of
the greatest challenges in the evolutionary history of the crop species (Allaby et al.,
2015). Ancient DNA (aDNA) studies provide tools with which to directly address
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evolutionary processes in crop plants. The technology of analyzing ancient genomes in

mostly carbonized plant remains is however still in its infancy, and has only recently
produced its first reliable results. In uncharred plant remains warm temperatures at the
most ancient archaeological sites limit preservation of ancient biomolecules to just a few
thousand years, which disables the investigation of the origins of domestication (Fig. 3).
Dispersal of agricultural
production into regions
beyond the Fertile
Crescent accompanied
adaptation processes
within the particular
founder crop species and
supported the recognition
and valuation of other
advantages by cultivators
and farmers, such as the
generally higher number
of grains per ear in the
Figure 3. Ancient DNA content of desiccated barley case of the six-row barley
seeds over time in Qasr Ibrim, North Africa (taken form. Remains of six-row
from Allaby et al., 2015)
barley have been
recovered at a number of
sites in southeastern Turkey, for example, at aceramic Neolithic Çatalhöyük around 9,000
BP and in late Neolithic Haçilar around 8,000 BP (D. Zohary, Hopf, & Weiss, 2012).
Dispersal into the Indian subcontinent and Egypt dates at least back to the 7th mill. BP,
and from there it was carried into the highlands of Ethiopia. Cultivation under a wide
range of ecological conditions led to the establishment of the secondary center of genetic
diversity that has been identified in this region (Backes et al., 2006). Dispersal to Tibet
has been dated to the Tibetan Late Neolithic period (6000–4500 BP) and the initial
adoption of barley is assumed to have enabled farmers to buffer shortfalls of other crops,
such as millet, below 2500 m.a.s.l. or during anomalously cold years (Barton, 2016;
d’Alpoim Guedes, Lu, Hein, & Schmidt, 2015).
The dispersal patterns of barley in Europe are highly diverse. While in Eastern Europe
the crop species was well reflected in the Neolithic, it occurs only sporadically in sites of
the early Linear Pottery Culture (LBK). Kreuz and Marinova (2017) interpret this pattern
as representing a deliberate reduction of crop species diversity “based on [the] expertise”
of the ancient farmers, which the authors seem to consider as being climate-related.
As outlined in section 2, cultivated barley is adapted to a broader range of environments

than any other cereal species. The optimal range is on well-drained, fertile loam soils
under relatively cool temperatures (15–30°C) and a moderate annual rainfall (500–1000
mm), thus the highest yields are achieved in central and northern Europe (MacGregor &
Bhatty, 1993). Its high tolerance for cool temperatures may also explain the importance of
Page 8 of 26


barley for breadmaking in northern Europe from medieval times onward (Körber-Grohne,
1987). In their retrospective of archaeobotanical barley finds in Central Europe, Šálková,
Benes, Komárová, and Vanecek (2012) emphasize the continuously increasing importance
of barley from the Neolithic until medieval times in association with its use for beer
brewing, which they consider to be supported by archaeological findings, such as
drinking sets and biochemical analyses of pottery residues.
In Europe poly-row forms have been cultivated as winter crops since at least medieval
times, while two-row forms prevail in summer cropping and are preferably used for beer
brewing. Macronutrient composition in taxa used for brewing differs from others in its
higher starch percentage and comparatively low protein content, both of which are
affected by climate; i.e. under dry and warm conditions the protein content is higher.
While being the earliest cereal known to be cultivated and domesticated in the Old World,
barley entered the New World only after 1492 AD, grown in a colony founded by
Columbus, and spread from there into other areas, mostly to North America, but also
Mexico. By 1958 it was distributed all over the continent with major cultivation in the
northernmost geographic regions, such as the Canadian states of Alberta, Sasketchewan,
Manitoba, and the US states of North Dakota and Montana (Hockett, 1991).
The Development of Barley Cultivation in the Near East and Europe

