Fact-checked by Grok 2 weeks ago

Founder crops

The founder crops, also known as the Neolithic founder crops or primary domesticates, comprise eight species of plants that were domesticated by early farming communities in the Fertile Crescent of Southwest Asia during the Neolithic Revolution approximately 10,000–11,000 years ago. These include three cereals—einkorn wheat (Triticum monococcum), emmer wheat (T. turgidum subsp. dicoccum), and barley (Hordeum vulgare)—four pulses—lentil (Lens culinaris), pea (Pisum sativum), chickpea (Cicer arietinum), and bitter vetch (Vicia ervilia)—and one oil and fiber crop, flax (Linum usitatissimum). All are annual, self-pollinating plants native to the region, with domestication traits such as non-shattering seed heads in cereals and indehiscent pods in pulses emerging through simple genetic mutations. Their during the period (ca. 11,600–8,500 years BP) represented a pivotal shift from to intensive , enabling sedentary settlements and population growth across the and beyond. Archaeological evidence from sites like , Cafer Höyük, and indicates that began with and lentils around 11,600–10,700 calibrated years BP, followed by full of the package by 10,700–9,600 calibrated years BP, with these crops forming the dietary staples that supported early societies. While the traditional list of eight has been foundational to understanding the origins of farming, recent analyses suggest that additional species, such as faba bean and certain wheats, may have played complementary roles in this agricultural transition, broadening the scope of early crop diversity.

Introduction

Definition

The term "founder crops" refers to the eight plant species that were the first to be domesticated in the ancient Near East, forming the basis of early agriculture. This concept was coined by botanists Daniel Zohary and Maria Hopf in their 1988 book Domestication of Plants in the Old World: The Origin and Spread of Domesticated Plants in Southwest Asia, Europe, and the Mediterranean Basin. The core group consists of three cereals—einkorn wheat (Triticum monococcum), emmer wheat (T. turgidum subsp. dicoccum), and barley (Hordeum vulgare)—four pulses (legumes)—lentil (Lens culinaris), pea (Pisum sativum), chickpea (Cicer arietinum), and bitter vetch (Vicia ervilia)—and the fiber crop flax (Linum usitatissimum). This traditional list of eight species forms the core of early agriculture, though recent research suggests additional plants, such as the faba bean (Vicia faba), may have played complementary roles in the Neolithic founder package. These crops were primarily domesticated in Southwest Asia, within the region known as the , which spans modern-day southeastern , , , , , , and northern . Initial domestication occurred during the period (ca. 11,600–8500 years BP, corresponding to 9600–6500 BCE), marking a pivotal shift from societies to sedentary farming communities. As foundational elements of the —the broader transition to in the region—these founder crops enabled the development of surplus food production and supported the growth of early agricultural economies across .

Historical Significance

The of founder crops played a pivotal role in the , marking a profound shift from to systematic in the around 10,000 BCE. This transition enabled reliable food production, which supported and facilitated population growth by providing a stable caloric base that exceeded what mobile could sustain. In turn, these developments laid the groundwork for the emergence of early civilizations, as settled communities could invest labor in non-subsistence activities like tool-making and trade. The economic and social impacts of founder crop cultivation were transformative, fostering surplus-based economies that allowed for and redistribution within communities. This surplus underpinned the growth of proto-urban villages, such as in the during the 9th–8th millennia BCE and in from the 8th millennium BCE, where dense populations exceeding 1,000 inhabitants relied on to maintain social complexity and ceremonial structures. By 7000 BCE, these agricultural practices had spread westward into via and eastward into , carried by migrating farmers who adapted the crops to new environments and integrated them with local resources. Archaeological evidence underscores the deep roots of this process, with pre-domestication harvesting of wild cereals documented at Ohalo II in the around 23,000 years ago, where fisher-hunter-gatherers used composite sickles to process grains like and progenitors. This intensive exploitation of wild stands preceded the full of founder crops by several millennia, culminating in morphologically distinct domesticated forms by approximately 10,000 BCE across the . Such evidence highlights a gradual intensification of plant use that bridged foraging and farming. The long-term legacy of founder crops extends to their role as the foundational package for subsequent agricultural diversifications, influencing the development of global food systems through hybridizations and . Descendants of and , in particular, remain major staples, contributing significantly to modern human intake—wheat alone accounts for about 20% of global caloric supply, with regional variations reaching up to 50% in wheat-dependent areas. This enduring impact underscores how these early domestications shaped dietary patterns, economic structures, and worldwide.

The Core Founder Crops

Cereals

The cereal founder crops of the in the consist of (Triticum monococcum), emmer wheat (Triticum turgidum subsp. dicoccum), and (Hordeum vulgare), which were among the first plants domesticated by early farmers around 10,000 years ago. These grasses provided reliable sources that supported the transition from societies to settled , forming the staple basis of early diets in the region. Einkorn wheat, derived from the wild ancestor Triticum boeoticum, is a diploid with 2n=14 chromosomes and features small, hulled grains that were primarily used in early for making and . Domestication selected for non-shattering heads, facilitating easier harvesting compared to the brittle rachis of its wild . Emmer wheat originated from the wild tetraploid species Triticum dicoccoides (2n=28) and produces hulled grains that require additional to separate from the glumes, a labor-intensive process in early farming. It served as a key ingredient for early flatbreads and acted as a direct progenitor to modern durum wheat (Triticum turgidum subsp. durum), influencing subsequent breeding. Barley, with its wild ancestor Hordeum spontaneum, developed both hulled and naked (hull-less) varieties during , offering versatility in processing. Known for its , it was utilized in early societies for , , and as animal fodder, with two-row and six-row forms differing in kernel arrangement and yield characteristics. As members of the family, these cereals share high yield potential in the nutrient-rich soils of the , where their wild progenitors demonstrated superior biomass and seed production compared to other regional grasses. Nutritionally, they provided essential energy, with einkorn containing about 12–18% protein by weight (varying by cultivar and conditions), higher than many modern varieties.

