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The Domestication of Barley

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The Domestication of Barley Barley ( P h o t o : R o b i n A p p e l . ) Figure 1: Cultivated barley. The domestication of barley The origin of agriculture is one “How great is the power of man in Wild barley of the seminal events in human accumulating by his Selection The wild ancestor of barley is well culture. The development of documented, and has been traditionally domestic animals and crops successive slight variations...” described as Hordeum spontaneum C. Koch ., a wild, large-seeded barley. Originally from wild species laid the HARLES ARWIN N THE RIGIN OF PECIES C D , O O S , 1859 considered a separate species, this plant is, in foundation for the Neolithic fact, a subspecies of the cultivated crop, revolution some 10,000 –12,000 cultivation of plants selected from the wild because the wild type is cross-compatible and years ago and resulted in the (but not yet genetically different from wild fully interfertile with cultivated barley. Since transition of hunter-gatherer plants) and terminates in fixation, through there are no biological barriers to groups to sedentary, pastoral human selection, of morphological (hence hybridisation between wild and cultivated and farming societies. genetic) differences distinguishing a barley, all forms are now deemed to belong to domesticate from its wild progenitor. These a single biological species ( H. vulgare ) and, differences make up the ‘domestication as a result of gene-flow, any rigorous by Ian Hornsey syndrome’ and usually render the domesticate taxonomic distinction between wild and less capable of surviving in the wild (i.e. cultivated forms is always going to be become dependent on man). As well as loss of problematical. Accordingly, the scientifically griculture in the Old World appears to seed dispersal mechanism, other features of correct designation for the wild type is A have arisen in several key centres, the syndrome include, increase in seed size, therefore H. vulgare L. subp. spontaneum including the Fertile Crescent in the Near East loss of seed dormancy and loss of chemical or (C.Koch) Thell. (although, henceforth, we’ll and the middleYangtze River Valley in China. mechanical protection against herbivores. call it H. spontaneum ). Then, strictly, There has long been an interest in the Cultivated barley ( Hordeum vulgare L. – cultivated barley becomes H. vulgare subsp. mechanisms by which agriculture was Figure 1), is a diploid (2n = 14) and mostly vulgare. disseminated from these sites of origin. Only self-pollinated crop of which hundreds of Both wild barley and the first cultivated by studying the origins and migrations of modern varieties and land races are known. forms were morphologically similar, and were cultivated crops such as barley are we likely Although mainly grown today for animal feed two-rowed and ‘hulled’ (i.e. the husk adhered to be able to ascertain the origins of brewing. and for brewing/distilling, barley was an to the grain). The husk is rich in silica and Since the onset of the genomic age, studies of important cereal food in the past. Any doubt would have to be removed by pounding barley domestication have progressed rapidly. about the use of barley in bread-making can before the grain is suitable for consumption. During this agricultural revolution, ancient be dispelled by referring to the Bible, where In ‘naked’ barleys, the husk does not adhere to people saved seeds from plants with favoured Ezekiel (4:9) says: “Take thou also unto thee the grain and can be easily removed by traits for the next generation (sowing), and wheat, and barley, and beans, and lentils, and threshing. over time they converted seemingly millet, and spelt, and put them in one vessel H. spontaneum (Figure 2) can still be found unpromising wild species into reliable, and make thee bread thereof...” growing today around the eastern productive crops. Domestication is the end- It is probably only since the Middle Ages that Mediterranean basin and western Asia (Figure point of a continuum that starts with the the role of barley as a staple food has declined 3), where it occupies primary habitats. exploitation of wild plants, continues through in most areas. Morphologically, it is rather difficult to Brewer & Distiller International • December 2010 • www.ibd.org.uk 41 Barley ( ( P I m h o a t g o e : : D O r s x f M o r a d r k U N n i e v s e b r i s t t i t a y n P d r D e s e s l . w ) e n S a m u e l . ) Figure 3. Distribution of wild barley. The area in which wild barley is massively spread is shaded – the ‘Fertile Crescent’. Dots represent additional sites, mainly weedy forms. Wild barley extends further eastwards – as far as Tibet. wild populations. It may, however, quickly be now submerged Late Stone Age, site on the selected for in a population that is subjected to southern shore of the Sea of Galilee, and the man-made environment of reaping, dating from ca. 17,000 BC. Other early threshing and sowing. Experimental field evidence of H.spontaneum collection from trials and computer simulations indicate that, the wild comes from another Late Stone Age under certain conditions, the recessive (non- site at Tell Abu Hureyra, in northern Syria brittle) genotype may become predominant in (9000 BC) ; Tell Mureybit (Late Stone Age, 8- a remarkably short time (around 20 years!). 