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Site of Einkorn Wheat Domestication Identified by DNA Fingerprinting
Abstract
The emergence of agriculture in the Near East also involved the domestication of einkorn wheat. Phylogenetic analysis that
was based on the allelic frequency at 288 amplified fragment length polymorphism molecular marker loci indicates that a wild
group of Triticum monococcum boeoticum lines from the Karacadağ mountains (southeast Turkey) is the likely progenitor of cultivated einkorn varieties. Evidence
from archeological excavations of early agricultural settlements nearby supports the conclusion that domestication of einkorn
wheat began near the Karacadağmountains.
... This region does not overlap with but is adjacent to the presumed core domestication area proposed for some of the founder crops of the Southwest Asian domestication center, that is, the Fertile Crescent (Abbo et al. 2010). The core area, located in the KaracaDag˘range near Diyarbakir in southern Turkey, would have been the domestication hearth of einkorn (Heun et al. 1997) and emmer (Ö zkan et al. 2002;Luo et al. 2007) wheat, chickpea (Abbo et al. 2003), and lentil (Ladizinsky 1999). How can this discrepancy be explained? ...
... The main arguments that have been advanced in favor of this location include genetic relatedness based on neutral markers, the current distribution of the wild progenitor of some of the 7 so-called founder crops, and the existence of archaeological sites containing archaeobotanical remains in the same general area. Heun et al. (1997) first identified a closer genetic relationship of the Triticum monococcum var. boeoticum population from the KaracaDag˘region with T. monococcum var. ...
... However, these same results also show that the KaracaDag˘population is an outlier among T. monococcum var. boeticum populations (see Figure 2A of Heun et al. 1997) and is genetically intermediate between T. monococcum var. boeticum and var. ...
... They bolstered their argument with recent genetic analyses of modern populations of some of these domesticates and their proposed progenitors-a technique that was new to archaeology at the time. In one of the first multi-locus genetic analyses of Near Eastern founder crop, Heun et al. (1997) concluded that domestic einkorn had a monophyletic origin that could be traced to a single wild population growing on the slopes of the Karacadağ volcano in southeastern Turkey. Subsequent genetic analysis of domestic emmer concluded that this crop plant also had a monophyletic origin with its closest living wild population found on the same Karacadağ volcano (Özkan et al. 2002). ...
... The 1990s and early 2000s also witnessed remarkable advances in the development of scientific methods directed at documenting domestication. Early genetic analyses of modern domesticates and their wild relatives were helping identify both the progenitors of future domesticates and the likely location of initial domestication of various plant (Heun et al. 1997;Özkan et al. 2002) and animal species (Bradley et al. 1996;Luikart et al. 2001). The promise of being able to extract replicable DNA from ancient specimens was on the horizon (Fernández et al. 2005(Fernández et al. , 2006. ...
Interdisciplinary teams investigating the origins of agriculture in the Eastern Fertile Crescent in the 1950s through 1970s considered the region a primary center of initial domestication and agricultural emergence. Political events then shifted the focus of archaeological investigation on agricultural origins to the Western Fertile Crescent. Decades of subsequent research appeared to indicate that the west was the earliest and most important center of agricultural origins in Southwest Asia, with the Eastern Fertile Crescent portrayed as a backwater that lagged behind transformative innovations from the west. The resumption of investigations in the east in the early 2000s, coupled with new scientific methods for documenting agricultural emergence, has reestablished the region as a heartland of domestication of both crop and livestock species. This broad topic is covered in two papers, beginning here with the history of this work from the 1950s through the early 2000s. The second paper will present a synthesis of recent work in the east, evaluating the continued relevance of early work in light of recent explanatory models for agricultural origins.
... Wheat is one of the most important cereal crops consumed as a staple food by humans on almost every continent in the world. It was the first domesticated crop about 10,000-12,000 years ago around Fertile Crescent and played an important role in initiating the agricultural revolution [1][2][3][4]. The wild form of durum wheat, wild emmer (Triticum turgidum ssp. ...
... Each season was accepted as a different environment and named as follows: E1 (2017-2018) and E2 (2018-2019). 3 The panel was sown in the plots, which had two 2-m rows with a row spacing of 20 cm and a genotype spacing of 10 cm. In each row, there were 20 seeds. ...
Durum wheat is an economically and nutritionally important cereal. The increase in durum wheat yield is mostly associated with improving grain traits. The grain size and shape-related traits are directly related to wheat yield. In addition, grain size influences the seed germination rate and seedling vigour, which play key roles in stand establishment and yield. Thus, it is important to investigate grain traits both agro-morphologically and genetically. In this study, a panel of durum wheat, consisted of 146 genotypes, was evaluated for grain traits agro-morphologically, and a genome-wide association study (GWAS) was conducted to dissect the genomic regions associated with these traits. As a result of GWAS, a total of 41 marker-trait associations (MTAs) were identified on different chromosomes of durum wheat. Of these MTAs, only 11 were stable across environments. A BLAST search for flanking sequences of every stable MTAs in the Svevo genome identified 18 putative candidate genes directly associated with seed traits of different plants, particularly wheat seeds. In conclusion, the annotation results and literature information provide strong evidence that identified stable MTAs and their candidate genes may have important functions in the formation of wheat grain traits. After the validation of these MTAs with different fine-mapping and functional characterization studies, these loci may provide valuable information for geneticists and breeders to improve wheat yield.
... monococcum (2n = 2x = 14, A m A m genome), is a model plant for wheat and other Triticeae species. T. monococcum is one of the first domesticated and oldest cultivated crops, with a history dating back about 12,000 years 8,9 . Being an agricultural founder crop with a long history of cultivation in various geographical and environmental regions, einkorn wheat is an important source of genes for improving modern wheat for resistance against biotic and abiotic stresses [10][11][12][13][14][15][16][17][18][19] . ...
... Einkorn wheat also offers a unique opportunity to study domestication and selection-related evolutionary history as it is the only diploid wheat species for which there are both wild and cultivated types 9,17,31 . Because of the contrasting characters of wild and cultivated einkorn 32 , mapping populations have been developed from the crosses between wild and cultivated einkorn accessions to understand the genetic architecture of domestication and agronomic traits in einkorn wheat 11,[33][34][35] . ...
Einkorn wheat (Triticum monococcum) is an ancient grain crop and a close relative of the diploid progenitor (T. urartu) of polyploid wheat. It is the only diploid wheat species having both domesticated and wild forms and therefore provides an excellent system to identify domestication genes and genes for traits of interest to utilize in wheat improvement. Here, we leverage genomic advancements for einkorn wheat using an einkorn reference genome assembly combined with skim-sequencing of a large genetic population of 812 recombinant inbred lines (RILs) developed from a cross between a wild and a domesticated T. monococcum accession. We identify 15,919 crossover breakpoints delimited to a median and average interval of 114 Kbp and 219 Kbp, respectively. This high-resolution mapping resource enables us to perform fine-scale mapping of one qualitative (red coleoptile) and one quantitative (spikelet number per spike) trait, resulting in the identification of small physical intervals (400 Kb to 700 Kb) with a limited number of candidate genes. Furthermore, an important domestication locus for brittle rachis is also identified on chromosome 7A. This resource presents an exciting route to perform trait discovery in diploid wheat for agronomically important traits and their further deployment in einkorn as well as tetraploid pasta wheat and hexaploid bread wheat cultivars.
... Using SNP tags located in the TtBtr1-A and TtBtr1-B "domestication loci" (3049 SNPs for A1 subgenome and 4724 SNPs for B1 subgenome), local phylogenies were constructed, which proved that wild emmer from the northern Levant was the closest group to all cultivated wheat [15]. In southeastern Turkey, direct wild ancestors of another wheat important in antiquity, T. monococcum, were also discovered [27], which led to the approval of the hypothesis on the domestication of T. dicoccum in this region. ...
... It consists of 620 genera and 8000 species. Plants are annuals, leaf-blades are flattened, inflorescence may have a spike, spikelets solitary at the nodes of the tough or fragile rachis, laterally compressed, 2-6(-9)-flowered (Cope, 1982;Heun et al., 1997). It is widely distributed in all the regions of the world (Anjum and Muhammad, 2012). ...
Wheat (Triticum aestivum L.) is one of the first domesticated crop plants and is the most widely grown crop in the world, both in terms of cultivated area and yield, and wheat is known to be consumed by about two-thirds of the world's population. The study was carried out to investigate the elemental and proximate composition of wheat against boron induced stress. Greenhouse experiments were conducted in October 2019 under natural light conditions to assess the chemical status of plants under various treatments such as the stress of inorganic boric acid. In the experiment, the pots were filled with sandy loam soil (1 kg). The pots were pre-treated with boron as boric acid (H 3 BO 3) only once before sowing, at the doses of 0 mg/kg (control), 3 mg/kg, 6 mg/kg, 9 mg/kg, 12 mg/ kg, 15 mg/kg and 18 mg/kg soil. The different concentrations of Boron (B) were prepared separately by taking a respective amount of boric acid. Pots without supplemented B constituted the control. No additional supplements were applied to the experimental soil. Each pot was irrigated with 100 mL of distilled water with intermittent intervals of 48 hours. All the treatments were replicated three times considering each pot as one replicate. Through EDX analysis, it was noticed that the boron stress treatment exhibited the maximum number of elements detected in (T 2 , T 5 , and T 6) wheat crops followed by (T 0 , T 1 , and T 3). A total of 17 elements were detected and observed in wheat including Ni in different treatments of B. It has been determined that the amount of each element varies according to the amount of B applied. Thus, the study performed under B stress helps understand elemental and proximate content in various treatments in wheat plants.. 2024. Proximate composition and minerals content of wheat (Triticum aestivum L.) under boric acid stress via energy-dispersive x-ray (EDX) fluorescence based analysis.