(Later Neolithic Era to the Bronze and Iron Ages)
Archaeobotanical and textual evidence from Bronze Age Mesopotamia suggests that
societies during this extended period of time were basically producing barley
monocultures (Riehl, Pustovoytov, Dornauer, & Sallaberger, 2012), and were likely also
trading large amounts of these cereals (Sallaberger, 2016). Barley was also a preferred
animal feed with which to fatten domesticated ungulates before slaughter, and was used
for feeding “plough oxen” and donkeys (Sallaberger, 2004). Today up to one third of the
barley supply is used for grain and straw for animal feed (MacGregor & Bhatty, 1993).
Reasons for the heavy focus on barley during the Bronze Age include an assumed
nutritional depletion of cultivated soils and issues of salinization (Zohary et al., 2012), as
well as climatic change at the end of the 3rd millennium BCE (Riehl, 2008, 2012).
As archaeobotanical findings suggest, two-rowed barley was also the most frequently
cultivated form in Bronze and Iron Age Near Eastern settlements, whereas in central and
northern Europe the two-row form was virtually absent at least until the end of the first
millennium BCE (Körber-Grohne, 1987). This may be related to the species’ high level of
drought tolerance and its short life cycle, resulting in sufficient yields, reaching a
minimum of 200–300 mm of mean annual precipitation. This is confirmed by ancient
textual evidence from northern Mesopotamia (Rattenborg, 2016), and as has been noted
by Charles (1984) biogeographic differences between the two-rowed barley cultivated in
upland dry-framing regions and the six-rowed varieties predominating in alluvial plains
have been observed in Iraq.
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Winter barley requires vernalization, i.e. a cold stimulus for initiation of floral
development. Spring barley is adapted to warmer environments and does not need
vernalization. Whether there have been regions in the past where barley was cultivated
as a winter crop is difficult to assess. In the Gezer calendar the sowing of cereals is
attributed to the months between October and January for a harvest in April, suggesting
cultivation of winter barley (Dalman, 1987, vol. 1). However, according to recent studies,
quality traits do not differ between autumn- and spring-sown conditions (Badeck, 2016).
The particular characteristics of barley can be considered as supportive for the

development of agricultural technologies after the establishment of Neolithic cultivation.
Its early maturity and harvests and general stress tolerance provide an ideal basis for
intercropping and crop rotation systems, also helping to control weeds and diseases
(Hockett, 1991).
Naked forms of barley appear in the eastern European record as early as the early
Neolithic period. Archaeobotanical differentiation of hulled and naked grains is, however,
limited by preservation affecting the surface layers of cereal grains. For southwestern
Europe Tereso et al. (2016) note the replacement of naked barley by hulled barley
throughout the Bronze Age, but do not offer an explanation. Sturite et al. (2018) tested
hull-less and hulled barley varieties in field experiments in Latvia and Norway in order to
assess the varieties’ adaptability across environments (sites) and found that grain yield of
hull-less barley varieties was significantly lower than for hulled barley varieties,
regardless of climatic conditions and management systems, but the effect was less
pronounced in organic farming systems. On the other hand, hull-less barley varieties
tended to contain more protein and β-glucans than hulled barley varieties. However, other
studies indicate that there are no significant effects on the fatty-acid composition of milk
from lactating dairy cows from the use of feed containing either hulled or hull-less barley
(Yang et al., 2018). Reasons for cultivating either hulled or naked barley in the past may
thus be found in ancient agricultural technology, and one may consider that while naked
barley may be less labor-intensive during threshing, hulled barley may have the
advantage of higher pest and damage resistance during storage.
Dietary Use of Barley

