Fertile Crescent Research Articles

Speleothems from the Middle East: An Example of Water Limited Environments in the SISAL Database

The Middle East (ME) spans the transition between a temperate Mediterranean climate in the Levant to hyper-arid sub-tropical deserts in the southern part of the Arabian Peninsula (AP), with the complex alpine topography in the northeast feeding the Euphrates and Tigris rivers which support life in the Southeastern Fertile Crescent (FC). Climate projections predict severe drying in several parts of the ME in response to global warming, making it important to understand the controls of hydro-climate perturbations in the region. Here we discuss 23 ME speleothem stable oxygen isotope (δ18Occ) records from 16 sites from the SISAL_v1 database (Speleothem Isotope Synthesis and Analysis database), which provide a record of past hydro-climatic variability. Sub-millennial changes in ME δ18Occ values primarily indicate changes in past precipitation amounts the result of the main synoptic pattern in the region, specifically Mediterranean cyclones. This pattern is superimposed on change in vapor source δ18O composition. The coherency (or lack thereof) between regional records is reviewed from Pleistocene to present, covering the Last Glacial Maximum (~22 ka), prominent events during deglaciation, and the transition into the Holocene. The available δ18Occ time-series are investigated by binning and normalizing at 25-year and 200-year time windows over the Holocene. Important climatic oscillations in the Holocene are discussed, such as the 8.2 ka, 4.2 ka and 0.7 ka (the Little Ice Age) Before Present events. Common trends in the normalized anomalies are tested against different climate archives. Finally, recommendations for future speleothem-based research in the region are given along with comments on the utility and completeness of the SISAL database.

Between domestication and civilization: the role of agriculture and arboriculture in the emergence of the first urban societies

The transition to urbanism has long focused on annual staple crops (cereals and legumes), perhaps at the expense of understanding other changes within agricultural practices that occurred between the end of the initial domestication period and urbanisation. This paper examines the domestication and role of fruit tree crops within urbanisation in both Western Asia and China, using a combination of evidence for morphological change and a database that documents both the earliest occurrence of tree fruit crops and their spread beyond their wild range. In Western Asia the domestication of perennial fruit crops likely occurs between 6500 bc and 3500 bc, although it accompanies a shift in location from that of the earliest domestications within the Fertile Crescent to Mesopotamia, where the earliest urban societies arose. For China, fruit-tree domestication dates between ca 4000 and 2500 bc, commencing after millet domestication and rice domestication in Northern and Southern China, respectively, but within the period that led up to the urban societies that characterised the Longshan period in the Yellow River basin and the Liangzhu Culture in the Lower Yangtze. These results place the domestication of major fruit trees between the end of the domestication of staple annual crops and the rise of urbanism. On this basis it is argued that arboriculture played a fundamental role within the re-organisation of existing land use, shifting the emphasis from short-term returns of cereal crops into longer term investment in the developing agricultural landscape in both Western and East Asia. In this respect perennial tree crops can be placed alongside craft specialisation, such as metallurgy and textiles, in the formation of urban centres and the shaping the organisational administration that accompanied the rise of urbanism.

Megalithic tombs in western and northern Neolithic Europe were linked to a kindred society

Paleogenomic and archaeological studies show that Neolithic lifeways spread from the Fertile Crescent into Europe around 9000 BCE, reaching northwestern Europe by 4000 BCE. Starting around 4500 BCE, a new phenomenon of constructing megalithic monuments, particularly for funerary practices, emerged along the Atlantic façade. While it has been suggested that the emergence of megaliths was associated with the territories of farming communities, the origin and social structure of the groups that erected them has remained largely unknown. We generated genome sequence data from human remains, corresponding to 24 individuals from five megalithic burial sites, encompassing the widespread tradition of megalithic construction in northern and western Europe, and analyzed our results in relation to the existing European paleogenomic data. The various individuals buried in megaliths show genetic affinities with local farming groups within their different chronological contexts. Individuals buried in megaliths display (past) admixture with local hunter-gatherers, similar to that seen in other Neolithic individuals in Europe. In relation to the tomb populations, we find significantly more males than females buried in the megaliths of the British Isles. The genetic data show close kin relationships among the individuals buried within the megaliths, and for the Irish megaliths, we found a kin relation between individuals buried in different megaliths. We also see paternal continuity through time, including the same Y-chromosome haplotypes reoccurring. These observations suggest that the investigated funerary monuments were associated with patrilineal kindred groups. Our genomic investigation provides insight into the people associated with this long-standing megalith funerary tradition, including their social dynamics.