7500 BC.), also in northern Syria, and Tell Aswad, some 25km. south-east of Damascus, Barley domestication at a site dated to 7800 – 7600 BC. The kernel The received wisdom has been that barley specimens recovered at these sites were domestication was a single event essentially identical to present-day wild (monophyletic) and that the domesticate barley, being two-rowed with a brittle rachis. Figure 2. Hordeum spontaneum – growing on spread with human migrations, but the small The earliest signs of the cultivation of barley the Plain of Konya, Turkey. differences between wild and domesticated emanate from a later (Neolithic; ca. 7500 BC barley and the apparent ease with which wild onward) level at Tell Abu Hureyra; from forms become domesticated, has led some phase II (ca. 6900-6600 BC) in Tell Aswad distinguish between wild H. spontaneum and workers to suggest that barley domestication and from 6400 BC pre-pottery Jarmo, Iraq. At cultivated two-rowed varieties, since they must have occurred more than once (i.e. was all of these sites cultivated, non-brittle, barley differ primarily in their methods of seed polyphyletic). Such agricultural historians heads have been found, together with the wild structure (Figure 4) and dispersal; H. and geneticists alike think that the crop was type. The earliest domesticated barley to be spontaneum has brittle ears (spikes) which, at domesticated in dispersed small areas and recovered (ca. 7500 – 6,400 BC) also had maturity, disarticulate into individual wedge- spread by human migrations and trade. two-rowed spikes. shaped spikelets. These fragments are highly Most archaeological evidence suggests that Cultivation of six-rowed barley was specialised devices which ensure the survival barley domestication went hand in hand with apparently a little later, with estimated dates of the plant in the wild. In wild barley, with the evolution of Neolithic villages in the being in the 6,800-6,000 BC range. Then, rudimentary lateral rows of spikelet, there is Levantine arc of the ‘Fertile Crescent’. around 5,000-4,000 BC, when barley became no impediment to seed dispersal after Late Stone Age peoples in this area became widely cultivated in the alluvial soils of shattering of the spike. increasingly dependent on plant foods for Mesopotamia (and, later, in Lower Egypt), the Wild barley has complementary genes, Btr1 their existence and may have cultivated six-rowed form soon became dominant and and Btr2, which control the formation of the plants. The earliest signs of the pre- ousted its two-rowed sister to become the brittle rachis and cultivated barley carries agricultural gathering of wild barley (together most important crop in these early mutant recessive alleles (btr1 and btr2) which with wild emmer wheat) are from Ohalo II, a civilizations. are tightly linked and result in a non-brittle About fifty years ago, Danish botanist Hans rachis. All cultivated barleys have either one Helbaek noted that evidence strongly of the two recessive alleles and there is suggested that two-rowed barley was brought evidence that most occidental cultivars have into cultivation earlier than its six-rowed the brt1 allele while most oriental forms carry counterpart, but that there were spatial btr2. In wild forms, seeds are regularly differences in their spread. Both two-and six- scattered on the ground as the stalk shatters – rowed barleys emerged in Greece between this makes them difficult for humans to 6,000 and 4,000 BC, but the six-rowed form gather. After the lethal (for wild populations) dominated in the Balkans and central Europe mutation, the seeds fail to drop and are (3,000 BC – to around the birth of Christ) and conveniently concentrated on the stalk for in southern Europe and North Africa humans to gather. (5,000 –2,000 BC) and he also noted that after The loss of a brittle rachis is one of the its disappearance from Mesopotamia and most critical traits in the evolution and ancient Egypt it does not reappear in the domestication of barley and the mutation Figure 4: Cross-sections of wild and archaeological record of these two regions mentioned above will rarely occur in totally cultivated barley. until about AD 900. Cultivation of two-rowed 42 Brewer & Distiller International • December 2010 • www.ibd.org.uk Barley ( P h o t o : D r P a m e l a R o s e , E E S .
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