Legumes

The legumes among the founder crops—lentil (Lens culinaris), (Pisum sativum), (Cicer arietinum), and bitter vetch ()—belong to the family and were domesticated in the alongside cereals, providing essential protein-rich complements to carbohydrate-heavy diets. These pulses share key biological traits, including symbiotic with rhizobial in root nodules, which enhances by converting atmospheric into usable forms. As a result, they were often grown in rotation with cereals to maintain and sustain long-term cultivation without synthetic fertilizers. When combined with cereals, legumes form complete proteins by supplying complementary , supporting balanced in early agricultural societies. Lentil originated from the wild progenitor Lens orientalis and features small, flat seeds measuring about 2.5–3.0 mm in the wild form, increasing slightly to up to 4.2 mm in domesticated varieties. With approximately 25% protein content, lentils served as a vital substitute in early diets, commonly prepared in soups and stews for their nutritional density. Among the pulses, lentil is considered the earliest to be domesticated, dating back to around 11,000 BCE in the . Pea derives from the wild Pisum sativum subsp. elatius, growing as vines that produce round seeds of 3–4 mm in wild forms, domesticated for larger 6–8 mm seeds and non-dormant pods that retain seeds for easier harvesting. Containing 22–25% protein, peas were a key protein source in diets, valued for their digestibility and role in diversifying consumption. Chickpea traces to the wild progenitor , with domesticated seeds reaching up to 6 mm—larger than the wild 3.5 mm—and exhibiting two main types: (small, angular, brown seeds) and kabuli (large, smooth, beige seeds). Known for drought resistance due to deep roots and efficient water use, chickpeas were roasted as snacks or incorporated into stews and curries, providing about 20% protein as a resilient dietary staple. Bitter vetch, with its wild ancestor also Vicia ervilia (showing minimal distinction between wild and cultivated forms), produces seeds that require processing to mitigate their bitter taste from neurotoxic compounds like β-cyanoalanine. Primarily used as a forage crop for due to its hardiness and soil-enriching properties, it was occasionally consumed by humans during famines after by soaking or cooking.

Flax

Flax (Linum usitatissimum) stands as the unique non-cereal, non-legume among the eight core founder crops of Neolithic Southwest Asia, domesticated primarily for its seed oil and stem fibers rather than direct consumption as grain or pulse. Its wild progenitor, Linum bienne, is an annual herb with slender, erect stems reaching up to 60 cm, narrow lanceolate leaves, and delicate blue flowers that bloom in summer. Native to the Mediterranean Basin and Southwest Asia, this wild form produces small seeds in dehiscent capsules, which shatter to disperse naturally. Domestication, occurring around 10,000 years ago in the Fertile Crescent, selected for non-shattering capsules and larger seeds (up to 4 mm), facilitating easier harvest while preserving the plant's annual lifecycle and floral traits. The plant's dual utility is evident in its seeds, which yield rich in omega-3 fatty acids—specifically alpha-linolenic acid () at 50–60% of total oil content—making it a prized resource for and . These seeds provided an early of essential ALA, supporting dietary health in ancient communities, while the extracted oil fueled lamps, flavored foods, and served medicinal roles, such as treating ailments and digestive issues. Complementing this, flax stems, measuring 80–120 cm in fiber varieties, undergo —exposure to moisture to degrade pectins and separate bast fibers—yielding long, flexible threads for production. Archaeological evidence confirms these as the earliest domesticated textiles, dating to 9,000 years ago in the . The fibers' exceptional tensile strength, two to three times that of , ensured durable fabrics resistant to wear. Flax favors cool, moist temperate climates with well-drained loamy soils and moderate rainfall (400–600 mm annually), allowing integration with cereals in mixed cropping systems to maximize field efficiency. Grown as a short-season (90–110 days to maturity), it produced oil for versatile applications in , cooking, and healing salves, alongside fibers that bolstered early economies. This combination of nutritional (ALA-rich seeds) and industrial (superior-strength fibers) values distinguished flax, enriching beyond the sustenance focus of other founder crops.

Domestication

Origins and Evidence

The domestication of the founder crops, comprising eight key plant species including , emmer wheat, , lentils, peas, chickpeas, bitter vetch, and , primarily originated in the region of Southwest Asia. Archaeological evidence points to initial cultivation and early processes centered in this area, spanning modern-day southeastern , , , , , and northern . Pre-domestication cultivation of wild cereals and legumes began around 10,000–8,500 BCE during the (PPNA) period, as indicated by increased densities of wild progenitor remains at sites like Mureybet and , suggesting systematic human management of plant populations prior to full . Key evidence for early domestication comes from archaeobotanical remains, including carbonized seeds, rachis fragments, and phytoliths, which reveal shifts from wild to domesticated morphologies such as non-shattering ears in cereals. In southeastern , at Çayönü Tepesi, domesticated-type and emmer wheat appear in Early (EPPNB) contexts dated to approximately 8,500–8,000 BCE, with wild types dominating earlier PPNA layers but transitioning to include a mix of forms by the mid-10th millennium BCE. Similarly, in the , barley domestication is evidenced at Tell Aswad in southern Syria, where about 30% of cereal chaff shows domesticated traits around 8,750–8,250 BCE, marking one of the earliest confirmed instances of non-brittle rachises. , used for and oil, shows early domesticated capsule fragments at (ancient ) in the , dated to 10,250–9,500 cal BP (approximately 8,300–7,550 BCE), representing the oldest secure evidence for its cultivation in the region. By around 7,500 BCE, full of the core founder crops had occurred across multiple sites in the , as seen in the predominance of non-shattering forms and increased seed sizes in assemblages from the PPNB period. This timeline aligns with broader archaeobotanical data from sites like 'Ain Ghazal and Yiftahel, where the founder package appears in settled communities. The debate on multiple centers of highlights a core focus in the northern (southeastern and northern ), with parallel developments in southern zones like the , supported by regional variations in crop appearance and patterns.