... Archaeobotanical studies from the Fertile Crescent (i.e., Southeast Anatolia) are among the regions where the first domestication attempts began worldwide and early farming sites were established (Lev-Yadun et al. 2000;Aurenche et al. 2001;Diamond 2002;Brown et al. 2009). Domestication of wild grains such as barley, einkorn wheat, emmer wheat, and rye during the Neolithic period is a challenging process that began with the showing of wild seeds and was identified with the agricultural benefits of some mutations (Nesbitt and Samuel 1996;Heun et al. 1997Heun et al. , 2008Lev-Yadun et al. 2000;Özkan et al. 2002;Luo et al. 2007;Dietrich et al. 2012;Weiss and Zohary 2011;Abbo and Gopher 2017). The earliest signs of cereal processing in the Fertile Crescent are thought to have occurred in the Levant Basin in the Late Epipaleolithic (Hillman et al. 2001;Liu et al. 2018;Eitam et al. 2015). ...
Paleoecological findings from soil samples recovered near the Göbekli Tepe indicate an environment that sustains a variety of multiproxy analyses, revealing prominent changes in ecological conditions.
Soil samples were collected in the vicinity of the pre-Neolithic site in the Göbekli Tepe. These sediment samples underwent comprehensive analysis utilizing a multiproxy approach, which included pollen analysis, total organic carbon (TOC) and total inorganic carbon (TOC) measurements, and X-ray fluorescence (XRF) elemental analysis.
Total organic carbon varied significantly in the SK-1 section, with lower levels at the bottom and higher levels at the top. The Ca/Ti ratio, which is indicative of weathering and moisture availability, typically increased in the early Holocene and briefly decreased at approximately 13,354 years BP. The Sr/Ti ratio, related to weathering and erosion processes, also suggests intensified weathering. The presence of deciduous forests and the increase in organic carbon values further support the notion of favorable environmental conditions during the early Holocene. In contrast, the lower levels of SK-1 show lower organic carbon values, lower Ca/Ti and Sr/Ti ratios, and greater abundances of herbaceous plants and steppes, indicating colder and drier conditions at 13,354 years BP.
A significant increase in organic carbon values and the presence of deciduous Quercus in the upper levels of the SK-3 section suggest a period of increased organic productivity and the existence of deciduous forests during the early Holocene at the Göbekli Tepe.
... For more than 10,000 years, wheat cultivation has held a significant place within the socio-economic fabric of the "Fertile Crescent"-an area within the Eastern Mediterranean and West Central Asia (Bilgic et al. 2016;Diamond 1997Diamond , 2002Heun et al. 1997). However, the region, which recently surpassed the critical 1.5 °C temperature increase threshold compared to pre-industrial levels, now faces significant climatic challenges due to the accelerating pace of climate change (IPCC 2022). ...
Numerous studies have addressed climate change's impact on agriculture, yet the specific effects on wheat production in Turkey remain under-explored. This research aims to bridge this gap by quantifying these impacts in Turkey over 25 years. As the region's principal wheat producer, Turkey offers a unique context, situated in the area where wheat was first domesticated. We analyzed a comprehensive dataset comprising over 20,000 records from 820 political districts, employing a panel data fixed-effect model to mitigate unobserved heterogeneity and reveal dynamic time-related patterns. We focused on critical climate variables: temperature, precipitation, and drought. The results show a significant 9% decrease in wheat yields following a 1.5 °C rise in temperature, affecting nearly 1 million hectares. A notable finding is the 'May effect', highlighting how climatic changes in May disproportionately affect wheat yields. This study provides detailed insights into the temporal and geographic aspects of climate change's impact on wheat production, emphasizing the need for targeted policy actions and strategic agricultural planning, particularly in dry areas.
... Найбільш ранніми одомашненими рослинами, так званими засновниками ("founder crops"), були плівчасті пшениці двозернянка і однозернянка та плівчастий ячмінь разом з бобовими -сочевицею, горохом, викою ервілією, нутом і також льоном (Zohary, 1996). За даними порівняльної генетики, імовірним районом походження культурної пшениці з дикої є околиці сучасного міста Діярбакир у Туреччині (Heun et al., 1997). Дикі предки культурних рослин ростуть і зараз в південно-західних районах Азії, в зоні так званого "Fertile Crescent" (Плодючого півмісяця). ...
Archaeobotany (palaeoethnobotany) is a science that studies fossil plant remains and, as a result of their analysis, establishes the time and place of origin of cultivated plants and their further distribution. Obtaining information is closely related to archaeological research. The results of archaeobotanical research are of interest to historians, botanists, ethnographers, anthropologists, agronomists, chemists, and linguists. Since the identification of random fossils, archaeobotany has come a long way in a relatively short period of time and has made many achievements along the way. Archaeobotany is directly related to social problems. After all, agricultural activity is the interaction of man with the environment in the process of formation of new plant communities (cultivated plants and weeds), deforestation, the emergence of new landscapes. Ukraine together with Moldova is the first region on the territory of the Eastern European plain, which found itself on the path of Neolithic farmers of Middle Eastern origin, who came here through the Balkans from the Middle East and brought with them the range of the first domesticated plants combined with traditional cultivation. Based on the study of archaeobotanical materials picture of the appearance and distribution of these cultivated plants in Ukraine is reproduced. The change in the composition of the most widely used edible plants, especially cereals: wheat, barley, rye, oats, millet, as well as legumes: peas, lentils and industrial crops – flax, hemp for thousands of years, in different chronological periods, starting from the appearance of the first Asia Minor agricultural tribes to the Middle Ages. In recent years archaeobotanists have been studying not only traditional fossil charred grains and seeds and their imprints, but also studying charred fragments of parenchyma, charcoals, and phytoliths. The chemical composition of plant origin, which is studied using a mass spectrometer, is taken into account, and studies of stable isotopes and the composition of ancient DNA are carried out. Genetic markers are used to determine the condition of fossil plants and animals, whether they are cultivated or wild, as well as to address the time and place of origin of domesticated plants and animals. Each of these studies requires modern technical equipment and special knowledge.
... The mountain, since then, has become a great source of many products having economic importance for people. For example, a previous study revealed that cultivation of einkorn cereals started around the mountain, and ancestors of many kinds of cereals still grow as wild cereals on the slopes of the mountain (Heun et al, 1997). Livestock remains one of the main activities of natives and one of the most important livestock practices, bee-keeping largely depends on the flora of the mountain. ...
Karacadağ Mountain, the shield volcano in Southeastern Türkiye, has been a treasure of biodiversity in terms of flora and fauna that serve a diverse number of agricultural practices and forestry. The floral composition is mainly composed and represented by the shrub plant, Astragalus spp. which is under threat by insect pests. This study aims to report the coleopteran woodboring species infesting Astragalus spp. along with their density and infestation rate on different altitude levels (lowest=1400-1500 m, middle=1600-1700 m, and upper=1850-1950 m) in 2016. Laboratory incubation of the specimens yielded six woodboring species: Agapanthia coeruleipennis Frivaldszky, 1878 and Xylotrechus sieversi (Ganglbauer, 1890) (Coleoptera: Cerambycidae), Sphenoptera (s.str.) anthracina Jakovlev, 1887, Sphenoptera (s.str.) coracina (Steven, 1829), Sphenoptera (s.str.) tragacanthae (Klug, 1829), and Anthaxia (s.str.) truncata Abeille de Perrin, 1900 (Coleoptera: Buprestidae). Among these species, S. anthracina is recorded for the first time in Turkish fauna. The infestation rate did not differ between altitudes while the number of woodborer pests per plant was highest at the highest level of altitude which did not differ between lowest and middle altitude levels.
... A modification of this hypothesis suggested by Kilian et al. (2007) was that a specific wild einkorn race, which arose without human intervention, underwent multiple independent domestication events. The second hypothesis, relying on ecological and molecular information, assumes that einkorn domestication was largely conscious, rapid, geographically localized in the Karacadag region and monophyletic (Heun et al., 1997;Brandolini et al., 2016). ...
... Wheat is one of the most important cereal crops and is consumed as a staple food by humans on almost every continent in the world. It was the first domesticated crop about 10,000-12,000 years ago around the Fertile Crescent and played an important role in initiating the agricultural revolution [1][2][3][4]. The wild form of durum wheat, wild emmer (Triticum turgidum ssp. ...
Durum wheat is an economically and nutritionally important cereal. The increase in durum wheat yield is mostly associated with improving grain traits, and the grain size- and shape-related traits are directly related to wheat yield. In addition, grain size influences the seed germination rate and seedling vigor, which play key roles in stand establishment and yield. Thus, it is important to investigate grain traits both agro-morphologically and genetically. In this study, a panel of durum wheat, consisting of 146 genotypes, was evaluated for grain traits agro-morphologically and a genome-wide association study (GWAS) was conducted to dissect the genomic regions associated with these traits. As a result of the GWAS, a total of 41 marker-trait associations (MTAs) were identified on different chromosomes of durum wheat. Of these MTAs, only 11 were stable across environments. A BLAST search for the flanking sequences of every stable MTA in the Svevo genome identified 18 putative candidate genes directly associated with the seed traits of different plants, particularly wheat seeds. In conclusion, the annotation results and literature information provide strong evidence that the identified stable MTAs and their candidate genes may have important functions in the formation of wheat grain traits. After the validation of these MTAs with different fine-mapping and functional characterization studies, these loci may provide valuable information for geneticists and breeders to improve wheat yield.
