1 Introduction

Malting Barley, scientifically known as Hordeum vulgare, holds a significant place in the history of agriculture and human civilization. The domestication of barley is believed to have originated in the Fertile Crescent around 10 000 years ago, making it one of the earliest cultivated crops (Badr et al., 2000; Mascher et al., 2016). Archaeological evidence suggests that barley was a staple in Neolithic societies and played a crucial role in the development of early agrarian cultures (Saisho and Purugganan, 2007; Riehl, 2019). The domestication process involved selecting traits that were beneficial for human use, such as non-brittleness of the rachis, which facilitated easier harvesting (Civáň et al., 2021).

The cultural importance of Malting Barley cannot be overstated. It has been used for various purposes, including baking, cooking, and most notably, beer brewing (Wei, 2024). The production of beer dates back to ancient civilizations such as Mesopotamia and Egypt, where it was a significant part of the diet and economy (Riehl, 2019). The versatility of barley, coupled with its adaptability to different climates, has made it a valuable crop across various cultures and regions (Saisho and Purugganan, 2007; Riehl, 2019). In modern times, barley continues to be a critical iingredient of the brewing industry, contributing to the global beer culture.

The evolutionary path of Malting Barley from its ancient domestication to modern global cultivation is a fascinating journey marked by genetic diversity and adaptation. Initially domesticated in the Fertile Crescent, barley spread to different parts of the world through trade and migration (Badr et al., 2000; Morrell and Clegg, 2007). Genetic studies have revealed that barley underwent multiple domestication events, contributing to its genetic diversity (Morrell and Clegg, 2007; Dai et al., 2014). Modern barley cultivars are the result of complex interactions between wild and domesticated varieties, shaped by regional selective pressures and gene flow (Russell et al., 2011; Civáň et al., 2021). Advances in genomic technologies have further illuminated the mosaic origins of cultivated barley, highlighting its evolutionary complexity (Dai et al., 2014; Pankin et al., 2018).

This study provides a comprehensive review of the global spread and brewing culture of Malting Barley, tracing its evolution from ancient domestication to modern agriculture. By examining the botanical traits and cultural diffusion of Malting Barley, it aims to understand the factors that have contributed to its widespread cultivation and cultural significance. The study analyzes genetic evidence of multiple domestication events, explores the cultural impact of Malting Barley, and assesses the current state of barley cultivation worldwide. Through this systematic review, it seeks to shed light on the complex history and enduring legacy of Malting Barley.

2 Botanical Characteristics and Domestication History of Malting Barley

2.1 Origins and early domestication of barley

Barley (Hordeum vulgare) is one of the earliest domesticated crops, with its origins tracing back to the Fertile Crescent around 10 000 years ago. Archaeological evidence indicates that barley was domesticated from its wild relative, Hordeum spontaneum, in this region, which played a crucial role in the transition from hunter-gatherer societies to agrarian communities (Badr et al., 2000; Riehl, 2019). The domestication process involved selecting traits that were beneficial for cultivation and harvesting, such as non-brittle rachis, which allowed grains to remain attached to the plant, facilitating easier collection (Pourkheirandish et al., 2015; Civáň et al., 2021).

Recent genetic studies have provided insights into the complex domestication history of barley. Evidence suggests that barley may have been domesticated more than once, with significant genetic contributions from regions both within and east of the Fertile Crescent. This polyphyletic origin is supported by differences in haplotype frequencies and genomic analyses, indicating that modern cultivated barley has genetic inputs from both the Near East and regions as far east as Tibet (Morrell and Clegg, 2007; Saisho and Purugganan, 2007; Dai et al., 2014). These findings highlight the importance of barley in early agricultural societies and its role in the spread of agrarian culture across Eurasia.

2.2 Evolution of botanical traits in barley

The evolution of barley's botanical traits has been driven by both natural selection and human intervention. Key traits that have influenced its domestication include grain retention, drought resistance, and adaptability to diverse climates. The transition from wild to domesticated barley involved mutations in genes responsible for grain retention, converting the brittle rachis of wild barley into a tough, non-brittle form that facilitated harvesting (Pourkheirandish et al., 2015; Haas et al., 2019). This trait was crucial for the establishment of barley as a staple crop in early agricultural societies.

Barley is known for its remarkable adaptability to various environmental conditions, including arid and saline soils. This adaptability is partly due to its genetic diversity, which has been shaped by multiple domestication events and subsequent selection pressures. Studies have shown that barley landraces from different regions exhibit distinct genetic patterns, reflecting their adaptation to local climates and agricultural practices (Russell et al., 2011; Riehl, 2019). Additionally, barley's ability to thrive in both cool and warm climates has contributed to its widespread cultivation and use in various cultural contexts, from food production to brewing (Riehl, 2019).