Key Changes in Domestication

The domestication of founder crops involved human selection for a suite of traits known as the , which enhanced their utility for while reducing in the wild. These adaptations primarily arose through the preferential harvesting and replanting of seed-bearing individuals, favoring mutations that retained seeds on the until harvest. Key changes included alterations in seed retention, size, , and palatability, often controlled by a small number of recessive genes. In cereals such as (Triticum monococcum), (Triticum dicoccum), and (Hordeum vulgare), a critical was the of a non-shattering rachis, where the brittle structure of wild forms—allowing natural via disarticulation at the rachis nodes—was replaced by a tough rachis that retained grains until manually threshed. This trait is governed by recessive mutations: a single locus (btr) in einkorn, two loci (Btr1 and Btr2) in barley acting complementarily, and two loci in emmer. Accompanying this was a 2–4-fold increase in , from narrower wild kernels to broader, thicker domesticated ones, improving yield and ease of processing. Additionally, reduced emerged, enabling more uniform under conditions. For legumes including peas (Pisum sativum), lentils (Lens culinaris), chickpeas (Cicer arietinum), and bitter vetch (Vicia ervilia), selection targeted pod morphology to prevent explosive dehiscence in wild forms, which scatters seeds. Domesticated varieties developed larger, non-dehiscent pods through single recessive mutations, such as the pod constriction gene in peas, resulting in higher seed retention and yield. Seed size increased notably, for example, from 3–4 mm in wild peas to 6–8 mm in domesticated forms, alongside smoother seed coats that reduced toxicity—particularly in bitter vetch, where levels of the anti-nutritional compound canavanine declined, making it suitable for human and animal consumption. These changes collectively boosted harvestable seed yield by retaining pods intact during reaping. In flax (Linum usitatissimum), domestication shifted from wild biennial forms to annual varieties with elongated capsules that were less prone to splitting, controlled by a single recessive mutation, facilitating easier seed and fiber harvest. Selection for fiber use produced taller stems with finer, longer bast fibers, while oil varieties emphasized larger seeds (up to 4.8 mm from wild sizes) with altered fatty acid composition, increasing linoleic and linolenic acid content for edible oil. These dual-purpose adaptations reflect early human preferences for both industrial and nutritional uses. Genetically, these traits exemplify the , where parallel selective pressures across species led to convergent adaptations despite diverse wild progenitors. In wheats, played a role, with arising from hybridization of diploid progenitors followed by chromosome doubling, buffering deleterious mutations and accelerating trait fixation. Under cultivation, key domestication alleles fixed rapidly—estimated at 10–20 generations for traits like non-shattering—due to strong selection via repeated sowing of harvested seeds, which imposed annual evolutionary rates of 0.1–1% for quantitative traits like seed size. This process transformed wild progenitors into obligate cultivars reliant on intervention.

Cultivation and Spread

Early Cultivation Methods

The transition to in the involved a pre- phase of intensive gathering and management of wild stands of founder crops, such as , emmer wheat, , lentils, peas, chickpeas, bitter vetch, and , which lasted approximately 1,000 years before the establishment of full farming practices around 10,500–9,500 cal BP. During this period, known as low-level food production, early foragers engaged in activities like vegetation clearance, controlled burning, and broadcasting seeds on disturbed lands, leading to unintentional selection for traits favorable to human harvesting, such as non-shattering rachises in cereals. These practices, evident at (PPNA) sites like I and Jerf el-Ahmar, gradually intensified resource exploitation without immediate full domestication. Early cultivation techniques relied on simple stone tools and dry farming methods suited to the region's variable rainfall and alluvial soils in river valleys like the and . Sickles with embedded flint blades were used to harvest wild stands by cutting brittle ears, as seen in PPNA assemblages from Netiv Hagdud and , while stone mortars and grinding slabs processed grains post-harvest. Dry farming predominated on fertile alluvial deposits, with fields prepared through hoeing or digging sticks; crop rotation involving fallowing periods allowed in rain-fed areas receiving 200–500 mm annual precipitation, as inferred from multi-year occupation layers at sites like . In semi-arid zones, supplemental drew from seasonal wadis, channeling flash floods to moisten fields, particularly in early Mesopotamian settlements such as Choga Mami around 7,950–7,550 cal . Agronomic practices emphasized sustainability and efficiency in mixed systems. Seeds were broadcast by hand across prepared plots, a method facilitating the cultivation of cereals alongside legumes like lentils and peas, which naturally fixed nitrogen to enhance soil fertility without deliberate fertilization. Storage occurred in subterranean pits lined with plaster or simple above-ground granaries, preserving surpluses as evidenced by large lentil hoards (up to 1.4 million seeds) at Yiftah'el and barley caches at Tell Aswad. Yields for early wheat varieties were modest, typically ranging from 500–1,000 kg/ha under these rain-dependent conditions, reflecting the challenges of nascent agriculture. Challenges included weed proliferation and pest pressures, managed through rudimentary means. Rye often appeared as a contaminant in cereal fields, thriving in disturbed soils at sites like Can Hasan III, requiring manual weeding or selective harvesting. Pest control involved periodic field burning to clear residues and reduce insect populations, a practice documented in charred plant remains and settlement fire layers from PPNA contexts. These methods, while effective for small-scale production, limited scalability until later innovations in the Pottery Neolithic.

Geographical Spread

The founder crops, originating in the , initially expanded to adjacent regions through and trade networks. By approximately 9000 BCE, domestication of emmer wheat, , and had begun in the , with cultivation spreading northward to by 8000 BCE and eastward into around the same period. From the , these cereals and like lentils reached by around 8000 BCE, facilitated by seasonal that supported irrigation-based farming. The diffusion continued westward into , where and arrived in by 7000 BCE via maritime routes from and the Aegean. By 5500 BCE, these crops had reached through the Linearbandkeramik culture, whose farmers practiced on soils, introducing mixed farming systems with cereals and . , valued for its fibers in production, accompanied this spread, with evidence of processing in European settlements by the 5th–4th millennium BCE. In , lentils diffused to sites like in the Indus region by around 6000–5000 BCE, while chickpeas reached the Indus Valley around 4000 BCE, integrating into local agricultural systems alongside native millets during the Early Harappan phase (c. 3300–2600 BCE). Peas reached via early trade routes during the Western around the 2nd century BCE, where they hybridized with indigenous varieties and adapted to temperate climates. Following European exploration after 1492 CE, founder crops such as and were introduced to the , where they became staples in colonial agriculture, particularly in and the . Today, these crops are cultivated in over 100 countries, with alone covering approximately 220 million hectares globally as of , underscoring their enduring adaptability. The geographical spread of founder crops was driven primarily by , including the dispersal of farmers from the , which carried seeds and knowledge across continents. Climate suitability played a key role, as temperate zones in and favored cereal growth, while hybridization with local wild relatives enhanced resilience and yield in new environments.