... Multiple natural and artificial einkorn introgressions into bread wheat containing agriculturally important genes have been described [6][7][8][9][10] . Population genetic analyses indicate that wild einkorn clusters into three distinct groups (races α, β and γ) and point to a region around the Karacadağ mountains in Southeastern Turkey as the site of einkorn domestication [11][12][13][14][15][16][17] . ...
Einkorn (Triticum monococcum) was the first domesticated wheat species, and was central to the birth of agriculture and the Neolithic Revolution in the Fertile Crescent around 10,000 years ago1,2. Here we generate and analyse 5.2-Gb genome assemblies for wild and domesticated einkorn, including completely assembled centromeres. Einkorn centromeres are highly dynamic, showing evidence of ancient and recent centromere shifts caused by structural rearrangements. Whole-genome sequencing analysis of a diversity panel uncovered the population structure and evolutionary history of einkorn, revealing complex patterns of hybridizations and introgressions after the dispersal of domesticated einkorn from the Fertile Crescent. We also show that around 1% of the modern bread wheat (Triticum aestivum) A subgenome originates from einkorn. These resources and findings highlight the history of einkorn evolution and provide a basis to accelerate the genomics-assisted improvement of einkorn and bread wheat.
... These earliest cultivated forms were diploid (genome AA) (einkorn) and tetraploid (genome AABB) (emmer) wheat's and their genetic relationships indicate that they originated from the south-eastern part of Turkey (Heun et al., 1997;Nesbitt, 1998;Dubcovsky and Dvorak, 2007). The study was undertaken to inheritance of grain yield and its components in ninety varieties (45 indigenous and 45 Exotic) of bread wheat along with four checks. ...
The present investigation was conducted at the Main Experiment Station (MES) of A.N.D. University of Agriculture and Technology, Kumarganj, Ayodhya (U.P.) during Rabi 2018-19. The study was undertaken to the inheritance of grain yield and its components in ninety varieties (45 indigenous and 45 Exotic) of bread wheat along with four checks. The experiment was laid out in Augmented Block Design. The wide range of variation for different characters and comparison of means of germplasm line indicates the existence of a very high degree of variability for all the characters in the indigenous as well as exotic lines of wheat. Almost high to moderate estimates of broad sense heritability was recorded for tillers per plant followed by days to maturity, spikelets per spike, harvest index, test weight, days to 50% flowering, grains per spike, and the peduncle length. The characters which exhibited higher estimates of PCV and GCV were harvest index and days to maturity. The high estimates of genetic advance in percent of mean were recorded for harvest index, days to maturity, tillers per plant, and for plant height.
... Einkorn wheat, Triticum monococcum, (2n = 2x = 14, A m A m ) is the first cultivated crop that led to the initiation of agriculture (Nesbitt and Samuel 1996). It was domesticated around Karacadağ mountain in Turkey (Kilian et al. 2007;Heun et al. 1997;Özkan et al. 2002). This is the only cultivated diploid wheat which has been grown for thousands of years in the Middle East, Central Asia, Europe, and North Africa. ...
Einkorn wheat is a diploid (AmAm genome) and is the first cultivated crop that initiated agriculture. It is related to durum and bread wheat, and it harbours unique genes that can be used for wheat improvement. Grain size and shape are the main breeding targets due to their direct relation to yield and milling quality. To understand the genetic control of the grain size and shape-related traits in Einkorn wheat, a biparental population of 150 F8 recombinant inbred lines (RILs) derived from a cross between an advanced einkorn line (ID1623) and a cultivar (MONLIS) was used. The RIL population was genotyped with SNP, Silico-DArT, and SSR markers and a genetic map comprising seven linkage groups (representing n = 7) was constructed. The map contained 3716 markers distributed across 760 loci with a total length of 1216.09 cM and an average density of one locus every 1.60 cM. Composite Interval Mapping was used to detect the quantitative trait loci (QTLs) controlling seven grain size and shape-related traits using genetic map and the phenotypic data collected from five different environments and the BLUP (Best Linear Unbiased Prediction) values. A total of 33 QTLs (25 novel QTLs) were detected, which were distributed on all the seven einkorn chromosomes. Of these, 14 QTLs distributed on four chromosomes (2Am, 3Am, 5Am, and 6Am) were stable across environments. Three QTL hot spots were observed on chromosomes 2Am, 5Am and 6Am. Seven QTLs, one each for the seven traits with highest PVE% (up to 14–26% PVE in individual environments) were recommended for marker-assisted recurrent selection for improvement of grain traits in einkorn wheat. The study thus provides novel and important genetic information to help understand the genetic control of grain size and shape-related traits and also the genomic resources for use in cultivated einkorn wheat breeding.
... A well-known example is common wheat (Triticum aestivum, 2n = 6x = 42, AABBDD). This crop originated from the hybridization between cultivated wheat T. turgidum (AABB) and the weed Aegilops tauschii (DD) ~ 10,000 years ago [8,9]. T. turgidum cultivation is still associated with weedy Ae. tauschii in Middle Eastern agroecosystems, which is thought to be the birthplace of common wheat [10]. ...
Background
Weeds are not only economically important but also fascinating models for studying the adaptation of species in human-mediated environments. Aegilops tauschii is the D-genome donor species of common wheat but is also a weed that influences wheat production. How shading stress caused by adjacent wheat plants affects Ae. tauschii growth is a fundamental scientific question but is also important in agriculture, such as for weed control and wheat breeding.
Result
The present study indicated that shade avoidance is a strategy of Ae. tauschii in response to shading stress. Ae. tauschii plants exhibited growth increases in specific organs, such as stem and leaf elongation, to avoid shading. However, these changes were accompanied by sacrificing the growth of other parts of the plants, such as a reduction in tiller number. The two reverse phenotype responses seem to be formed by systemically regulating the expression of different genes. Fifty-six genes involved in the regulation of cell division and cell expansion were found to be downregulated, and one key upstream negative regulator (RPK2) of cell division was upregulated under shading stress. On the other hand, the upregulated genes under shading stress were mainly enriched in protein serine/threonine kinase activity and carbon metabolism, which are associated with cell enlargement, signal transduction and energy supply. The transcription factor WRKY72 may be important in regulating genes in response to shading stress, which can be used as a prior candidate gene for further study on the genetic regulation of shade avoidance.
Conclusions
This study sheds new light on the gene expression changes and molecular processes involved in the response and avoidance of Ae. tauschii to shading stress, which may aid more effective development of shading stress avoidance or cultivars in wheat and other crops in the future.
... U pšenice jednozrnky (Tritium monococcum) výrazně promluvila do domestikačních schémat archeogenetická analýza současných divokých populací pšenic v Anatolii a na Předním východě. Na základě genové diverzity recentních populací pšenice boetské Triticum boeoticum (předek jednozrnky) bylo jako domestikační místo stanovena docela malá oblast v regionu Karacadağ v jihovýchodním Turecku (Heun et al. 1997; obr. 3.1.18), ...
Book in the Czech language, English summary. Full-text PDF. The text of the book is split into four sections. The first section, The Prologue, is devoted to the history of research into the beginnings of agriculture. We consider it very important as without a detailed depiction of the development of concepts and historical research results it is not easy to understand the contemporary views of this issue. Much of the study deals with environmental archaeology as this science has been at the forefront of research into the beginnings of agriculture, but we also mention the development of paleoecology and agronomic sciences. The second section, with the name The Frames, describes in detail the basic palaeoecological factors (climate and vegetation paleoecology) and social conditions under which humans of the end of the Pleistocene entered the scenery of the agricultural beginnings. We also pay our attention to anthropological research of “relict“ hunting and gathering groups as these archaic communities provide model testimony of possible paleoeconomic practices that may be cautiously used as a reconstructive tool for the situation in prehistory. In the thematically regional core of our study (the third section), called The Centers, we depict events and processes in important parts of the world with the exception of the Americas. For the area of the Near East and China and to a certain extent also Africa and Europe, we used a uniform scheme of interpretation which includes the development of the local climate and vegetation, followed by the description of local communities. The final section of the book summarizes the results.
... Wild einkorn wheat, ssp. aegilopoides, displays wide genetic diversity [12][13][14] and is a valuable resource for disease resistance and grain quality-related characteristics in wheat breeding, as well as drought stress tolerance [15][16][17][18][19][20][21][22]. ...
Allopolyploid speciation is a major evolutionary process in wheat (Triticum spp.) and the related Aegilops species. The generation of synthetic polyploids by interspecific crosses artificially reproduces the allopolyploidization of wheat and its relatives. These synthetic polyploids allow breeders to introduce agriculturally important traits into durum and common wheat cultivars. This study aimed to evaluate the genetic and phenotypic diversity in wild einkorn Triticum monococcum ssp. aegilopoides (Link) Thell., to generate a set of synthetic hexaploid lines containing the various Am genomes from wild einkorn, and to reveal their trait characteristics. We examined the genetic diversity of 43 wild einkorn accessions using simple sequence repeat markers covering all the chromosomes and revealed two genetically divergent lineages, L1 and L2. The genetic divergence between these lineages was linked to their phenotypic divergence and their habitats. L1 accessions were characterized by early flowering, fewer spikelets, and large spikelets compared to L2 accessions. These trait differences could have resulted from adaptation to their different habitats. We then developed 42 synthetic hexaploids containing the AABBAmAm genome through interspecific crosses between T. turgidum cv. Langdon (AABB genome) as the female parent and the wild einkorn accessions (AmAm genome) as the male parents. Two of the 42 AABBAmAm synthetic hexaploids exhibited hybrid dwarfness. The phenotypic divergence between L1 and L2 accessions of wild einkorn, especially for days to flowering and spikelet-related traits, significantly reflected phenotypic differences in the synthetic hexaploids. The differences in plant height and internodes between the lineages were more distinct in the hexaploid backgrounds. Furthermore, the AABBAmAm synthetic hexaploids had longer spikelets and grains, long awns, high plant heights, soft grains, and late flowering, which are distinct from other synthetic hexaploid wheat lines such as AABBDD. Utilization of various Am genomes of wild einkorn resulted in wide phenotypic diversity in the AABBAmAm synthetic hexaploids and provides promising new breeding materials for wheat.