Although today rarely used as a component of bread, barley remains in archaeological
finds in Neolithic lakeside settlements have been interpreted as being derived from bread
(Heiss et al., 2017).
Barley bread with admixture of other plant seeds like lentil can be traced as a very old
tradition in some of the eastern Mediterranean regions, such as Egypt and the Southern
Levant, and is found in biblical texts (Löw, 1967; Dalman, 1964). Mixed barley bread is
also known to have been produced in southwestern Germany from intercropping yields of
barley and lentil stands in the 19th century (Körber-Grohne, 1987), although this was
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mostly done in times of distress, for example, when cereal flour was rare and
economically expensive.
Gluten content of barley grain is 27% lower than in wheat, which older sources (e.g.,
Braun, 1995) have seen as an explanation of the less common use of barley for bread
baking, as gluten is responsible for the rising of dough. Instead, as far back as antiquity,
barley was used in the human diet in the form of roasted grain, flat unleavened bread,
cakes, porridge, or groats. Although roasting of unhardened barley grain seems to have
been a common practice in antiquity, the potential beneficial effects of this practice on
the digestibility of the grain have not been considered.
Generally, the use of barley for grain consumption by humans and other animals has
received less scholarly attention than its role in beer brewing. Today malt production is
the second most important processing form of barley, and numerous research articles
document prehistoric and historic knowledge of the fermentation process, including
alcohol production (e.g., Dietler, 2006; McGovern, 2009; Meussdoerffer, 2009). The aspect
of the particular health benefits of fermented foods, in terms of digestibility of the food
and bioavailability of micronutrients, has regained attention recently in relation to
increasing rates of gastrointestinal diseases.
The use of fermented starch to produce alcoholic beverages and foods has been globally
documented in a broad range of different plant-utilizing cultures, starting as early as the
aceramic Neolithic.
During fermentation, organic substances are transferred into acids, gases, and alcohol,
with involvement of bacteria and enzymes, in most cases without the involvement of
oxygen, although oxidative fermentation also exists, for example, in acetification. While
malolactic fermentation is the precondition for cheese production, alcoholic fermentation
of plant starch is used in beer production. An ethnohistorical classification of beer
according to the starting product, type of processing and microbes involved is provided in
Rosenstock and Scheibner (2017). Today there is an extremely broad range of beer styles
whose characteristics, cultural origin, and geography are described in the scientific and
popular literature (e.g., Patterson & Hoalst-Pullen, 2014). An overview of the processing
steps for the production of beer in modern industry is provided in Appendix 2 of OECD
(2004).
Despite the fact that many authors seem to be fascinated by the idea of Paleolithic people
hunting down alcoholic products, and often use highly suggestive and manipulative
writing styles to impose their ideas on their readers, direct archaeological evidence is
missing and even for later periods archaeobotanical evidence of malted grain is rare (Fig.
4), not least because malting results in comparatively higher fragility, thus reducing the
preservability of the grain. Malting requires the grain to be steeped in water for
germinating or sprouting, followed by drying or roasting it in a kiln, a process which in
prehistory may have accidentally led to carbonized grain; this is taken, then, as a record
of malting.
Page 11 of 26


Residues of oxalate are frequently cited as direct evidence for malt production from
cereal grain (e.g., Dietrich, Heun, Notroff, Schmidt, & Zarnkow, 2012). Oxalate is a
characteristic component of virtually all plant species, and occurs in comparatively low
amounts in beer (1.5 mg/100g), compared to other foodstuffs; it is frequently documented
in the form of archaeobotanical remains, such as the seeds of different cabbage species
(up to 7.5 mg/100g) or members of the Chenopodiaceae (up to several hundred mg/100g).
Oxalate is even present in meat in similar concentrations as in beer. Therefore, referring
to oxalate residues in prehistoric bowls or large-size vessels as evidence for prehistoric
beer production should be considered as highly tenuous. A more promising methodology
may be the analysis of enzymatically affected starch grains from archaeological samples
(Wang et al., 2017). Such methods, however, need to be integrated into multiple lines of
evidence to allow the differentiation of naturally sprouted grain from active fermentation
technologies.
Malted barley grain has been recorded at various archaeological sites in Europe and the
Aegean, dating to the Iron and Bronze Age (e.g., Stika, 2011; Valamoti, 2018). Besides
analytical attempts in archaeology to prove the production of beer, there are also works
that focus on the social implications of beer consumption in economics, status, and ritual
(e.g., Arthur, 2014; Dietler, 2006). In particular, the spiritual aspect of unification with the
gods and ancestors is to be mentioned here. Also as a tribute beer, malt and “beer bread”
is frequently mentioned in the texts (Meussdoerffer, 2009). Due to its generally short
period of storability compared to wine, according to Hayashida (2015), beer was never
extensively produced.
From the Bronze Age onwards, however, beer production represented, an important
sector of productivity in Mesopotamia (Sumerian kaš, Akkadian šikāru), Egypt, and other
geographic regions (cf. Tamang & Samuel, 2010; see also the excellent overview on the
textual evidence by Rattenborg, 2016). The considerable role of beer in the everyday diet
is documented in grain-to-beer ratios from studies of Middle Bronze Age textual records
in the Jazīrah and the Iraqi alluvium; these are 0.5 ~ 2 measures of grain to beer
throughout the Early and Middle Bronze Ages (Rattenborg, 2016). The records of the
Southern Levant present a strong contrast with these results. Although the evidence is
from the Iron Age period, using cereals for beer brewing purposes was likely less
important in the Levant, where wine production was much more relevant during classical
and historical times. This is also suggested by the ethnographic record on traditional
farming in the Southern Levant. In the seven volumes of Dalman’s (1987) ethnohistorical
description of agriculture in Palestine, beer production or consumption is almost
unmentioned, except one passage of the text in which beer is mentioned as a rare import
from Egypt for Passover. However, there are ancient Rabbinic texts describing the
production of beer (Lutz, 1922, p. 81), and in fact the assumption that viticultural
societies were ignorant of beer brewing may be a modern artifact. Certainly, recent
archaeobotanical research from Bronze Age Aegean settlements supports the argument
that the opposite may have been the case (Valamoti, 2018). Similar arguments have been
made based on the iconographic record from Linear B tablets (Weilhartner, 2016).
Page 12 of 26