Additional and Proposed Founder Crops

Other Domesticated Plants

(Secale cereale) originated from the wild progenitor Secale montanum in the and is considered a secondary or accidental domesticate, emerging as a weed in early and fields rather than through intentional selection. Unlike the core founder crops, rye was cultivated as early as the (ca. 11,600–10,700 years ago), but full likely occurred later, around 6,000–3,500 years ago during the or earlier, in or the broader , where its tolerance for poor, marginal soils allowed it to persist and eventually be cultivated for grain and production. This opportunistic adaptation made rye valuable in regions unsuitable for , contributing to its spread across by the . Figs (Ficus carica) represent one of the earliest tree crops domesticated in the , with evidence of parthenocarpic (seedless) varieties—selected for their larger, more edible fruits—dating to the period at the site of Gilgal I in the Lower . These cultivated figs, derived from wild populations, were stored in large quantities around 11,400–11,200 years ago, predating the domestication of the core and founder crops and suggesting figs as a pioneering that supported year-round food availability through drying and . Almonds (Prunus dulcis) were domesticated through selection for sweet, non-bitter kernels from wild bitter varieties native to the , with early archaeological evidence of use appearing in the Mesolithic layers of Franchthi Cave in around 7,000 BCE, though full domestication occurred later, approximately 6,300–5,300 years ago in the . This process transformed almonds from a sporadically gathered wild resource into a reliable crop, valued for its nutritional density and portability. Other notable Near Eastern domestics from the same era include grapes (Vitis vinifera), domesticated around 8,000–6,000 years ago in the Transcaucasus and Near East for fruit and wine production, olives (Olea europaea), cultivated starting in the Levant by 6,000–5,000 BCE for oil extraction, and dates (Phoenix dactylifera), originating approximately 7,000 years ago in the Middle East for their sweet, storable fruits. These fruit and tree crops complemented the core founder package by providing diverse, perennial resources that enhanced dietary variety, offered off-season nutrition through preservation, and supported emerging economies via trade in oils, wines, and dried goods.

Debates on the List

The concept of founder crops, as originally defined by Zohary and Hopf in 1988, encompasses eight plant species— (Triticum monococcum), emmer wheat (Triticum turgidum subsp. dicoccum), (Hordeum vulgare), (Lens culinaris), (Pisum sativum), (Cicer arietinum), bitter vetch (Vicia ervilia), and (Linum usitatissimum)—that formed the basis of early agriculture in southwest . This canonical list has faced scholarly scrutiny, with proposals to expand it based on emerging archaeobotanical suggesting additional early domesticates or cultivated species. Archaeobotanical , including work by George Willcox on pre-domestic cultivation, supports including (Secale cereale) due to its presence in Early sites along the Middle Euphrates, where it appears alongside the core cereals in processed assemblages, indicating intentional cultivation. Similarly, (Pistacia spp.) has been proposed as a candidate for inclusion, given its exploitation in Pre-Pottery contexts across the and , though it is often classified as a gathered resource rather than a fully domesticated founder due to lacking clear morphological changes. More recent findings, such as analysis identifying a "new wheat" variant related to Triticum timopheevii in early sites, further challenge the original list by highlighting overlooked cereal diversity in the domestication package. Faba bean (Vicia faba) has also been proposed as an additional founder crop, with of by 10,700–10,200 calibrated years BP and its translocation to by 8,800 years BP, complementing the pulse component of early agriculture. Central to these debates are the criteria for "founder" status, which require a species to demonstrate early cultivation, integration into the Neolithic crop package, and foundational economic roles in sustaining sedentary communities. Scholars argue that these standards are inconsistently applied, as evidenced by ongoing discussions about figs (Ficus carica), which genetic and archaeobotanical data suggest were domesticated around 11,400 years ago in the Jordan Valley—predating cereal cultivation by millennia through parthenocarpic mutants propagated via cuttings. However, critics contend that early fig remains may represent selective gathering of naturally occurring parthenocarpic variants rather than deliberate domestication, complicating their classification as pre-cereal founders. Later crops like oats (Avena spp.) are routinely excluded because they emerged as secondary domesticates, likely as weeds in primary cereal fields, rather than integral to the initial Neolithic package. Regional variations across the underscore the list's limitations, with distinct crop assemblages in northern and southern zones reflecting local ecologies and practices. In southern areas like the , pulses such as lentils and peas dominate early assemblages, comprising a larger proportion of the diet than in northern sites, where cereals like prevail. These differences suggest that the "founder package" was not uniform but adapted regionally, challenging the notion of a singular, monolithic origin. Recent genetic studies post-2018 have intensified these debates for specific crops, revealing evidence of multiple independent events for , with distinct wild progenitor populations contributing to cultivated lineages across the and beyond, as shown by analyses indicating polyphyletic origins rather than a single event. Critiques also highlight potential biases in the traditional list, which may prioritize a "core" set of species from central sites, underrepresenting peripheral or non-cereal contributions from broader southwest Asian contexts.

Modern Relevance

Contemporary Uses

The founder crops, through their modern descendants, remain staples in global agriculture, with , , and among the most widely cultivated. In the 2024/25 marketing year, global reached approximately 793 million metric tons, making it one of the world's top crops essential for . Barley stood at about 143 million metric tons in the same period, primarily used for and . output expanded to roughly 7.6 million metric tons, with as a leading producer at around 1.8 million metric tons, driven by demand for pulse-based diets. These crops serve diverse contemporary roles in human diets and industries. Wheat descendants form the basis for , , and baked goods worldwide, providing caloric and nutritional foundations in staple foods. Barley is key in production and feed, while also contributing to human consumption in porridges and health foods. Legumes like lentils and chickpeas offer plant-based proteins, with chickpeas prominently featured in products such as and , supporting vegetarian and vegan diets. Flaxseeds are valued for their omega-3 fatty acids in supplements and functional foods, alongside flax fibers for textiles and industrial applications. Regional production hotspots reflect both historical legacies and modern adaptations. In the , traditional diets continue to emphasize , , lentils, and chickpeas in dishes like and lentil soups, sustaining cultural cuisines. and dominate wheat exports, with the producing over 122 million tons and contributing significantly to global trade. In , is cultivated for uses akin to teff, including in flatbreads and as a resilient highland crop for food and fodder. Despite their importance, founder crop cultivation faces challenges from and agricultural shifts. Droughts, intensified by , can cause chickpea yield losses of up to 50% in vulnerable regions, prompting calls for resilient varieties. The widespread adoption of high-yield hybrids has diminished the cultivation of varieties derived from original founder crops, raising concerns for and in breeding programs.