... Diploid einkorn wheat (2n = 14) is an ancient crop that mankind has been cultivating for 10 thousand years (Heun et al., 1997). The grain of this wheat is a valuable product for a healthy diet (Hidalgo, Brandolini, 2014). ...
Diploid einkorn wheat (2n = 14) is an ancient crop that people cultivate for 10 thousand years. The grain of this wheat is a valuable product for a healthy diet, which determines the increasing interest in einkorn wheat by scientists and agricultural producers. Meanwhile, the wide use of this crop is hindered by several shortcomings that complicate the usage of modern technologies: low yield, ear fragility, a tendency to lodging, and difficult grain threshing. Nevertheless, there are some preconditions for improving the agronomic properties of this crop. We carried out crosses in seven combinations with the use of three wheat species (T. boeoticum, T. monococcum, T. sinskajae) to improve the diploid einkorn wheat in terms of productivity and threshing. In total, the hybrid seed set in the crosses varies from 6.3 % to 79.7 %. In the combination of cultivated wheat T. sinskajae with wild T. boeoticum, differences in the results of reciprocal crosses are observed specifically in the hybrid seed set (in the forward cross it equals 6.3 %; in the reverse one, 48.9 %). Hybrids from reciprocal crosses of T. monococcum var. sofianum UA0300649 and T. sinskajae f. aristata were equivalent in seed set (72 and 82 %) and inheritance patterns and had similar quantitative traits. In other combinations, the seed set varied from 12.5 to 45.6 %. Hybrid depression was the most frequent (22 cases out of 49) inheritance type of the F1 quantitative traits in einkorns; dominance of the parent form with a large trait manifestation was registered in 11 cases, and heterosis in four cases. In hybrids, the inheritance of spike length is correlated with the inheritance type of the ear number (r = 0.92) and the grain number (r = 0.78) per spike. The dominance degrees after these two traits are also highly correlated (r = 0.89). The combination UA0300400 T. boeoticum var. thaoudar ARM / UA0300224 T. sinskajae var. sinskajae RUS, which manifested heterosis for kernel number per spike (Hp = 1.2), the weight of spike (Hp = 11.8) and weight of kernels per spike (Hp = 5.4) is of particular interest. The combination UA0300222 T. monococcum var. hohensteinii / UA0300224 T. sinskajae var. sinskajae is promising for creating easily threshed material.
... According to this view, the eight Neolithic founder crops were selected and domesticated once, in a rapid event that took place in a single region or 'core area' located in southeast Turkey, as suggested primarily by some genetic studies (e.g. Ladizinsky and Adler 1976;Heun et al. 1997;Mori et al. 2003;Özkan et al. 2005;Luo et al. 2007). From here, domesticated or semi-domesticated plants (and consequently agriculture) radiated to other regions Kilian et al. 2007;Özkan et al. 2011). ...
Zohary and Hopf coined the term ‘founder crops’ to refer to a specific group of eight plants, namely three cereals (einkorn, emmer and barley), four legumes (lentil, pea, bitter vetch and chickpea), and a fibre/oil crop (flax), that founded early Neolithic agriculture in southwest Asia. Zohary considered these taxa as the first cultivated and domesticated species, as well as those that agricultural communities exploited and eventually spread to Europe and other regions. As a result, these eight species soon become the hallmark of the Neolithic plant-based subsistence. However, the ‘founder crops’ concept was defined at the end of the 1980s, when the development of agriculture was considered a rapid event, and therefore, terms like domestication, agriculture and plant cultivation were used interchangeably in the literature. The aim of this paper is thus to revisit concept of the ‘Neolithic founder crops’. Through a critical review of the archaeobotanical evidence gathered in the last 40 years, we evaluate the relative contribution of the ‘eight founder crop’ species to the plant-based subsistence across different periods of the southwest Asian Neolithic. We conclude that multiple groups of ‘founder’ species could be defined depending on whether one seeks to represent the most exploited plants of the Neolithic period, the first cultivated and domesticated crops, or the species that agricultural communities cultivated and eventually spread to Europe. Improved understanding of Neolithic plant-based subsistence in general, and agriculture in particular, will be attained by moving beyond conventional narratives and exploring the evolutionary history of plants other than the original ‘founder’ species.
... Major cultivated species of wheat include: Triticum aestivum, which is a hexaploid species and is widely cultivated in the world; Triticum durum, the only tetraploid form of wheat widely used today, and the second most widely cultivated wheat; Triticum monococcum, a diploid species with wild and cultivated variants; Triticum dicoccum, a tetraploid species. These earliest cultivated forms were diploid (genome AA) (einkorn) and tetraploid (genome AABB) (emmer) wheat's and their genetic relationships indicate that they originated from the south-eastern part of Turkey (Heun et al., 1997;Nesbitt, 1998;Dubcovsky and Dvorak, 2007). ...
An experiment was conducted in a randomized complete block design with two
replications at experimental area of Research Farm, College of Agriculture, Gwalior,
(M.P.) during 2014-2015. The study was undertaken to inheritance of grain yield and its
components in thirty six wheat genotypes. Analysis of variance revealed that genotypes
were significantly differed for all characters and suggested considerable variability for all
yield and its attributing characters in the present material. Among genotypes tested, HI
1610 seems to be the best, because, it is highest yielder, early flowering, early maturing,
and bold grain size and dwarf.Almost equal magnitude of GCV and PCV showed for days
to heading, 1000 grain weight and days to maturity indicating that these characters are less
influenced by the environment while grain yield and plant height exhibited high value of
PCV over its GCV value, indicating relatively large amount of variation for these
characters was due to environment alone. The Higher heritability was observed for days to
heading, 1000 grain weight and plant height. High heritability coupled with high genetic
advance recorded for traits 1000 grain weight, plant height, days to heading and grain yield
and that these characters are governed by additive gene effects; a directional selection for
these traits would be more effective for desired genetic improvement.
... boeoticum Bois and T. dicoccoides Koern. ex Schweinf) growing in the Near East, based primarily on yield and other characteristics like most non-brittle rachis and freethreshing naked kernels (Heun et al 1997;Shewry 2009). ...
The East Mediterranean is a hotspot of agricultural biodiversity in Europe and the center of origin for wheat and other basic food crops. Hulled wheat species (einkorn, emmer and spelt) are among the most ancient cereal crops, popular within the region for hundreds of years. At a certain point in history, however, the introduction of higher-yielding, freethreshing wheats caused them to fall into a state of neglect, to such an extent that they have become a relic crop. For social, cultural or simply economic reasons, hulled wheats today are becoming popular once again and an exclusive and fashionable food for consumers.
Because of the dynamic growth of niche markets and broad demand for organic products derived from ancient or hulled wheat, organic restoration activities have started based on the conservation of genetic resources and application of evolutionary breeding. In Greece, activities carried out over the last two decades by the AEGILOPS NGO, Plant Breeding and Genetic Resources Institute, Greek Gene Bank, laboratories of universities, and seed savers have contributed to on farm-maintenance under organic farming conditions and practices (organic restoration) of hulled wheat and the creation of a growing niche market.
In this article, the organic restoration case of “Kaploutzas” an old Greek einkorn landrace, is examined and also several multi-sector market-based strategies are proposed for enforcing the necessary dynamic evolutionary processes to maintain hulled wheat and enhance cereal biodiversity before it is lost.
In order to continue humanity, we need to feed the growing population and practice agriculture in harmony with the changing environmental conditions. For this reason, we require high-yielding cultivars capable of withstanding environmental stress factors. The only solution to developing these varieties is the utilization of genetic resources. Among all plants, wheat stands out as the most important plant genetic resource. Being the first plant cultivated by humans, wheat owes its status to its high adaptability in diverse environmental conditions. Hence, it has been grown for thousands of years in almost any climate and region, becoming the primary food source for humans due to its ease of cultivation and rich nutrient content.
Local wheat varieties and their wild relatives possess gene resistant to both biotic and abiotic factors. Plants face various stresses such as drought, cold, heat, diseases, pests, and global warming. Wild wheat relatives and local populations cultivated by farmers in stressed regions for extended periods have acquired the ability to cope with these stress factors. Local varieties have gained significance not only for their resistance/tolerance to stress factors but also for their suitability to meet the changing nutritional habits and needs of today. Despite their lower yields, enthusiasts and some farmers grow these wheat varieties. Scientific studies on wheat genetic resources have been carried out for many years. Although collections of these genetic resources have been amassed worldwide, the conservation efforts for local varieties, especially in the hands of farmers, have recently started to gain importance. Just as in the past, the value of wheat genetic resources will increase in the future.