Numerous studies of ancient beer brewing are available for the Near East, where
archaeobotanical and written evidence can be combined for conclusive results (Samuel,
1996). Ancient administrative and literary texts from Bronze Age Mesopotamia provide
some insight into the brewing process, and often include a list of components and
ingredients, which are not always clearly identifiable (Civil, 1964). Meussdoerffer (2009)
reports that the Sumerians produced at least nine different beer types from barley and
barley malt, and that in Babylonian times up to 70 different types of beer were
manufactured, mostly from emmer wheat and barley.
The most comprehensive knowledge on ancient beer brewing is available for Egypt,
including archaeobotanical, chemical, written, and iconographic evidence. The most
common component of ancient Egyptian beer residues is barley, whereas pure emmer
beer seems to have been rarely produced, but was more frequently mixed with barley
(Samuel, 2000). Lutz (1922) also notes that the commonest beer was prepared from
barley, as can be gathered from the following detailed description of the production
process (p. 78).
Take fine clean barley and moisten it for one day and draw it off or also lay it up in a
windless place until morning and again wet it six hours. Cast it into a smaller perforated
vessel and wet it and dry it until it shall become shredded and when this is so pat it (i.e.,
shake, or rub) in the sun-light until it falls apart. For the must(?) is bitter. Next grind it
and make it into loaves adding leaven, just like bread and cook it rather raw and
whenever (the loaves) rise, dissolve sweetened water and strain (it) through a strainer or
light sieve. Others in baking the loaves cast them into a vat(?) with water and they boil it
a little in order that it may not froth nor become luke-warm and they draw up (= absorb)
and strain it and having prepared it, heat (it) and examine (it).
As outlined in the ancient texts most Egyptian beers were composed of a special type of
bread or sourdough, which was used as a source of yeast and lactic acid (Meussdoerffer,
2009). This important step in historical beer production is also known as the preparation
of “beer loaves,” lightly baked dough based on barley groats, water, and yeast, which
were broken up afterward, washed through a sieve with water, and then left for
fermentation. The preparation of such “beer loaves” has likely contributed to scholarly
discussions on whether domestication developed due to the human needs of cereals for
bread or for beer production, as in the case of the most recently discovered remains, that
of 14,000-year-old cereal dough from the Jordanian site of Shubayaqa I (Arranz-Otaegui,
Gonzalez Carretero, Ramsey, Fuller, & Richter, 2018).
The most ancient chemically analyzed beer residues have been discovered in large
predynastic vats at Hierakonpolis (Upper Egypt) and provided radiocarbon dates of 3500–
3400 BC. The residues contained grains and spikelets of wheat and barley, as well as
fragments of dates and grape pips (Maksoud, El Hadidi, & Amer, 1994). A recent study of
six of the beer vats suggests that only emmer was used for beer brewing (Attia, Marinova,
Fahmy, & Baba, 2018).
Page 13 of 26