μCT trait analysis reveals morphometric differences between domesticated temperate small grain cereals and their wild relatives.

SummaryWheat and barley are two of the founder crops domesticated in the Fertile Crescent, and currently represent crops of major economic importance in temperate regions. Due to impacts on yield, quality and end‐use, grain morphometric traits remain an important goal for modern breeding programmes and are believed to have been selected for by human populations. To directly and accurately assess the three‐dimensional (3D) characteristics of grains, we combine X‐ray microcomputed tomography (μCT) imaging techniques with bespoke image analysis tools and mathematical modelling to investigate how grain size and shape vary across wild and domesticated wheat and barley. We find that grain depth and, to a lesser extent, width are major drivers of shape change and that these traits are still relatively plastic in modern bread wheat varieties. Significant changes in grain depth are also observed to be associated with differences in ploidy. Finally, we present a model that can accurately predict the wild or domesticated status of a grain from a given taxa based on the relationship between three morphometric parameters (length, width and depth) and suggest its general applicability to both archaeological identification studies and breeding programmes.

Wheat tandem kinases provide insights on disease-resistance gene flow and host-parasite co-evolution.

Stripe (yellow) rust, caused by the fungus Puccinia striiformis f. sp. tritici (Pst), is a destructive disease of wheat spread globally. Wild emmer wheat (Triticum turgidum ssp. dicoccoides; WEW) is known as a source for novel Pst resistance genes (R-gene), but our knowledge on wheat-Pst co-evolution in natural populations is limited. Yr15 is a WEW (accession G25) gene, which confers a broad-spectrum resistance to Pst, and encodes a tandem kinase-pseudokinase protein designated as WTK1. Exon-intron comparisons of multiple WTK1 homoeologous and paralogous copies scattered in allopolyploid wheat genomes enabled us to develop functional molecular markers (FMMs), which were used for population genetic study. The functional allele (Wtk1) was absent in a worldwide collection of 513 wheat cultivars, except for 32 introgression lines with Yr15 from G25, as well as in 84% of the 382 tested WEW accessions collected across the Fertile Crescent. Yr15 was found to be distributed along a narrow axis from Mt Carmel to the Anti-Lebanon Mountains ridge, mostly at elevations above c. 500m, where the climatic conditions are favorable for disease development, therefore providing insights on gene flow and host-parasite co-evolution in WEW natural habitats. Moreover, the worldwide absence of Wtk1 in cultivated wheat and in WEW natural populations from southeast Turkey, where wheat is believed to have been domesticated, proposes that Yr15 was rather left behind, than lost during domestication. Our results highlight the importance of conservation of WEW populations in their natural habitats for discovery of novel R-genes and studies of host-parasite co-evolution.

Cover Image: Barley and wheat biology

Wheat and barley were among the first crops in the Fertile Crescent. From there they started to successfully conquer our plates. Haas et al. (204–225) give an update to what we know about the domestication genomics of these two major cereals. Their highly complex genomes only recently became accessible for more detailed analyses through advanced sequencing techniques.