Both the wheat species Triticum aestivum L. and Triticum durum Desf., which have wheat cultivars that are most important for small grain field crops, are grown worldwide as food sources for a rapidly increasing world population. Because of recent global climate changes, pathogenic diseases, pests, and abiotic stress factors on wheat crops have become more detrimental than ever before. Increases in atmospheric temperature have stimulated drought stress as well as having caused increased vector-borne viruses and other systemic disease pathogens. Another influence caused by ongoing global climate change is that sporadic local virus diseases are becoming more widespread by vector-borne transmissions. Wheat cultivars are naturally found susceptible to at least 30 virus diseases under field conditions. However, five of those viruses, namely, yellow dwarf viruses (YDVs), wheat streak mosaic virus (WSMV), wheat dwarf virus (WDV), soilborne wheat mosaic virus (SBWMV), and wheat spindle streak mosaic virus (WSSMV), occur in all wheat-growing areas of the world and cause substantial grain yield losses. Thus, this chapter will examine host range and disease incidence geographically in the world, disease symptoms, the morphological and molecular structure of viruses, and the annual disease cycle. It will also review disease management. In conclusion, the impact of these wheat virus diseases on food security is evaluated in this chapter.
Wheat is a very important cereal plant that has been serving the nutrition of human beings for about 10,000 years and has become a strategic product of countries today. Since the day it was cultivated, it has been under the influence of many different biotic and abiotic stress factors in the regions where it grows. Harmful insects are the leading biotic stress factors. Insects appear in different phenological periods of the wheat plant in different parts of the world and sometimes cause economic damage by feeding on the root, stem, leaf, spike, flower, and grain of the plant. Throughout the world, there are many harmful insect species belonging to different orders and families that feed on the wheat plant. Today, it is known that the global potential crop loss due to harmful insects in wheat is 7.9% on average. This yield loss varies according to countries, regions, years, climatic conditions, and wheat genotypes. Preventing the damage and product losses caused by insects in wheat is an important plant protection problem of the countries. The solution to this problem passes in recognizing the harmful insects in question and successfully combating them.
In this chapter, a brief description of the life cycle, ecology, geographical distribution damages in plants, economic importance, host plants, natural enemies, and management of harmful insects that cause economic loss in wheat are given.
Spring wheat is grown in winter in temperate regions where winter temperatures do not fall below −5 ºC and only in summer in regions with very cold winters. In Türkiye, spring wheat is sown in the coastal zone such as the Aegean and Mediterranean, where winters are not very cold, and in the Southeastern Anatolia Region, where winters are mild. Wheat is a long-day crop. In conditions where the day length is 12–14 h, they pass into the generative period.
Wheat is a very important and strategic product for Türkiye. Since it was first produced, it has been in the first place for nutrition in the world. Although it changes every year, annual production changes around 20 million tons. Facultative and winter wheat varieties account for 60–70% of the wheat produced, and the rest is spring wheat, which is usually grown under rainfed or irrigation conditions in the coastal and southeastern regions of the country. The vast majority of grown spring wheat consists of varieties and germplasm originating from the International Maize and Wheat Improvement Center (CIMMYT).
Wheat research in Türkiye first started at Eskişehir Sazova Seed Improvement Station, which was reported to be established under decree dated December 13, 1925. This was followed by studies in Yeşilköy, Adapazarı, Adana, and Samsun Seed Stations established in 1926. Currently, eight research institutes established in Samsun, Sakarya, İzmir, Antalya, Adana, Kahramanmaraş, Diyarbakır, and Şanlıurfa province are studying spring wheat.
The genus Clavibacter (family Microbacteriaceae) includes Gram-positive actinobacterial species infecting diverse agricultural crops such as alfalfa, maize, pepper, potato, tomato, and wheat. Clavibacter tessellarius causing bacterial mosaic of wheat was reported for the first time in 1976 in Nebraska (USA). Since then, the pathogen was known to be restricted to North America i.e. Canada and USA. While bacterial strains suspected to be C. tessellarius have occasionally been isolated from wheat samples outside North America, occurrence of the pathogen in the Old World has not yet been confirmed by the plant protection authorities. In spring 2020 (January to May) dozens of wheat fields with young plants showing leaf chlorosis and mild mosaic symptoms were surveyed and sampled in Southern Iran. Among 192 surveyed wheat fields, Gram-positive orange-pigmented actinobacterial strains were isolated from samples of 53 fields. Altogether, 61 bacterial strains were identified as C. tessellarius based on the phenotypic features, PCR-based molecular tests, and pathogenicity assays. Multilocus sequence analysis of five housekeeping genes i.e. atpD, dnaK, gyrB, ppk, and rpoB using ten representative strains from diverse geographic areas in Southern Iran confirmed the phylogenetic status of the strains within C. tessellarius. Our data indicate that the bacterial mosaic pathogen occurs in four southern Iranian provinces including Bushehr, Fars, Kerman, and Khuzestan. Thus, occurrence of C. tessellarius outside North America is confirmed. Potential risks associated with the emergence of the pathogen in the Old World, and the need for strict quarantine measures in the area is further discussed.
Wheat dwarf virus (WDV) has caused considerable economic loss in the global production of grain crops. Knowledge of the evolutionary biology and population history of the pathogen remain poorly understood. We performed molecular evolution and worldwide phylodynamic analyses of the virus based on the genes in the protein-coding region of the entire viral genome. Our results showed that host-driven and geography-driven adaptation are major factors that affects the evolution of WDV. Bayesian phylogenetic analysis estimates that the average WDV substitution rate was 4.240 × 10⁻⁴ substitutions/site/year (95% credibility interval, 2.828 × 10⁻⁴–5.723 × 10⁻⁴), and the evolutionary rates of genes encoding proteins with virion-sense transcripts and genes encoding proteins with complementary-sense transcripts were different. The positively selected sites were detected in only two genes encoding proteins with complementary-sense, and WDV-barley are subject to stronger purifying selection than WDV-wheat. The time since the most recent common WDV ancestor was 1746 (95% credibility interval, 1517–1893) CE. Further analyses identified that the WDV-barley population and WDV-wheat population experienced dramatic expansion-decline episodes, and the expansion time of the WDV-barley population was earlier than that of the WDV-wheat population. Our phylogeographic analysis showed that the WDV population originating in Iran was subsequently introduced to Europe, and then spread from Eastern Europe to China.
This paper investigates the causes and the consequences of the emergence of agriculture in the Middle East. Agriculture has emerged in many parts of the world since the end of the last Ice Age about 15 000 years ago. The paper first surveys the Palaeolithic Period to understand why agriculture did not emerge earlier. Then the paper considers the processes that led to the emergence of agriculture in the Middle East. The question is approached as a problem in economic history, and the archaeological record is understood in terms of changes in labour productivity, as measured by calories produced per hour of work, and in the size of the associated agricultural surplus (the difference between the calories produced by a food procurement system and the calories required to sustain the people trying to support themselves with that system). The shift from foraging to the cultivation of wild seeds and the later impact of the emergence of domestic seed and improvements in tool design are assessed in this framework. It is also used to analyse the impact of agriculture on human wellbeing, the origin of manufacturing (e.g. pottery), and the emergence of inequality, states, and warfare.
Background and aims:
The development and morphology of crop plants have been profoundly altered by evolution under cultivation, initially through unconscious selection without deliberate foresight, and later by directed breeding. Wild wheats remain an important potential source of variation for modern breeders, however, the sequence and timing of morphological changes during domestication are not fully resolved.
Methods:
We grew and measured 142 wheat accessions representing different stages in wheat evolution, including three independent domestication events, and compared their morphological traits to define the morphospace of each group.
Key results:
The results show that wild and domesticated wheats have overlapping morphospaces, but each also occupies a distinct area of morphospace from one another. Polyploid formation in wheat increased leaf biomass and seed weight but had its largest effects on tiller loss. Domestication continued to increase the sizes of wheat leaves and seeds, and made wheat grow taller with more erect architecture. Associated changes to the biomass of domesticated wheats generated more grains and achieved higher yields. Landrace improvement subsequently decreased tiller and spike numbers, to focus resource allocation to the main stem, accompanied by a thicker main stem and larger flag leaves. During the Green Revolution, wheat height was reduced to increase the harvest index and therefore yield. Modern wheats also have more erect leaves and larger flower biomass proportions than landraces.
Conclusions:
Quantitative trait history in wheat differs by trait. Some trait values show progressive changes in the same direction (e.g. leaf size, grain weight), others change in a punctuated way at particular stages (e.g. canopy architecture), while other trait values switch directions during wheat evolution (e.g. plant height, flower biomass proportion). Agronomically valued domestication traits arose during different stages of wheat history, such that modern wheats are the product of >10,000 years of morphological evolution.
Stripe rust, caused by Puccinia striiformis f. sp. tritici (Pst), is one of the most destructive fungal diseases of wheat. Cultivated einkorn (Triticum monococcum ssp. monococcum, 2n=2x=14, AmAm), one of the founder crops of agriculture, harbors unexploited genetic sources for wheat improvement. An advanced wheat line Z15-1949 with 42 chromosomes, selected from the hybrids of Pst-susceptible common wheat cultivar Crocus and resistant T. monococcum accession 10-1, exhibits high resistance to a mixture of the Chinese prevalent Pst races. Genetic analysis on F1, F2, and F2:3 generations of the cross between Z15-1949 and Pst-susceptible common wheat SY95-71 indicated that the resistance of Z15-1949 was conferred by a recessive gene, tentatively designated as YrZ15-1949. This gene was mapped to the short arm of chromosome 7D using the Wheat 55K SNP array, flanked by markers KASP-1949-2 and KASP-1949-10 within a 3.3 cM genetic interval corresponding to 1.12 Mb physical region in the Chinese Spring reference genome V2.0. The gene differs from previously reported Yr genes on 7D based on their physical positions, and is probably a novel gene. YrZ15-1949 would be a valuable resource for developing Pst-resistant wheat cultivars and the linked markers could be used for the marker-assisted selection.