Despite the astonishing similarities between the methods used for beer brewing across
the different Near Eastern regions (Lutz, 1922), differences have been recognized as
existing over time (Meussdoerffer, 2009) and between geographic regions separated from
each other by a small distance (Ishida, 2005). Modern, high-quality barley for beer
production should have a low protein content, which means a high proportion of starch,
i.e. potentially fermentable sugar. Typical nitrogen levels of 1.5–1.7% have been reported
for two-rowed malting barley in Europe, and 1.8–2.0% for six-row barley in North
America (Bamforth & Barclay, 1993). Overviews on the general chemical composition of
barley are found in various contributions (e.g., MacGregor & Bhatty, 1993; Hockett,
1991).
Figure 4. Archaeobotanical records of malted grain

clearly showing the germ bud and impression. From
Jacomet (2009; drawings) and Stika (2011; photos).
The 1-mm scale applies to both drawings and photos.
Modern Economic Significance of Barley and Sustainable Agro-

Production
Although barley was traditionally a major food crop, it accounted for only 5% of
worldwide cereal production in 2018. There are, however, a number of countries
producing considerable amounts of barley. In terms of production volume Russia,
Germany, France, and Canada are the main producers, while in Ireland, Cyprus, Qatar,
Norway, and Denmark the barley production percentage per ton is still higher than 40%
among all cereals cultivated (FAOSTAT, 2018). The diversity of climates of the countries
focusing on barley cultivation documents the broad agro-ecological range of the crop. The
major proportion of the world’s barley production, i.e., approximately 85%, is used for
feeding animals (mostly pig and cattle), while the rest is used for malt and seed
production, and food consumption. A minor part is used to produce starch, either for food
use (major component of the daily diet in Morocco, India, China, and Ethiopia) or for the
chemical industry (OECD, 2004).
Aside from its general nutritional qualities, the crop provides minerals, vitamins,
especially vitamin E, and other antioxidants, primarily polyphenolics. The endosperm cell
walls are rich in β-glucans that positively affect blood cholesterol and glucose levels,
which can in turn impact cardiovascular health and diabetes control. For this reason,
barley has recently been rediscovered by Western civilizations for the purposes of food
production (Baik & Ullrich, 2008). Over the past 20 years, allergy, intolerance, and
sensitivity to cereals have been discussed to an increasing extent. About 1–3% of the
human population are documented as showing tendencies toward cereal intolerances,
Page 14 of 26


and increased use of gluten in food production industry and the consumption of wheat-
flour-based products have contributed to the increased prevalence of celiac disease,
which has been rising since at least the 1950s.The causal proteins, such as gliadins and
glutenins in wheat or hordein in barley, are together commonly termed “gluten.” Gluten
intolerance or celiac disease manifested as an autoimmune disease occurring in
individuals with the human leucocyte antigen (HLA) DQ2 and/or DQ8 genotype (Gilissen,
van der Meer, & Smulders, 2014), requiring the individual’s diet to be low in gluten input.
Reports on the lower gluten content of barley compared to wheat (Braun, 1995) may be
worth consideration in future food research.
Harry V. Harlan is a major figure in barley research and as a result of his impressive
research output is even considered by some as the “Godfather of modern barley.”3 His
research, which aimed at diversity and genetic improvement in barley cultivation, is
summarized in his autobiography (H. V. Harlan, 1957).
Beside the wild progenitor Hordeum vulgare ssp. spontaneum, only Hordeum bulbosum is
capable of producing fertile progeny when crossed to the domesticated species, and thus
is considered as a resource for the genetic improvement of barley (Backes et al., 2006), in
particular in relation with diseases affecting Hordeum vulgare, but not Hordeum
bulbosum (Morrell & Clegg, 2011; Pickering, 2000; Pickering, Niks, Johnston, & Butler,
2004). Apart from this, landraces of the domesticated Hordeum vulgare variety provide a
valuable gene pool for crop improvement and sustainable agriculture in the face of
climate change. Such landraces are particularly numerous in arid environments such as
those of North Africa, and are genetically very heterogeneous; i.e., no crossing barriers
have been developed between the wild and domesticated forms, and therefore
spontaneous and artificial crosses are easily obtained.
The breeding programs of the last 100 years have resulted in the creation of thousands of
commercial barley varieties, and meanwhile breeding has developed in a kind of race
against time, with the goal of shortening the breeding cycle and recombination; this is
made obvious by the increasing number of barley-breeding programs worldwide, which
are also carried out at scientific venues, such as the International Barley Genetics
Symposium.4 Quality criteria for breeding are determined according to the respective
uses (processing characteristics and nutritional value) of barley (OECD, 2004). Existing
varieties are distinguished by: their use as feed or malting barley, their winter or spring
growth habit, and whether they are two- or poly-row, or hulled vs. naked varieties, as well
as their starch amylose/amylopectin ratio. An example of breeding success in malting
barley is a significant reduction of seed dormancy. As long dormancy is undesirable in
malting, barley breeding has reduced it to 1–3 days, whereas in the poly-row varieties
which have been continuously bred for use as animal feed, dormancy has not undergone
strong selection and its dormancy period is similar to that in the wild form, i.e. variable
and occasionally very long (Von Bothmer, 1992).
Since 2006 the barley genome has been extremely well investigated by the members of
the International Barley Sequencing Consortium (IBSC),5 who aim to physically map and
Page 15 of 26