Genetic Research and Conservation

Genetic research on founder crops has advanced significantly through whole-genome sequencing efforts, providing detailed insights into their evolutionary history and genetic architecture. The first for ( vulgare), a key founder crop, was assembled in 2012, integrating physical, genetic, and functional sequence data to map the gene space across its approximately 5 Gb genome. This assembly has been iteratively improved, with high-quality chromosome-scale assemblies of wild genomes published in 2023, enhancing resolution for and mapping. Similarly, seminal work has identified key genes, such as the Q gene in (Triticum spp.), which encodes an AP2-like that suppresses spike shattering, a critical for non-dispersal in cultivated varieties. Current research emphasizes developing climate-resilient varieties using genomic tools, particularly in response to environmental stresses affecting founder crops. For instance, CRISPR-Cas9 editing has been applied to enhance in like lentils (Lens culinaris) by targeting involved in water use efficiency and stress response, building on broader advances in editing for resistance in crops. hotspots in the serve as focal points for these studies, where genebanks collect and characterize wild relatives to identify alleles for traits such as heat and drought tolerance. Conservation efforts prioritize ex situ preservation to safeguard the of founder crops against loss from and intensive agriculture. The Center for Agricultural Research in the Dry Areas (ICARDA), a center, maintains over 152,000 accessions of cereals, legumes, and forages from the , including wild progenitors of founder crops like , , and chickpeas, facilitating breeding and research. Global seed banks, such as the , store duplicate samples of like (Triticum monococcum), ensuring long-term viability for restoration if needed. Additionally, initiatives promote the revival of such as (Triticum dicoccum) in systems, leveraging their superior nutritional profiles—higher in protein, fiber, and micronutrients compared to modern —for sustainable, low-input production. Post-2020 findings have refined our understanding of founder crop origins, revealing multiple domestication and diversification events. Genomic analyses of chickpeas (Cicer arietinum) indicate at least two independent post-domestication origins for the kabuli type, driven by selection on a gene during migration along historical trade routes from the . These crops also play a pivotal role in , supporting low-input farming through nitrogen-fixing that enhance and reduce reliance on synthetic fertilizers in diverse cropping systems.