Araştırmada, Zenit ile B27 yerel makarnalık buğday çeşidi ve bu çeşitlerin melezlenmesi sonucunda elde edilen 11 adet makarnalık buğday genotipi, 8 adet allel spesifik DNA markörü kullanılarak bazı hastalık ve kalite ile ilgili allellerin tespiti yapılmıştır. Moleküler tarama sonucunda kullanılan 8 DNA markörü 25 adet allel üretirken, çalışmada kullanılan DNA markörlerinin ortalama polimorfizm bilgi içeriği (PIC) değeri 0.9775 olarak tespit edilmiştir. En yüksek polimorfizm bilgi içeriği değeri 0.99 olarak hesaplanırken, en düşük polimorfizm bilgi içeriği değeri 0.95 olarak hesaplanmıştır. Çalışmada DNA markörleri tarafından üretilen 25 allel kullanılarak oluşturulan dendrogramda, ebeveynlere göre iki ana grup meydana gelmiştir. Zenit×B27-7, Zenit×B27-9 ve Zenit×B27-11 ile Zenit×B27-5, Zenit×B27-6, Zenit×B27-8, Zenit×B27-10 melez kombinasyonları % 100 benzer bulunmuştur. Araştırma sonuçlarına göre Zenit×B27-1, Zenit×B27-2 ve Zenit×B27-3 genotiplerinde Waxy (Wx-A1) özelliğine ait genler belirlenirken, Zenit×B27-3 ve Zenit×B27-5 genotiplerinde yüksek proteine (Gpc-B1) ait gen bölgesinin yer aldığı saptanmıştır.
Bu çalışma, 2019-2020 buğday yetiştirme sezonunda Mardin ili Artuklu İlçesi yağışa dayalı şartlarında Augmented Deneme Desenine göre planlanarak Güneydoğu Anadolu Bölgesine özgü 80 adet yerel makarnalık ve 10 adet standart çeşit kullanılarak yürütülmüştür. Deneme, her blokta 20 adet yerel genotip ve 10 adet kontrol çeşit olacak şekilde düzenlenmiştir. Güneydoğu Anadolu Bölgesine özgü bu yerel makarnalık çeşitlerin karakterizasyonunun amaçlandığı çalışmada; yerel buğday çeşitlerinin ve kontrol çeşitlerinin alınan gözlemleri sırasıyla tane verimi 229 kg/da- 371 kg/da; biyolojik verimleri 1313 kg/da –1218 kg/da; bin tane ağırlıkları 42.9 g – 40.15 g, başaklanma gün sayısı 117.8 gün-111 gün; protein oranı % 18.02 - % 14.94 arasında değişmiştir. Elde edilen verilere göre, yerel buğday çeşitlerinin bitki boyu, biyolojik verim, tanede protein oranı bakımından kontrol çeşitlerinden daha yüksek değerlere sahip olduğu tespit edilirken, tane verimi bakımından ise kontrol çeşitlerin daha yüksek değerlere sahip olduğu tespit edilmiştir. Yerel çeşitlerin, modern ıslah çeşitlerine göre daha geç başaklandığı ve yeşil kalma süresinin daha uzun olduğu gözlemlendiği çalışmada, yerel çeşitlerin daha çok kışlık gelişme tabiatına sahip özellikler taşıdığı, özellikle tanede protein oranın ve biyolojik verimin artırılması çalışmalarında gen kaynağı olarak kullanılabilecek çok sayıda potansiyel yerel çeşit olduğu tespit edilmiştir. Elde edilen verileri GGE biblot analiz metodolojisine göre değerlendirilmiş, yapılan değerlendirmede, tane verimi (TV), başakta tane sayısı (BSTS), başakta tane ağırlığı bakımından ST8 (kontrol) G80 (yerel) genotipleri en yüksek değerlere sahip olmuşlardır. Yerel genotipler bin tane ağırlığı (BTA), peduncle uzunluğu (PU), bitki boyu, tanede protein oranı (TPO), biyolojik verim (BV) ve başaklanma gün sayısı (BGS) bakımından daha yüksek değerlere sahip olurken, bu özellikler açısından G8 (yerel) en yüksek değerlere sahip olmuştur. İncelenen özellikler bakımından geniş bir varyasyona sahip olduğu tespit edilen bu yerel çeşitlerinin korunması ve gelecek kuşaklara aktarılması konusunun önemli olduğu ve aynı zamanda ulusal ve uluslararası buğday ıslah programlarında kullanılabilecek yararlı özelliklere sahip olduğu tespit edilmiştir.
Einkorn wheat (Triticum monococcum) is one of the oldest cereal crops to be domesticated by human beings, playing essential role in early agriculture development. Today, it is considered an important genomic resource for modern wheat improvement, especially for resistance genes against pests and diseases. However, the exploration and utilization of useful genes from T. monococcum is rather limited due to the lack of a high-quality genome and annotation for this species despite two
draft genome assemblies released recently. Here, we report a near-complete genome assembly for T. monococcum with a total length of 5.11 Gb with a contig N50 of 131.2Mb and scaffold N50 of 728.66Mb, representing a genome assembly of highest quality for any wheat genome reported. Phylogenomic analysis confirmed T. monococcum is closely-related to T. urartu, the progenitor of wheat A subgenomes. A 4AL/5AL terminal translocation is present in the diploid species T. urartu and T.
monococcum, taking place before wheat polyploidization. The genome sequence confirmed the introgression of T. monococcum rust resistance genes at 5AmL into modern bread wheat varieties. This near-complete reference genome of T. monococcum will be an essential resource for wheat functional and evolutionary
genomic studies and expedite the cloning of useful genes in T. monococcum for future wheat improvement.
Türkiye is one of the origin centers of durum wheat and among the important producer countries of durum wheat. The aim of this study is to examine some durum wheat genotypes with different characteristics in terms of some agronomic and quality characteristics and to determine the relationships between features. The study was carried out in four different environments with supplementary irrigated and based of rainfed in Diyarbakir province conditions. Trial design carry out according to Randomized Complete Blocks Split Plots Experiment Design and three replications. It was determined that there were significant differences at the p<0.01 level between genotypes in all the traits examined. According to the research results, change range of average values in durum wheat varieties were determined as; heading time (HT) 170.33-178.42 days, plant height (PH) 93.0-139.2 cm, the number of spikes per square meter (SN) 441.50-567.50 spikes/m2, the number of grains per spike (GN) 40.7-65.5 grains/spike, thousand grain weight (TGW) 32.4-47.0 g, test weight (TW) 77.5-85.6 kg/hl, protein ratio (PR) 12.72-17.21%, sedimentation amount (SA) 9.58-25.08 ml, b yellowness value (YV) 18.27-27.90, vitreousness ratio (VR) 75.42%-85.42%. Plant height exhibited a positive correlation with protein content (r = 0.728**), and sedimentation amount also demonstrated a positive correlation with the b yellowness value (r = 0.649**). As a general trend, genotypes with spring attributes were positioned ahead of those with winter characteristics. It has been observed that winter genotypes have a heading time 5-6 days later than that of spring genotypes. Since Fırat-93 (TGW), Kunduru 1149 (PR), Urfa 2005 (TW and VR) and Candidate 1 (SA and YV) genotypes are at the forefront in terms of quality parameters. It would be beneficial to use these genotypes as parents and to protect them as genitor.
In this chapter, the taxonomical complexities of the genus Triticum are presented. Following the biological concept of species, the genus contains six species, two diploids, two tetraploid, and two hexaploids. The characteristic morphology of the genus and that of the wild forms, their geographic distribution, and ecological affinities, as well as their preadaptation for domestication and the processes leading to wheat domestication are reported. The origin and evolution of the diploid species, and the genome analysis of the allopolyploids are reviewed. Origin of the A, B, and D subgenomes of allopolyploid wheats, are presented. The relationships between Triticum species and other Triticineae are discussed.
The chapter deals with the various steps, periods, and processes that led to the domestication of the wheat as well as with the archaeological sites where domestication took place. Additionally, the chapter describes the ecogeographical characteristics of the area of wheat domestication, the selection of non-brittle rachis, large grain size, rapid and synchronous germination, free-threshing grains, and yield. The genetic basis of non-brittle rachis and free-threshing grains are delt with in details. The formation of hexaploid wheat, T. aestivum , and the spread of its free-threshing form to almost all parts of the globe to become the main cultivated wheat, are reviewed. The production of synthetic Triticum aestivum , and Triticale are also referred to in this chapter.