sequence the barley genome to accelerate crop improvement (The International Barley
Genome Sequencing Consortium, 2012). The haploid genome size of barley is, at 5.3 Gb,
one of the largest in cereal crops and is twice the size of the human genome. Meanwhile,
a number of traits in barley that enable it to withstand abiotic stress have been
genetically identified. Only to mention a few of these, they include resistance to water/
drought stress, seedling desiccation tolerance, yield and agronomic performance in the
Mediterranean environments under rain-fed conditions, salt tolerance, boron and
aluminum tolerance, manganese deficiency, and low-temperature and frost tolerance
(Lakew, 2009).
The composition and particular traits of the barley grain are submitted to the
environmental conditions predominating during its development, thus agronomic quality
of the barley grain is an important aspect in modern profit-oriented agriculture. For
example, dry weight per grain and final-grain starch content are increased with lower
temperatures, because the plant-internal mechanism of survival reduces the duration of
grain filling at elevated temperatures (Duffus & Cochrane, 1993). Fertilizer policy, and in
particular the twofold effects of nitrogen application with increasing yields on the one
hand, and reduced protein quality on the other, plays an enormous role in barley
agronomy. Increased nitrogen content in the grain is mainly due to the presence of
hordein (Duffus & Cochrane, 1993), more recently discussed in relation with nutritional
disorders such as gluten sensitivity and celiac disease. In general, nitrogen levels are a
critical issue in barley cultivation, because barley appears to be sensitive to nitrogen, not
only in relation to the protein content in grains, but also as a concern during the growing
period, as it is less resistant to lodging6 compared to wheat and oats, and thus replies
negatively to excessive fertilization (Hockett, 1991). The number of barley diseases is
considerable,7 while most of these occur under irrigated or high-moisture conditions.
Drought is a permanent constraint to agricultural production and an important factor

threatening the survival of people in the developing world. Furthermore, the impact of
climate and the aim of safeguarding future agricultural production has generated the
valuation of biodiversity in crops’ wild relatives and their preservation, providing a vital
resource for breeding new crop varieties that are better adapted to environmental stress.
Conservation initiatives are accessible through gramene.org (Morrell & Clegg, 2011).
Genetic mapping is highly valuable for the localization of economically important traits,
thus recent conservational works make heavy use of these technologies (e.g., Allel, Ben-
Amar, Lamine, & Abdelly, 2017). A particular focus in genetic research is on exposure to
salinity. As 40% of the world’s food is produced under irrigation, genetic research on
barley also aims to discover genes related to its salinity tolerance, which is a major
research area at the International Center for Biosaline Agriculture.8
Because of its broad range of environmental adaptability, barley is the most widely grown
cereal crop, with highly diverse breeding objectives; these include, beside its primary
economic properties, research into different forms of resistance to diseases, pests, and its
responses to various environmental stress factors (Hockett, 1991). An extensive number
of resistance genes for different diseases have been mapped in barley with DNA markers
Page 16 of 26