References

  1. [1]
    The Neolithic Southwest Asian Founder Crops : Their Biology and ...
    This article reviews the available information on the founder grain crops (einkorn wheat, emmer wheat, barley, lentil, pea, chickpea, and flax) that started ...
  2. [2]
    On the 'lost' crops of the neolithic Near East - PMC - PubMed Central
    Feb 23, 2013 · The claim that the 'classic' eight 'founder crop' package (einkorn wheat, emmer wheat, barley, lentil, pea, chickpea, bitter vetch, and flax) ...
  3. [3]
    Revisiting the concept of the 'Neolithic Founder Crops' in southwest ...
    Apr 8, 2023 · Zohary and Hopf coined the term 'founder crops' to refer to a specific group of eight plants, namely three cereals (einkorn, emmer and ...
  4. [4]
    (PDF) Domestication of Plants in the Old World - The Origin and ...
    founder crops have a rather limited distribution. Wild emmer wheat and wild ... Zohary · Maria Hopf · View full-text. Chapter. Full-text available. Vegetables ...
  5. [5]
    Archaeogenomic analysis of the first steps of Neolithization in ...
    Nov 22, 2017 · The Neolithic transition in west Eurasia occurred in two main steps: the gradual development of sedentism and plant cultivation in the Near East ...
  6. [6]
    Evidence for food storage and predomestication granaries 11,000 ...
    Jul 7, 2009 · Food storage is a vital component in the economic and social package that comprises the Neolithic, contributing to plant domestication, ...
  7. [7]
    Early Neolithic genomes from the eastern Fertile Crescent - Science
    From there, farming spread into surrounding regions, including Anatolia and, later, Europe, southern Asia, and parts of Arabia and North Africa. Whether the ...
  8. [8]
    Composite Sickles and Cereal Harvesting Methods at 23,000-Years ...
    Nov 23, 2016 · Use-wear analysis of five glossed flint blades found at Ohalo II, a 23,000-years-old fisher-hunter-gatherers' camp on the shore of the Sea of ...Missing: ago | Show results with:ago
  9. [9]
    The Origins of Agriculture in the Near East | Current Anthropology
    Agriculture in the Near East arose in the context of broad-based systematic human efforts at modifying local environments and biotic communities.<|control11|><|separator|>
  10. [10]
    The grain quality of wheat wild relatives in the evolutionary context
    Wheat is one of the world's most important crops. It provides 19% of human calorie intake and 21% of protein intake (Shiferaw et al. 2013). Hexaploid bread ...Missing: percentage | Show results with:percentage
  11. [11]
    On the Origin of the Non-brittle Rachis Trait of Domesticated Einkorn ...
    Jan 4, 2018 · Einkorn and emmer wheat together with barley were among the first cereals domesticated by humans more than 10,000 years ago, long before durum ...
  12. [12]
    On the Origin and Domestication History of Barley (Hordeum vulgare)
    Remains of barley (Hordeum vulgare) grains found at archaeological sites in the Fertile Crescent indicate that about 10,000 years ago the crop was domesticated ...
  13. [13]
    [PDF] Rediscovering Ancient Wheats - Cereals & Grains Association
    The first wild ancestors of wheat were diploid wild einkorn (T. boeoticum, genome AmAm) and red wild einkorn (T. urartu, genome AA), both of which are separated ...
  14. [14]
    Einkorn, Emmer, Spelt and Hard Red Spring Wheat - PubMed Central
    Apr 2, 2021 · Einkorn (Triticum monococcum L. ssp. monococcum) is a diploid (2n = 2x = 14) hulled wheat carrying the A genome [2]. Cultivated emmer wheat ...
  15. [15]
    Domestication quantitative trait loci in Triticum dicoccoides ... - PNAS
    Abstract. Wild emmer wheat, Triticum dicoccoides, is the progenitor of modern tetraploid and hexaploid cultivated wheats. Our objective was to map domestication ...
  16. [16]
    Emmer | American Society of Baking
    Emmer is also a tetraploid species, containing 2n=28 chromosomes. Emmer wheat is a potentially valuable genetic resource for providing economically ...
  17. [17]
    Emmer Wheat | Grown in Marin
    On threshing, a hulled wheat spike breaks up into spikelets. These require milling or pounding to release the grains from the glumes. IVFGWheatThresh6. Emmer ...Missing: flatbreads parent
  18. [18]
    Re-discovering ancient wheat varieties as functional foods - PMC
    Most tetraploid wheats (e.g. emmer and durum wheat) are derived from wild emmer, T. dicoccoides. Wild emmer is itself the result of a hybridization between ...Missing: parent | Show results with:parent
  19. [19]
    The Importance of Barley Genetics and Domestication in a Global ...
    (2004) hypothesized that naked barley originated from wild barley directly or it originated from hulled domesticated barley, which is now extinct. High-density ...
  20. [20]
    To Grow, or Not to Grow Row Barley? - Cornell Small Farms
    Jan 13, 2020 · Two-Row, Six-Row ; Greater drought tolerance, which means less chance of increased protein levels due to moisture stress. Lower range of protein ...Missing: bread | Show results with:bread
  21. [21]
    Barley - Syngenta
    Barley is a versatile crop which is used in various foods such as bread, cereals, and soups. One of its major uses is in the production of beer and whiskey and ...Missing: fodder | Show results with:fodder
  22. [22]
    2-Row vs 6-Row - Barley Breeding Program | Montana State University
    Six-row types have a rounded head appearance with all six kernels developing. The tightly packed seeds around the six-row head tend to be smaller.
  23. [23]
    Were Fertile Crescent crop progenitors higher yielding than other ...
    Mar 11, 2015 · A comparison across nine grass species from the Fertile Crescent showed that cereal crop progenitors had the potential to produce a higher ...
  24. [24]
    Rediscovering Ancient Wheats - Cereals & Grains Association
    About 95% of the total wheat produced is common wheat, and about 5% is durum wheat. Spelt, emmer, and einkorn play only minor roles in terms of production and ...Missing: flatbreads parent
  25. [25]
    Advancing Grain Legumes Domestication and Evolution Studies ...
    In the Near East, the Neolithic founder legumes include pea, lentil, bitter vetch (Vicia ervilia), chickpea (Cicer arietinum) and presumably faba bean ...Missing: characteristics | Show results with:characteristics
  26. [26]
    Unraveling Origin, History, Genetics, and Strategies for Accelerated ...
    Legume crops play a significant role in increasing indigenous nitrogen production in addition to meeting demands of human population for protein and energy.
  27. [27]
    [PDF] Legume Crops Phylogeny and Genetic Diversity for Science and ...
    Oct 24, 2014 · This article discusses legume crops' phylogeny and genetic diversity for science and breeding, published in Critical Reviews in Plant Sciences.
  28. [28]
    Polyphenols, Saponins and Phytosterols in Lentils and Their Health ...
    Lentils, like most legumes, have been gaining popularity for their nutritional significance in human diets, with nearly 60% carbohydrate and 25% protein as ...
  29. [29]
    Molecular Evidence for Two Domestication Events in the Pea Crop
    Nov 6, 2018 · Both P. sativum and P. abyssinicum share traits that are typical of domestication, such as non-dormant seeds. Non-dormant seeds were also found ...
  30. [30]
    Genomic Tools in Pea Breeding Programs: Status and Perspectives
    Pea (Pisum sativum L.) is an annual cool-season legume and one of the oldest domesticated crops. Dry pea seeds contain 22–25% protein, complex starch and fiber ...
  31. [31]
    Chickpea (Cicer arietinum L.) Biology and Biotechnology
    Desi seeds are small, colored, and angular in shape, but kabuli seeds are huge, shaped like an owl's head, and beige in color [16]. For trade, seed size and ...Missing: curries | Show results with:curries
  32. [32]
    Chickpea seed types a) desi b) kabuli c) gulabi d) black e) green