Allelic Diversity of Hordein-Coding Loci Hrd A and Hrd B in Cultivated (Hordeum vulgare L.) and Wild (Hordeum spontaneum C. Koch) Barley from Jordan (as a Part of the Fertile Crescent)

Allelic Diversity of Hordein-Coding Loci Hrd A and Hrd B in Cultivated (Hordeum vulgare L.) and Wild (Hordeum spontaneum C. Koch) Barley from Jordan (as a Part of the Fertile Crescent)

The ‘Wheat Puzzle’ and Kartvelians route to the Caucasus

Wheat (Triticum L.) originated in the Fertile Crescent approximately 10 kya BP and has since spread worldwide. The ‘Wheat Puzzle’ was termed the observation that wild predecessors of five Georgian endemic wheat subspecies are found in Fertile crescent quite far from the South Caucasus. Kartvelian peoples are the ethno-linguistic group of speakers of Kartvelian languages (Georgian, Megrel, Laz, Svan). Kartvelian language is one of the seven language families of Eurasiatic superfamily, that may have arisen from a common ancestor over 15 kya BP (Pagel et al. in Proc Natl Acad Sci USA 110:8471–8476, 2013). One of the possibility to explain ‘Wheat Puzzle’ is that speakers of Protokartvelian language could be separated from Protoeurasiatic language speakers after migration from Africa to the Arabian peninsula and later moved to the northern part of Mesopotamia where wheat was domesticated. Kartvelian speakers could migrate further to South Caucasus together with domesticated wheat subspecies.

Domestication and crop evolution of wheat and barley: Genes, genomics, and future directions

Wheat and barley are two of the founder crops of the agricultural revolution that took place 10,000 years ago in the Fertile Crescent and both crops remain among the world's most important crops. Domestication of these crops from their wild ancestors required the evolution of traits useful to humans, rather than survival in their natural environment. Of these traits, grain retention and threshability, yield improvement, changes to photoperiod sensitivity and nutritional value are most pronounced between wild and domesticated forms. Knowledge about the geographical origins of these crops and the genes responsible for domestication traits largely pre-dates the era of next-generation sequencing, although sequencing will lead to new insights. Molecular markers were initially used to calculate distance (relatedness), genetic diversity and to generate genetic maps which were useful in cloning major domestication genes. Both crops are characterized by large, complex genomes which were long thought to be beyond the scope of whole-genome sequencing. However, advances in sequencing technologies have improved the state of genomic resources for both wheat and barley. The availability of reference genomes for wheat and some of its progenitors, as well as for barley, sets the stage for answering unresolved questions in domestication genomics of wheat and barley.

20,000 years of societal vulnerability and adaptation to climate change in southwest Asia.

The Fertile Crescent, its hilly flanks and surrounding drylands has been a critical region for studying how climate has influenced societal change, and this review focuses on the region over the last 20,000 years. The complex social, economic, and environmental landscapes in the region today are not new phenomena and understanding their interactions requires a nuanced, multidisciplinary understanding of the past. This review builds on a history of collaboration between the social and natural palaeoscience disciplines. We provide a multidisciplinary, multiscalar perspective on the relevance of past climate, environmental, and archaeological research in assessing present day vulnerabilities and risks for the populations of southwest Asia. We discuss the complexity of palaeoclimatic data interpretation, particularly in relation to hydrology, and provide an overview of key time periods of palaeoclimatic interest. We discuss the critical role that vegetation plays in the human–climate–environment nexus and discuss the implications of the available palaeoclimate and archaeological data, and their interpretation, for palaeonarratives of the region, both climatically and socially. We also provide an overview of how modelling can improve our understanding of past climate impacts and associated change in risk to societies. We conclude by looking to future work, and identify themes of “scale” and “seasonality” as still requiring further focus. We suggest that by appreciating a given locale's place in the regional hydroscape, be it an archaeological site or palaeoenvironmental archive, more robust links to climate can be made where appropriate and interpretations drawn will demand the resolution of factors acting across multiple scales.This article is categorized under:Human Water > Water as Imagined and RepresentedScience of Water > Water and Environmental ChangeWater and Life > Nature of Freshwater Ecosystems