(Backes et al., 2006). These aspects, alongside its high valuation in food production, have
been crucial for the development of barley, and will guarantee its high value for human
societies in the future.
Suggested Reading
Baik, B.-K., & Ullrich, S. E. (2008). Barley for food: Characteristics, improvement,
and renewed interest.Journal of Cereal Science, 48(2), 233–242.
Gutaker, R. M., & Burbano, H. A. (2017). Reinforcing plant evolutionary genomics

using ancient DNA. Current Opinion in Plant Biology, 36, 38–45.
McGovern, P. E. (2009). Uncorking the past: The quest for wine, beer, and other alcoholic
beverages. Berkeley/London: University of California Press.
Morrell, P. L., & Clegg, M. T. (2011). Hordeum. In C. Kole (Ed.), Wild crop relatives:
Genomic and breeding resources—Cereals (pp. 309–319). Berlin: Springer.
OECD. (2004). Consensus document on compositional considerations for new varieties of

barley (Hordeum vulgare L.): Key food and feed nutrients and anti-nutrients. OECD.
Report no. 2.
Rattenborg, R. (2016). The scale and extent of political economies of the Middle
Bronze Age Jazirah and the Bilad al-Sam (c. 1800–1600 BCE). (Doctoral
Dissertation). Durham University, Durham.
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Notes:
(1.) Nomenclature according to Hanelt, Kilian, and Kilian (2001).
(2.) According to Hanelt, Kilian, and Kilian (2001), this is a hybrid offspring of ssp.
spontaneum and six-rowed barley.
(3.) See [Mark], “Harry Harlan: The godfather of modern barley,” Sprowtlabs (2017, 22
November).
(4.) See website for the symposium here: http://ibgs2016.org/.
(5.) “Mission statement,” International Barley Sequencing Consortium. (n.d.): http://

www.public.iastate.edu/~imagefpc/IBSC%20Webpage/IBSC%20Template-home.html
(6.) The displacement of stems or roots from their vertical and proper placement, which
leads to reduced yields.
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www.apsnet.org/publications/commonnames/Pages/Barley.aspx.
Page 25 of 26


(8.) “Home.” ICBA: Farming for tomorrow (n.d.). www.biosaline.org.
Simone Riehl
Environmental Archaeologist, Eberhard Karls Universität Tübingen
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Summary and Keywords

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Systematics and Taxonomy of Wild and

Barley species are

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Ecology and Biogeography of Barley

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Evolutionary History of Wild and Domesticated

Ancient DNA studies have contributed to an assessment of selection during different

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Barley in Human Subsistence (Pleistocene Through the Transition to

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agriculture, and probably determined their dominance in archaeobotanical assemblages

The evidence of crop-cultivating communities changes dramatically during the pre-

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The Evolution and Dispersal of Domesticated Barley Species From

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evolutionary processes in crop plants. The technology of analyzing ancient genomes in

As outlined in section 2, cultivated barley is adapted to a broader range of environments

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The Development of Barley Cultivation in the Near East and Europe

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The particular characteristics of barley can be considered as supportive for the

Dietary Use of Barley

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Figure 4. Archaeobotanical records of malted grain

Modern Economic Significance of Barley and Sustainable Agro-

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Drought is a permanent constraint to agricultural production and an important factor

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Gutaker, R. M., & Burbano, H. A. (2017). Reinforcing plant evolutionary genomics

OECD. (2004). Consensus document on compositional considerations for new varieties of

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Barton, L. (2016). The cultural context of biological adaptation to high elevation

Charles, M. P. (1984). Introductory remarks on the cereals. Bulletin on Sumerian

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Dietler, M. (2006). Alcohol: Anthropological/archaeological perspectives. Annual

Duffus, C. M., & Cochrane, M. P. (1993). Formation of the barley grain—Morphology,

FAOSTAT, (2018). Retrieved from http://www.fao.org/faostat/en/#data/QC.

Gutaker, R. M., & Burbano, H. A. (2017). Reinforcing plant evolutionary genomics