    Seeds of desi chickpea are small, angular with rough brown color testas while kabuli types are relatively large, smooth and cream colored testas. In addition a ...
  33. [33]
    Genetic relationships between wild and cultivated Vicia ervilia (L ...
    The wild form of bitter vetch, Vicia ervilia, grows mainly in Turkey. Wild and cultivated V. ervilia have the same chromosome number (2n =14), share the ...
  34. [34]
    Environmental imposition or an ancient farmers' choice? A study of ...
    Sep 27, 2019 · As a result, some species of legumes have been regarded as “inferior” to human consumption. Two known examples are bitter vetch (Vicia ervilia) ...
  35. [35]
    When and where will vetches have an impact as grain legumes?
    Hungarian vetch and Bitter vetch for example are often the legume crops of ... vetches is very small, they have been used frequently as food during famines.
  36. [36]
    Flax domestication processes as inferred from genome-wide SNP data
    Mar 13, 2025 · Flax (Linum usitatissimum L.) is known to be one of the Neolithic Southwest Asian founder crops domesticated for oil and fiber uses in the Near ...
  37. [37]
    Genetic Relationships of Cultivated Flax and Its Wild Progenitor as ...
    Jun 3, 2024 · Flax (Linum usitatissimum L.) was the first founder crop domesticated for oil and fiber uses in Near Eastern agriculture roughly 8000 years ago ...
  38. [38]
    Flax and flaxseed oil: an ancient medicine & modern functional food
    Flax fibers are amongst the oldest fiber crops in the world. The use of flax for the production of linen goes back at least to ancient Egyptian times. Flax ...<|control11|><|separator|>
  39. [39]
    Linseed Oil - an overview | ScienceDirect Topics
    It contains 9.0 g SFA/100 g, 18.4 g MUFA/100 g, and 67.8 g PUFA/100 g of which 53.4 g is ALA. This unusually high content of ALA makes this oil oxidize rapidly, ...
  40. [40]
    [PDF] Flax Fiber Offers Cotton Cool Comfort - USDA
    Flax is two to three times stronger than cotton, making it one of the ... The ARS scientists are now entering a research agreement with. Clemson University ...
  41. [41]
    Regional diversity on the timing for the initial appearance of cereal ...
    Dec 5, 2016 · In Turkey, the earliest evidence includes domesticated-type emmer, barley, and one-grained einkorn, which are found around 10.3–10.2 ka Cal BP ...
  42. [42]
    Pathways to plant domestication in Southeast Anatolia based on ...
    Jan 22, 2021 · This suggests that at least some of the Çayönü Phase Ib charred Triticum grains can be attributed to domesticated-type einkorn and emmer wheat.
  43. [43]
    On the Origin of the Non-brittle Rachis Trait of Domesticated Einkorn ...
    Jan 3, 2018 · In barley, Btr1 and Btr2 act as complementary genes, with loss-of-function mutations in either of them generating the non-brittle rachis ...
  44. [44]
    Grain Disarticulation in Wild Wheat and Barley - Oxford Academic
    Jun 28, 2022 · In barley, independent mutations in two genes responsible for grain disarticulation, called Non-brittle rachis 1 and Non-brittle rachis 2 (btr1 ...
  45. [45]
    (PDF) Reconsidering Domestication Of Legumes Versus Cereals In ...
    Aug 6, 2025 · We explain the biological aspects of Near Eastern plant domestication by a comparative analysis of legume vs. cereal crop evolution.
  46. [46]
    Advancing Grain Legumes Domestication and Evolution Studies ...
    Pod shattering. The loss of pod dehiscence or shattering is one of the key traits that humans selected for in domesticated grain legumes. Pod shattering is ...Genetic Mapping Of... · Pod Shattering · Seed Dormancy
  47. [47]
    Locus-specific view of flax domestication history - PMC
    A subsequent flax domestication process occurred that probably involved multiple domestications and includes lineages that contain oil, fiber, and winter ...
  48. [48]
    Genomic Comparison and Population Diversity Analysis Provide ...
    Apr 24, 2020 · That is, oil-use flax is shorter, has more branches, and produces larger seeds that contain ∼40% oil, and fiber-use flax is comparatively taller ...
  49. [49]
    The Domestication Syndrome Genes Responsible for the Major ...
    Most domestication genes responsible for the major morphological change in cereal crops are transcriptional regulators, suggesting that this class of genes ...Missing: generations fixation
  50. [50]
    Dispersed emergence and protracted domestication of polyploid ...
    Jul 6, 2022 · We reveal that domesticated polyploid wheat emerged from the admixture of six founder wild emmer lineages, which contributed the foundation of ancestral ...
  51. [51]
    Patterns and processes in crop domestication: an historical review ...
    Aug 13, 2012 · The domestication syndrome may evolve over thousands of generations, as desirable traits are selected for in the agricultural environment and ...Missing: founder | Show results with:founder
  52. [52]
    The Neolithic Southwest Asian Founder Crops : Their Biology and ...
    Frequently it grows side by side with bitter vetch (Vicia ervilia). As argued by Zohary (1999), the rich chromosomal polymorphism found in the wild ...
  53. [53]
    [PDF] Plant domestication in the Neolithic Near East
    One of the tenets of the pre- domestication cultivation concept is its gradual unfolding via increased human intervention in both the environment and the life.
  54. [54]
    (PDF) Recent lessons from Near Eastern archaeobotany: wild cereal ...
    PDF | On Jan 1, 2008, D.Q.Fuller published Recent lessons from Near Eastern archaeobotany: wild cereal use, pre-domestication cultivation and tracing ...<|separator|>
  55. [55]
    Agronomic conditions and crop evolution in ancient Near East ...
    May 23, 2014 · Here we investigate the evolution of agronomic conditions in this region by reconstructing cereal kernel weight and using stable carbon and nitrogen isotope ...
  56. [56]
    Agronomic conditions and crop evolution in ancient Near East ...
    May 23, 2014 · Following the aforementioned environmental and genetic changes, yield estimates during the Neolithic were in general lower than 0.8 t ha−1, but ...
  57. [57]
    (PDF) Plant domestication in the Neolithic Near East - Academia.edu
    Rather, we contend that the biological idiosyncrasies of the Near Eastern founder crops depict a picture of a knowledge-based and conscious domestication that ...
  58. [58]
    Origins of agriculture - PlantsPeoplePlanet
    Eight Neolithic founder crops, the first known domesticated plants in the world, have been isolated in the Fertile Crescent of southwest Asia, and it was these ...<|separator|>
  59. [59]
    Agriculture in the Fertile Crescent & Mesopotamia
    Mar 22, 2023 · Wheat and goats were domesticated in the Levant by 9000 BCE, followed by peas and lentils in the Fertile Crescent and northern Egypt around 8000 ...<|control11|><|separator|>
  60. [60]
    A Complete History Of Barley: From Ancient Grain To CRISPR ...
    Sep 26, 2025 · Barley cultivation reached Central Europe's loess soils by 4,500 BCE via Linearbandkeramik farmers who cleared forests using slash-and-burn ...Missing: textiles | Show results with:textiles
  61. [61]
    The First Farmers | Europe before Rome - Oxford Academic
    The first farming communities in Europe are known from Greece and the Aegean area and date to around 7000 bc. By 4000 bc, farming had penetrated to the ...
  62. [62]
    Linear Pottery culture | Archaeology News Online Magazine
    The Linear Pottery culture was a Neolithic culture that emerged in Central Europe around 5500 BCE and lasted until approximately 4500 BCE.
  63. [63]
    The History of Neolithic Flax Fiber Processing - ThoughtCo
    May 20, 2019 · Flax was originally domesticated about 4000 years earlier in the Fertile Crescent region, for its oil-rich seeds: the cultivation of the plant ...
  64. [64]