The distribution and the Caracterization of constitutive heterochromatin in A-Am genomes of diploid wheat (progenitor), polyploid wheat (hybrids) and triticales (primary and secondary) are analyzed and compared by C-bands. The Comparison of zones rich in highly repeated DNA sequences marked by C bands on the all chromosomes of Am-A genomes revealed an important structural heterogeneity. Four chromosomes of Triticum monococcum (1Am-3Am-4Am-5Am) are almost similar to their homologues in wheat (Triticum durum , Triticum aestivum) and triticale, by the presence or absence of C bands. Contrary to the chromosomes 2Am (rich in heterochromatin), 6Am-7Am (absence of C bands) show a great differentiation compared to their homologues of Triticum durum and Triticum aestivum and x-Triticosecale Wittmack. In the triticales, A genome chromosomes are richer in heterochromatin compared to theirs homologous of polyploid wheats. This is explained by a "genome shock The confrontation of a C-bands genome (Triticum monococcum) with a C+ bands genome (durum wheat / or common wheat) produces an interspecific hybrid which at the sixth generation reveals C+ bands (triticales).The variations observed in our vegetal material indicated the existence of an intervarietal and interspecific heterochromatic polymorphism. The presence of B chromosomes in triticales, could be explained as a manifestation of their adaptation.
Since prehistory, men and innovations had flowed from East to West for centuries along the Mediterranean and had transformed its economies and societies. Later commercial city-states emerged on the shores of the Inner Sea and reinforced this process favoured by the low sea transport costs. Several of them became maritime empires or thalassocracies.
Powdery mildew caused by Blumeria graminis f. sp. tritici ( Bgt ), is one of the serious diseases that attack wheat plants during the growing season. The Bgt virulence was screened against 16 Egyptian wheat cultivars and 21 powdery mildew differential lines carrying different ( Pm ) resistance genes in two regions (Kafr-Elsheikh and El-Minya) in Egypt 2021–2022. Broad-spectrum virulence and high diversity were observed at both regions. Regional populations of Bgt in Egypt are independent of each other. Genes, Pm2a, Pm3a, Pm1b, Pm3b, Pm21, Pm34, Pm36, Pm37, Pm53, PmNCA6, PmMIAG12, and PmNCAG13 were nationally effective, while Pm4a, Pm4b , Pm6 , and Pm8 appeared to be defeated throughout the country affected by broad-spectrum virulence. Field responses showed that only four cultivars, Misr-1, Misr-2, Sakha-95 and Shandweel-1 were resistant in both regions. A strong correlation was recorded between FDS and AUDPC which implies that wheat pathologists and breeders may be able to assess their lines by a single scoring at an appropriate time. Phenotypic and genotypic data proved that ineffective Pm8 was present in four susceptible cultivars, Sakha-93, Sakha-94, Gemmeiza-10, and Gemmeiza-11, while it was present in combination with effective Pm3a in two resistant cultivars, Sakha-95 and Shandweel-1. This suggests that cultivar susceptibility may be attributed to the existence of ineffective gene Pm8 potentially suppressed by effective Pm3a in hexaploid-resistant wheat. Pyramiding effective resistance genes particularly those that have a suppression effect like Pm3a may be a viable option to avoid the risk of broad-spectrum Bgt virulence at a regional scale.
L'approccio olistico del Comfederalismo Democratico ha trovato in Medio Oriente (MO) un territorio di sperimentazione concreta in cui le donne sono diventate ‘eliche’ del cambiamento. Le donne del movimento curdo organizzano formazioni in cui il dialogo intellettuale è fuso con l’esperienza concreta, dove sono recuperati e connessi i saperi, dedicando tempo alla comprensione della polarità femminile. La riscoperta del territorio marginale per eccellenza, il corpo della donna, da cui sono partite tutte le colonizzazioni, attiva un cambiamento dalla portata storica non solo per il MO. Accademie informali fondate sulle montagne del Kurdistan e poi diffusesi in Europa, MO, Americhe; accademie spontanee all’ombra delle querce, davanti ad un çay, lungo un corso d’acqua, creano esperienze di formazione lontane dai centri formali, edificano un mondo pluri-verso (Borghi 2020). Formazione è cambiamento, è costruzione di società. Nei campi di Jineoloji le donne scrivono insieme la storia, condividono leggende locali di cui mettono in luce i legami con la cultura neolitica della Grande Madre, praticano una pedagogia influenzata dalle esperienze di Freire, producono co-narrazione e cultura. Creatività e libertà, due concetti che emergono dai casi studio come fattori propulsivi della trasformazione della società e delle persone: possibilità di estendere il proprio immaginario oltre le barriere che fino a poco prima sembravano invalicabili; prendere coscienza della responsabilità individuale (morale/etica) nelle relazioni (con tutti gli organismi come fattori propulsivi della trasformazione della società e delle persone: possibilità di estendere il proprio immaginario oltre le barriere che fino a poco prima sembravano invalicabili; prendere coscienza della responsabilità individuale (morale/etica) nelle relazioni (con tutti gli organismi).
In many regions worldwide wheat (Triticum aestivum L.) plants experience terminal high temperature stress during the grain filling stage, which is a leading cause for single seed weight decrease and consequently for grain yield reduction. An approach to mitigate high temperature damage is to develop tolerant cultivars using the conventional breeding approach which involves identifying tolerant lines and then incorporating the tolerant traits in commercial varieties. In this study, we evaluated the terminal heat stress tolerance of 304 diverse elite winter wheat lines from wheat breeding programs in the US, Australia, and Serbia in controlled environmental conditions. Chlorophyll content and yield traits were measured and calculated as the percentage of non-stress control. The results showed that there was significant genetic variation for chlorophyll retention and seed weight under heat stress conditions. The positive correlation between the percent of chlorophyll content and the percent of single seed weight was significant. Two possible mechanisms of heat tolerance during grain filling were proposed. One represented by wheat line OK05723W might be mainly through the current photosynthesis since the high percentage of single seed weight was accompanied with high percentages of chlorophyll content and high shoot dry weight, and the other represented by wheat Line TX04M410164 might be mainly through the relocation of reserves since the high percentage of single seed weight was accompanied with low percentages of chlorophyll content and low shoot dry weight under heat stress. The tolerant genotypes identified in this study should be useful for breeding programs after further validation.
Siyez buğdayı (Triticum monococcum), ülkemizde özellikle Kastamonu ilinde yetiştirilen ve yaklaşık on bin yıl önce kültüre alınmış günümüz buğdaylarının atasıdır. Morfolojik yapısı itibariyle diğer buğdaylardan farklı olarak kavuzuyla hasat edilmekte, kepek tabakası endosperme daha sıkı bağlı olduğundan diğer buğdayların işlendiği modern değirmenlerde öğütülememektedir. Bu nedenle hasadından sonra selektörden geçirme, kavuzdan ayrıma işlemleri yapılmakta, tam buğday formunda değirmende kırıldıktan sonra elenerek kepek tabakası ayrılmaktadır. Nişastalı endospermce zengin %25 düzeyinde kepek değirmen atık/yan ürünü olarak ayrılmaktadır. Siyez kepeği iri taneli endosperm içermesi nedeniyle gıda sektöründe kullanım alanı bulamamakta ve hayvan yemi olarak değerlendirilmektedir. Bu çalışmada, buğday unu yerine siyez kepeğinden spontan fermantasyon ile üretilen ekşi mayanın ekmekte kullanımının optimizasyonu yapılmış, ekmek kalite ve biyoaktif nitelikleri üzerindeki etkileri araştırılmıştır. Optimizasyon sonucu Hamur Verimi (HV):200 oranında hazırlanan siyez kepeği ekşi mayasının ekmekte %34.23 oranında kullanılması gerektiği buna karşılık formülasyondaki su oranın da %52.78 düzeyinde olması gerektiği belirlenmiştir. Üretilen optimize ekmek, ekmeklik buğday kepeği içerikli ekmeğe göre daha hacimli, yeme kalitesi, lezzet ve iç dokusu daha iyi olarak nitelendirilmiştir. Aynı zamanda fitik asit degradasyonu, mineral madde ve protein içeriğinde de artış tespit edilmiştir.
Plastids and mitochondria are organelles of plant cells with small genomes, which may exhibit discordant microevolution as we earlier revealed in pea crop wild relatives. We sequenced 22 plastid and mitochondrial genomes of Pisum sativum subsp. elatius and Pisum fulvum using Illumina platform, so that the updated sample comprised 64 accessions. Most wild peas from continental southern Europe and a single specimen from Morocco were found to share the same organellar genome constitution; four others, presumably hybrid constitutions, were revealed in Mediterranean islands and Athos Peninsula. A mitochondrial genome closely related to that of Pisum abyssinicum, from Yemen and Ethiopia, was unexpectedly found in an accession of P. sativum subsp. elatius from Israel, their plastid genomes being unrelated. Phylogenetic reconstructions based on plastid and mitochondrial genomes revealed different sets of wild peas to be most related to cultivated P. sativum subsp. sativum, making its wild progenitor and its origin area enigmatic. An accession of P. fulvum representing ‘fulvum-b’ branch, according to a nuclear marker, appeared in the same branch as other fulvum accessions in organellar trees. The results stress the complicated evolution and structure of genetic diversity of pea crop wild relatives.
A novel DNA fingerprinting technique called AFLP is described. The AFLP technique is based on the selective PCR amplification
of restriction fragments from a total digest of genomic DNA. The technique involves three steps: (i) restriction of the DNA
and ligation of oligonucleotide adapters, (ii) selective amplification of sets of restriction fragments, and (iii) gel analysis
of the amplified fragments. PCR amplification of restriction fragments is achieved by using the adapter and restriction site
sequence as target sites for primer annealing. The selective amplification is achieved by the use of primers that extend into
the restriction fragments, amplifying only those fragments in which the primer extensions match the nucleotides flanking the
restriction sites. Using this method, sets of restriction fragments may be visualized by PCR without knowledge of nucleotide
sequence. The method allows the specific co-amplification of high numbers of restriction fragments. The number of fragments
that can be analyzed simultaneously, however, is dependent on the resolution of the detection system. Typically 50—100 restriction
fragments are amplified and detected on denaturing polyacrylamide gels. The AFLP technique provides a novel and very powerful
DNA fingerprinting technique for DNAs of any origin or complexity.