Variation in Stripe Rust Resistance and Morphological Traits in Wild Emmer Wheat Populations

Wild emmer wheat (Triticum dicoccoides), the tetraploid progenitor of cultivated wheats, is indigenous to the Near East Fertile Crescent. An important center of distribution is found today in and around the catchment area of the upper Jordan Valley in Israel and surrounding regions. In the current study, the field stripe rust resistance and morphological traits were analyzed using 98 sample accessions that represented the geographical distribution of wild emmer populations in Israel and its vicinity. The resistance tests at two field locations revealed that the majority of the wild emmer accessions possess quantitative resistance against stripe rust. This could be due to the high frequency of Yr36 in the wild emmer populations. The identification of potentially novel stripe rust resistance in this set of germplasm is highly significant. In total, 11 morphological traits were examined in this study. Wide range of natural variation was revealed in the tested morphological traits. Most of the morphological traits had significant correlations with climate variables, indicating that the local environmental conditions have a profound effect on shaping the genetic structure of wild emmer wheat. Our results suggest that wild emmer wheat has the enormous potential to improve stripe rust resistance and various important agronomical traits in wheat.

Conservation gap analysis of crop wild relatives in Turkey

AbstractCrop wild relatives (CWR) are wild plant taxa relatively closely related to crops that can contribute beneficial traits for crop improvement, such as biotic and abiotic stress resistance. Turkey has a rich flora of approximately 11,000 higher plant taxa, has two Vavilov centres of crop diversity (i.e. the Mediterranean and the Near East), is recognized as the cradle of agriculture providing the northern boundary of the Fertile Crescent, and has recently been identified as the country with the highest concentration of CWR diversity. The objective of this paper is to present the results of a gap analysis of CWR genetic diversity in Turkey using existing data sources of 458 of the 764 priority CWR taxa with available georeferenced data. In total, 27,597 presence points were obtained from the Global Biodiversity Information Facility, Aegean Agricultural Research Institute in Menemen and Field Crops Central Research Institute, Ankara. Geographic Information System (GIS) software was used to identify taxon richness, sampling bias, future ex situ population collection and location where existing protected sites could form the basis of national network of in situ genetic reserves. CWR taxon richness was located along the Aegean Coast, Syrian border and southern Mediterranean coast. Current ex situ representation of CWR taxa is inadequate and further collection across the entire country is required. The highest priority in situ reserve location is found in Izmir, Sanliurfa and Antalya province, which reflects overall CWR richness.

Ancient cattle genomics, origins, and rapid turnover in the Fertile Crescent

Ancient cattle genomics, origins, and rapid turnover in the Fertile Crescent

PHARMACOGENOMIC ARCHITECTURE OF CLOZAPINE PLASMA CONCENTRATIONS

As the climate became warmer in the mid-latitudes in the wake of the conclusion of the last ice age, humankind entered a civilized society based on farming and herding. The first civilization grew as farming began in the Fertile Crescent located in the east of the Mediterranean Sea. Then, civilization spread mainly to the regions in similar latitudes where the climatic conditions were relatively comparable, thereby prompting the development of civilizations along the east–west axis. The supply of nutrients to humanity greatly changed compared with those supplied in the age of the hunter-gatherers, exerting a considerable influence on the disease pattern. In addition, animal domestication provided a hotbed for various infectious diseases.

Cognitive links between mountains and lowlands: balance weights from the Early Bronze Age site of Shengavit (Armenia) in context

The timespan from the mid 4th to the mid 3rd millennium BC is characterised by essential social transformations in the Near East and surrounding regions. The growing complexity is reflected, among others, also in technological innovations and active trade relations, which led to the appearance of balance weights and corresponding weight systems. This contribution, based on evidence from the settlement of Shengavit, Armenia, argues that balance weights were in use also beyond Mesopotamia, within the area of the spreading of Kura-Araxes cultural community, from the southern Caucasus to the Fertile Crescent. The use of such weight systems at Kura-Araxes sites implies possible cognitive links to the Near Eastern centres, reflecting the transfer of knowledge between mountains and lowlands.