    100 years of Discovery of Harappan Civilisation | Current Affairs
    Oct 17, 2024 · Early Harappan (6000 BCE-2600 BCE) is a formative phase of the ... Main crops: Wheat, Rice, millets, barley, lentil, chickpea and sesame etc.
  65. [65]
    Exchanges of economic plants along the land silk road - PMC
    Tibetan barley, barley, wheat, and jujube were introduced into China before the Silk Road; while mustard, lettuce, buckwheat, chickpea, alfalfa, walnut, ...Missing: lentils | Show results with:lentils
  66. [66]
    Ancient nomads spread earliest domestic grains along Silk Road ...
    Apr 2, 2014 · While these crops have been known to exist much earlier in ancient China and Southwest Asia, finding them intermingled in the Bronze Age burials ...
  67. [67]
    The Columbian Exchange
    When Europeans first touched the shores of the Americas, Old World crops such as wheat, barley, rice, and turnips had not traveled west across the Atlantic, and ...
  68. [68]
    Coming to America: Wheat sailed with Columbus
    Wheat was unknown in the Americas before Columbus. Spaniards brought it to Mexico, and other explorers to the US East Coast. The first Kansas crop was 350 ...
  69. [69]
    [PDF] Impacts of climate change on global wheat production and supply ...
    Feb 12, 2025 · Wheat also has the largest harvested area globally ... 219 million hectares in 126 countries in the last decade, followed only by maize at 193.<|separator|>
  70. [70]
    The Development of Agriculture - National Geographic Education
    May 29, 2025 · The development of agricultural about 12,000 years ago changed the way humans lived. They switched from nomadic hunter-gatherer lifestyles ...
  71. [71]
    Evolutionary history of barley cultivation in Europe revealed by ...
    Nov 2, 2011 · Wild and most cultivated barleys have hulled grains where the outer lemma and inner palea adhere to the pericarp epidermis at maturity. This ...
  72. [72]
    Genetic diversity and relationship between domesticated rye and its ...
    Because of this complex history, rye is considered a secondary domesticate (Preece et al., 2017). Rye belongs to the small genus Secale with only three taxa, ...
  73. [73]
    Population genomic analysis reveals domestication of cultivated rye ...
    Mar 7, 2022 · The geneticist NI Vavilov proposed that cultivated rye had been domesticated from weedy rye, rather than directly from wild species like other crops.
  74. [74]
    Early domesticated fig in the Jordan Valley - PubMed
    Here we report the discovery of nine carbonized fig fruits and hundreds of drupelets stored in Gilgal I, an early Neolithic village, located in the Lower Jordan ...Missing: carica BCE
  75. [75]
    Naturalization of almond trees (Prunus dulcis) in semi-arid regions ...
    The almond tree is probably one of the earliest domesticated trees in the Old World, with archaeological evidence suggesting domestication about 6300–5300 yr.
  76. [76]
    Editorial: Origins and Domestication of the Grape - PMC - NIH
    Archaeological data suggest that domestication of the grapevine began 6,000–8,000 years ago in the Transcaucasian region, between the Black Sea and Iran, from ...
  77. [77]
    Olive domestication and diversification in the Mediterranean Basin
    Nov 24, 2014 · Primary domestication and early uses of the emblematic olive tree: palaeobotanical, historical and molecular evidence from the Middle East.
  78. [78]
    Cross-species hybridization and the origin of North African date palms
    Date palm (Phoenix dactylifera L.) is a major fruit crop of arid regions that were domesticated ∼7,000 y ago in the Near or Middle East. This species is ...
  79. [79]
    https://www.researchgate.net/publication/265281637_Pre-Domestic_Cultivation_during_the_Late_Pleistocene_and_Early_Holocene_in_the_Northern_Levant
  80. [80]
    Targeted resequencing reveals genomic signatures of barley ...
    Mar 12, 2018 · We discovered multiple selective sweeps that occurred on all barley chromosomes during domestication in the background of several ancestral wild ...
  81. [81]
    https://nph.onlinelibrary.wiley.com/doi/10.1111/nph.15077
  82. [82]
    Barley - USDA Foreign Agricultural Service
    In 2024/2025, the top barley producers are the EU (35%), Russia (11%), and Australia (9%). The 2024/2025 production is 143.33 million metric tons.
  83. [83]
    Lentils Market Update - March 2025: Seedea
    Mar 5, 2025 · Global lentil production in the 2024/2025 season is estimated to increase by 15%, reaching approximately 6.6 million tonnes, up from 5.7 million tonnes in the ...Global Lentils Market in 2024 · Lentils Production in Australia
  84. [84]
    Lentil Markets 2024: India's Supply Paradox, Global Risks, and Price ...
    Jun 10, 2025 · Indian Production: Official estimate at 1.8 million tonnes (18 lakh); trade sources see actual output 10-12 lakh tonnes—a significant downward ...
  85. [85]
    Production - Wheat - USDA Foreign Agricultural Service
    10 Year Average MY 2015-2024. 768.31 Million Metric Tons ; 10-Year Compound Average Growth. 0.81% MY 2015-2024 ; 2023/2024. Production. 792.34 Million Metric Tons.
  86. [86]
    Impact of High Temperature and Drought Stresses on Chickpea ...
    Such changes in climate will impact chickpea production and yield and result in grain yield decreases of up to 19% in chickpea [4]. The crop generally ...1. Introduction · 2. Effect Of Drought On... · 4.1. Drought Escape And...
  87. [87]
    A physical, genetic and functional sequence assembly of the barley ...
    Oct 17, 2012 · Here we present an integrated and ordered physical, genetic and functional sequence resource that describes the barley gene-space in a structured whole-genome ...
  88. [88]
    High-quality wild barley genome assemblies and annotation with ...
    Aug 10, 2023 · The draft sequence assembly of barley cultivar (cv.) Morex was reported in 2012, and it was further improved in 2017, especially in the ...
  89. [89]
    Duplication and partitioning in evolution and function of ... - PNAS
    The Q gene encodes an AP2-like transcription factor that played an important role in domestication of polyploid wheat. The chromosome 5A Q alleles (5AQ and ...
  90. [90]
    Genetic Improvement in Leguminous Crops Through Genome Editing
    Genome editing techniques have been successfully used in different legume crops, mainly on model crops and grain legumes such as chickpea, soybean and cowpea.1 Introduction · 2.2 Modern Genome Editing... · 3 Crispr-Mediated Genome...<|separator|>
  91. [91]
    Review of Crop Wild Relative Conservation and Use in West Asia ...
    May 13, 2024 · The 'fertile crescent' is the crescent shaped region to the North and West of the Arabian desert, in current-day Turkey, Syria, Lebanon, Iraq, ...
  92. [92]
    ICARDA genebank
    It contains around 152,224 accessions of major winter cereals, food legumes, forage, and rangeland species drawn from four major Centers of Origin of the Crops ...Missing: founder | Show results with:founder
  93. [93]
    Svalbard Global Seed Vault - Crop Trust
    The Svalbard Global Seed Vault is our insurance against the loss of crop diversity and is the world's largest backup facility for seeds.
  94. [94]
    Ancient Wheats—A Nutritional and Sensory Analysis Review - PMC
    The purpose of this review is to provide a critical evaluation of the nutritional and sensory properties of ancient wheats (spelt, emmer, einkorn, and kamut)
  95. [95]
    Historical Routes for Diversification of Domesticated Chickpea ...
    May 9, 2023 · Abstract. According to archaeological records, chickpea (Cicer arietinum) was first domesticated in the Fertile Crescent about 10000 years ...
  96. [96]
    Keeping the Spotlight on Pulses: “Roots” for Sustainable Agriculture ...
    Feb 10, 2023 · For farmers, pulses are an important crop. When farmers include pulses in multiple cropping systems, they promote sustainable farming practices ...