The magnitudes of the systematic biases involved in sample heterozygosity and sample genetic distances are evaluated, and formulae for obtaining unbiased estimates of average heterozygosity and genetic distance are developed. It is also shown that the number of individuals to be used for estimating average heterozygosity can be very small if a large number of loci are studied and the average heterozygosity is low. The number of individuals to be used for estimating genetic distance can also be very small if the genetic distance is large and the average heterozygosity of the two species compared is low.
This paper deals with carbonized grains of the two-seeded variety of wild einkorn wheat (Triticum boeoticum var. thaoudar) and of wild barley (Hordeum spontaneum) from Tell Mureybit in northern Syria. Radiocarbon dates suggest an age between about 7500 and 8400 B.C. for the levels from which the seeds were recovered. The carbonized seeds are compared with modern wild eintorn and barley. It is likely that especially the wild einkorn was not collected in the vicinity of Tell Mureybit, but in the adjacent part of Turkey, at a distance of at least 100 km from the site.
Amplified fragment length polymorphism (AFLP) analysis is a PCR-based technique capable of detecting more than 50 independent loci in a single PCR reaction. The objectives of the present study were to: (1) assess the extent of AFLP variation in cultivated (Gycine max L. Merr.) and wild soybean (G. soja Siebold & Zucc.), (2) determine genetic relationships among soybean accessions using AFLP data, and (3) evaluate the usefulness of AFLPs as genetic markers. Fifteen AFLP primer pairs detected a total of 759 AFLP fragments in a sample of 23 accessions of wild and cultivated soybean, with an average of 51 fragments produced per primer pair per accession. Two-hundred and seventy four fragments (36% of the total observed) were polymorphic, among which 127 (17%) were polymorphic in G. max and 237 (31%) were polymorphic in G. soja. F2 segregation analysis of six AFLP fragments indicated that they segregate as stable Mendelian loci. The number of polymorphic loci detected per AFLP primer pair in a sample of 23 accessions ranged from 9 to 27. The AFLP phenotypic diversity values were greater in wild than in cultivated soybean. Cluster and principal component analyses using AFLP data clearly separated G. max and G. soja accessions. Within the G. max group, adapted soybean cultivars were tightly clustered, illustrating the relatively low genetic diversity present in cultivated soybean. AFLP analysis of four soybean near-isogenic lines (NILs) identified three AFLP markers putatively linked to a virus resistance gene from two sources. The capacity of AFLP analysis to detect thousands of independent genetic loci with minimal cost and time requirements makes them an ideal marker for a wide array of genetic investigations.
Studies of diversity permit conclusions about the genetic structure of species. Systematically developed core collections are a particularly good tool for such purposes. The objective of this study was to evaluate the genetic structure between and within gene pools of a core collection of wild Phaseolus vulgaris L. using the technique of amplified restriction polymorphism (AFLP) analysis. DNA fingerprint patterns of 114 genotypes were analyzed by the NTSYS program. Major groups or gene pools were recognized in Mesoamerica, Colombia, the northern Andes of Ecuador and northern Peru, and the southern Andes, although the separation among gene pools was not wide. A unique segment of germplasm from northern Peru showed greater genetic distance than the other gene pools. Although most Mexican wild beans formed a rather homogenous group, a small number of accessions clustered with the Andean gene pool, and accessions from Guatemala tended to cluster apart from Mexican accessions. In the southern Andean gene pool, more discrete groups were formed which were associated with certain regions such as Argentina, possibly reflecting greater geographic and genetic isolation. Colombian accessions appeared to be highly introgressed with germplasm from other areas. AFLP analysis produced a large amount of data in a short period, thus permitting greater insights into the genetic structure of wild beans than had been possible with other methods of analysis.
Methods for measuring the degree of agreement between dendrograms (which correspond uniquely to hierarchic nested classifications) are discussed. While some methods take the relative levels of the subsets into account, most presently available methods do not. A previously proposed consensus index is shown to have a maximum possible value that is (for a given number of objects) a function of the balance (the degree to which the subsets of each set are equal in size) of the consensus of the two classifications being compared. A new consensus index is proposed that does not have this defect. In addition, a new index is proposed that is the proportion of all possible classifications that contain the sets that are in the consensus of the two dendrograms being compared. A table giving the numbers of possible bifurcating dendrograms (up to t=100 objects) is furnished to assist in the computation of this latter index.
The assumptions involved in maximum likelihood estimation of evolutionary trees from quantitative character data are described. A strict maximum likelihood method applied to the case of two populations encounters singularities in the likelihood surface, and even when restrictions are placed on the parameters to avoid this the resulting estimate is converged to the wrong value as more characters are considered. These problems arise because new nuisance parameters are introduced every time a new character is added. If the data are assumed to consist only of the differences between population phenotypes, and a maximum likelihood solution based on this transformed data is found, this restricted maximum likelihood (REML) method behaves well. Two computational techniques, the pruning algorithm and the pulley principle, are described which allow rapid computation of the restricted likelihood. They allow construction of an iterative procedure for finding the maximum of the restricted likelihood within a given tree topology. Combined with an algorithm for searching among similar tree topologies, this allows construction of a computer program to find the REML estimate of the tree. The fact that the REML estimate is a maximum likelihood estimate obtained from transformed data allows use of likelihood-ratio testing of evolutionary hypotheses. Constancy of evolutionary rate per unit time can be tested, but it appears that discrimination between gradualist and punctuated-equilibrium hypotheses will require fossil data.
AFLP marker technology allows efficient DNA fingerprinting and the analysis of large numbers of polymorphic restriction fragments on polyacrylamide gels. Using the doubled haploids from the F1 of the cross Proctor Nudinka, 118 AFLP markers were mapped onto a barley (Hordeum vulgare L.) RFLP map, also including five microsatellite and four protein marker loci. The AFLP markers mapped to all parts of the barley chromosomes and filled in the gaps on barley chromosomes 2L, 4L and 6 in which no RFLP loci had been mapped. Interestingly, the AFLP markers seldom interrupted RFLP clusters, but grouped next to them. The combined map covers 1873 cM, with a total of 282 markers. The merging of AFLP and RFLP markers increased the total map length; 402 cM were added to the map at the tips of chromosomes or in regions corresponding to earlier gaps. Another 375 cM resulted from mapping AFLP markers near to RFLP clusters or in between non-clustered RFLP markers.
It is not unusual for several classifications to be given for the same collection of objects. We present a method, called
majority rule, which can be used to define a consensus of these classifications. We also discuss some mathematical properties
of this consensus tree.
A measure of genetic distance (D) based on the identity of genes between populations is formulated. It is defined as D = -logeI, where I is the normalized identity of genes between two populations. This genetic distance measures the accumulated allele differences per locus. If the rate of gene substitution per year is constant, it is linearly related to the divergence time between populations under sexual isolation. It is also linearly related to geographical distance or area in some migration models. Since D is a measure of the accumulated number of codon differences per locus, it can also be estimated from data on amino acid sequences in proteins even for a distantly related species. Thus, if enough data are available, genetic distance between any pair of organisms can be measured in terms of D. This measure is applicable to any kind of organism without regard to ploidy or mating scheme.
A method based on mutation distances as estimated from cytochrome c sequences is of general applicability.
If we accept the evidence at face value, we are led to conclude that emmer was probably domesticated in the upper Jordan watershed and that einkorn was domesticated in southeast Turkey. Barley could have been domesticated almost anywhere within the arc bordering the fertile crescent. All three cereals may well have been harvested in the wild state throughout their regions of adaptation long before actual farming began. The primary habitats for barley, however, are not the same as those for the wheats. Wild barley is more xerophytic and extends farther downslope and into the steppes and deserts along the wadis. It seems likely that, while all three early cereals were domesticated within an are flanking the fertile crescent, each was domesticated in a different subregion of the zone.
Lest anyone should be led to think the problem is solved, we wish to close with a caveat. Domestication may not have taken place where the wild cereals were most abundant. Why should anyone cultivate a cereal where natural stands are as dense as a cultivated field? If wild cereal grasses can be harvested in unlimited quantities, why should anyone bother to till the soil and plant the seed? We suspect that we shall find, when the full story is unfolded, that here and there harvesting of wild cereals lingered on long after some people had learned to farm, and that farming itself may have originated in areas adjacent to, rather than in, the regions of greatest abundance of wild cereals.
We need far more specific information on the climate during incipient domestication and many more carefully conducted excavations of sites in the appropriate time range. The problem is far from solved, but some knowledge of the present distribution of the wild forms should be helpful.
Systematik Geschichte und Verwendung
- E Schiemann Weizen Roggen Gerste
- M Nei
- Nei M.
) reported that in the Konya plain of Turkey a wild T. m. boeoticum population exists outside of the Fertile Crescent. Two T. m. boeoticum lines sampled at a distance of 50 km from this site were not closely related to cultivated einkorn
- Samuel Nesbitt
Trees (16) were generated as follows: NEI 72 NEI-UB and Rogers-W with NJ and FITCH; average euclidean and euclidean squared distances with FITCH. PHYLIP (Phylogeny Inference Package) was used in computing trees and for the REML clustering method
- Rogers-W From
Harris in Foraging and Farming. The Evolution of Plant Exploitation
- G C Hillman
- S M Colledge