The Eurasian Steppe is an important goat propagation route: A phylogeographic analysis using mitochondrial DNA and Y-chromosome sequences of Kazakhstani goats.

Goats (Capra hircus) were domesticated in the Fertile Crescent and propagated all over the world. The Silk Road through the Eurasian Steppe belt is a possible propagation route for domestic goats to Central Asia. Kazakhstan is in close geographical proximity to domestication centers and covers the majority of the Eurasian Steppe belt. In this study, we examined the genetic diversity and phylogeographic structure of Kazakhstani goats. The mtDNA sequences of 141 Kazakhstani goats were categorized into haplogroups A, C, and D, of which haplogroup A was predominant (97%), whereas haplogroups C and D were detected at low frequencies (1.4% each). The Kazakhstani haplotypes C were thzen categorized into Asian mtDNA type. Sequence analysis of the SRY gene on the Y-chromosome in 67 male Kazakhstani goats revealed two haplotypes: Y1A (64%) and Y2A (36%). Analysis of the distribution of mtDNA haplogroups and SRY haplotypes from Eurasia and Africa demonstrated genetic similarity among animals from Kazakhstan, Mongolia, and Northwest China located on the Eurasian Steppe belt. These phylogeographic results suggested that the Eurasian Steppe belt was an important propagation route for goats to Central Asia.

Repetitive DNA in the Architecture, Repatterning, and Diversification of the Genome of Aegilops speltoides Tausch (Poaceae, Triticeae).

The genome’s adaptability to environmental changes, especially during rapid climatic fluctuations, underlies the existence and evolution of species. In the wild, genetic and epigenetic genomic changes are accompanied by significant alterations in the complex nuclear repetitive DNA fraction. Current intraspecific polymorphism of repetitive DNA is closely related to ongoing chromosomal rearrangements, which typically result from erroneous DNA repair and recombination. In this study, we addressed tandem repeat patterns and interaction/reshuffling both in pollen mother cell (PMC) development and somatogenesis in the wild diploid cereal Aegilops speltoides, with a focus on genome repatterning and stabilization. Individual contrasting genotypes were investigated using the fluorescent in situ hybridization (FISH) approach by applying correlative fluorescence and electron microscopy. Species-specific Spelt1 and tribe-specific Spelt52 tandem repeats were used as the markers for monitoring somatic and meiotic chromosomal interactions and dynamics in somatic interphase nuclei. We found that, the number of tandem repeat clusters in nuclei is usually lower than the number on chromosomes due to the associations of clusters of the same type in common blocks. In addition, tightly associated Spelt1–Spelt52 clusters were revealed in different genotypes. The frequencies of nonhomologous/ectopic associations between tandem repeat clusters were revealed in a genotype-/population-specific manner. An increase in the number of tandem repeat clusters in the genome causes an increase in the frequencies of their associations. The distal/terminal regions of homologous chromosomes are separated in nuclear space, and nonhomologous chromosomes are often involved in somatic recombination as seen by frequently formed interchromosomal chromatin bridges. In both microgametogenesis and somatogenesis, inter- and intrachromosomal associations are likely to lead to DNA breaks during chromosome disjunction in the anaphase stage. Uncondensed/improperly packed DNA fibers, mainly in heterochromatic regions, were revealed in both the meiotic and somatic prophases that might be a result of broken associations. Altogether, the data obtained showed that intraorganismal dynamics of repetitive DNA under the conditions of natural out-crossing and artificial intraspecific hybridization mirrors the structural plasticity of the Ae. speltoides genome, which is interlinked with genetic diversity through the species distribution area in contrasting ecogeographical environments in and around the Fertile Crescent.

Genetic dissection of quantitative and qualitative traits using a minimum set of barley Recombinant Chromosome Substitution Lines

BackgroundExploring the natural occurring genetic variation of the wild barley genepool has become a major target of barley crop breeding programmes aiming to increase crop productivity and sustainability in global climate change scenarios. However this diversity remains unexploited and effective approaches are required to investigate the benefits that unadapted genomes could bring to crop improved resilience. In the present study, a set of Recombinant Chromosome Substitution Lines (RCSLs) derived from an elite barley cultivar ‘Harrington’ as the recurrent parent, and a wild barley accession from the Fertile Crescent ‘Caesarea 26–24’, as the donor parent (Matus et al. Genome 46:1010–23, 2003) have been utilised in field and controlled conditions to examine the contribution of wild barley genome as a source of novel allelic variation for the cultivated barley genepool.MethodsTwenty-eight RCSLs which were selected to represent the entire genome of the wild barley accession, were genotyped using the 9 K iSelect SNP markers (Comadran et al. Nat Genet 44:1388–92, 2012) and phenotyped for a range of morphological, developmental and agronomic traits in 2 years using a rain-out shelter with four replicates and three water treatments. Data were analysed for marker traits associations using a mixed model approach.ResultsWe identified lines that differ significantly from the elite parent for both qualitative and quantitative traits across growing seasons and water regimes. The detailed genotypic characterisation of the lines for over 1800 polymorphic SNP markers and the design of a mixed model analysis identified chromosomal regions associated with yield related traits where the wild barley allele had a positive response increasing grain weight and size. In addition, variation for qualitative characters, such as the presence of cuticle waxes on the developing spikes, was associated with the wild barley introgressions. Despite the coarse location of the QTLs, interesting candidate genes for the major marker-trait associations were identified using the recently released barley genome assembly.ConclusionThis study has highlighted the role of exotic germplasm to contribute novel allelic variation by using an optimised experimental approach focused on an exotic genetic library. The results obtained constitute a step forward to the development of more tolerant and resilient varieties.

‘The king, my lord, wrote me in a tablet: “…” ’ (EA 149:54 – 55): direct citation and communication in the syro-canaanite el-amarna letters

This paper examines the use of direct citation in the correspondence between the Egyptian Pharaoh and his officials and his Syro-Canaan vassals in the El-Amarna archive. A frequent phenomenon in the modern and ancient world alike, direct citation serves to support the writer’s claims, lay the blame on or absolve others, demonstrate loyalty, and depict distress. The examples adduced fall into four categories: authoritative, third-party, confirmative, and fabricated/ imaginary speech. Raising the question of the validity of applying modern linguistic theories to ancient sources, these types are analysed in the light of the international diplomacy that flourished in the Fertile Crescent during the 14th-century BCE.

Food resources in the economy and ritual practices of the Northern Mesopotamia population during the transition to the Neolithic

The ultimate establishment of the agricultural economy in the central zone of the Fertile Crescent took place in the late Pre-Pottery Neolithic (PPNB), while the heyday of symbolism, establishing complex social relations among the population of Northern Mesopotamia occur in the era of the early Pre-Pottery Neolithic (PPNA). In this period, the domestication of any plant species is not yet registered in South-Eastern Anatolia (an area where the long-term intertribal cult center of Gbekli Tepe was found) unlike the neighboring Levant. The paper discusses possible models for the producing economy establishment in the region, analyzes materials suggesting that the ritual practices of the transition period to the Neolithic in some cases could contribute to the emergence and new economic strategies spread on the territory of Northern Mesopotamia. At the same time, the comparison of the climatic changes scientific studies results, archeobotanical and archaeozoological collections and material evidence of the development of social and spiritual life from Epipaleolithic and early Neolithic monuments of Northern Mesopotamia shows the coevolution/mutual influence of people and the surrounding natural environment. In our opinion, on the basis of the available data it is impossible to assert the primacy of the symbol revolution in the process of Neolithization in relation to early attempts at plant cultivation.