2.3 Earliest records of barley in brewing

The use of barley in brewing dates back to ancient civilizations, with some of the earliest records found in Bronze Age Mesopotamia and Egypt. Barley was a key ingredient in the production of beer, which played a significant role in the diet and economy of these societies (Riehl, 2019). The evolution of barley's botanical traits, such as grain retention and adaptability, made it an ideal crop for brewing. The non-brittle rachis allowed for efficient harvesting and processing of grains, while its adaptability ensured a reliable supply of raw material for beer production.

As barley spread across different regions, its use in brewing also expanded. The genetic diversity of barley, resulting from multiple domestication events and regional adaptations, contributed to the development of various brewing traditions and techniques. For instance, the Tibetan hulless barley (Qingke) is believed to have been used in early brewing practices in the region, highlighting the cultural diffusion of barley and its significance in brewing (Dai et al., 2014). The continued selection for traits beneficial to brewing, such as high starch content and favorable fermentation properties, has further cemented barley's role as a cornerstone of beer production throughout history.

3 Spread and Cultural Integration of Malting Barley

3.1 Spread of barley from the fertile crescent to europe

Barley, one of the earliest domesticated crops, originated in the Fertile Crescent around 10 500 years ago and played a crucial role in the spread of agriculture across Europe. Genetic evidence suggests that barley was introduced into Europe through multiple routes, reflecting the complex nature of early agricultural expansion. The Neolithic spread across Europe occurred via two primary routes: one along the Mediterranean coasts and the other following the major river axes. This dual-route hypothesis is supported by DNA analysis, which indicates that domesticated barley in Neolithic Europe falls into three distinct genetic groups, each potentially originating from different centers in southwest Asia (Jones et al., 2013). This genetic diversity underscores the role of barley in facilitating cultural exchanges between early farming communities in Europe and southwest Asia.

Further genetic studies have revealed that barley was domesticated more than once, with a second domestication event occurring east of the Fertile Crescent. This second domestication contributed significantly to the genetic diversity of barley in Central Asia and the Far East, while the Fertile Crescent domestication influenced European and American cultivars (Morrell and Clegg, 2007). The phylogeographic analysis of barley landraces supports the notion of differential migration routes, with European and North African barley types being more closely related to Near Eastern populations, while Asian barley types show distinct genetic patterns (Saisho and Purugganan, 2007). These findings highlight the importance of barley in the cultural diffusion and agricultural development of early European societies.

3.2 Status of Malting Barley in medieval europe

During the medieval period, barley became a staple crop in Europe, significantly influencing the rise of brewing culture. Barley’s adaptability to diverse climates and its resilience in marginal environments made it an ideal crop for medieval European agriculture. By the fifth and fourth millennia BC, barley cultivation had become more common among early European farmers, particularly those in Central and Northern Europe (Jones et al., 2013). This period also saw the development of various barley varieties suited to different growing conditions, further promoting its widespread cultivation.

The brewing of beer, which had ancient origins in societies such as Bronze Age Mesopotamia and Egypt, became a significant cultural and economic activity in medieval Europe. Barley was a key ingredient in beer production, and its cultivation was closely linked to the rise of brewing culture. The use of fermented barley for producing alcoholic beverages is well-documented in archaeological contexts, and by the medieval period, beer had become an important part of the European diet and economy (Riehl, 2019). The spread of barley cultivation and the rise of brewing culture were mutually reinforcing processes, with the demand for beer driving the expansion of barley farming across Europe.

3.3 Global dissemination of Malting Barley to the Americas and Asia

The global dissemination of barley was significantly influenced by colonization and trade, particularly during the age of exploration. Barley, which had already spread across Eurasia by 2000 BC, was introduced to the Americas following European colonization. The introduction of barley to the New World after 1492 CE marked a significant expansion of its cultivation, leading to its global distribution by the 1950s (Riehl, 2019). This period of globalization saw barley being grown in diverse environments, from the high altitudes of the Andes to the varied climates of North America.

In Asia, the spread of barley was facilitated by ancient trade routes such as the Silk Road. Genetic analyses reveal that barley from the Fertile Crescent and a second domestication center in the eastern Iranian Plateau contributed to the genetic diversity of Asian barley populations (Saisho and Purugganan, 2007; Wang et al., 2015). The Silk Road played a crucial role in the exchange of barley genes between Eastern and Western populations, promoting the spread of barley cultivation across Central and East Asia (Wang et al., 2015). Archaeobotanical evidence supports the notion that barley, along with other crops, was part of the trans-Eurasian crop exchange, which facilitated cultural and agricultural interactions between prehistoric farming communities (Jones et al., 2016; Dong et al., 2017). This exchange laid the foundations for the widespread cultivation of barley in Asia, contributing to its status as a globally important crop.

4 Modern Cultivation of Malting Barley and Agricultural Evolution

4.1 Modern breeding and improvement of Malting Barley varieties

The development of Malting Barley varieties through selective breeding and genetic technologies has significantly advanced in recent years. Modern breeding efforts have focused on enhancing genetic diversity and improving agronomic traits to suit various environmental conditions. For instance, a study on Australian barley cultivars revealed that modern varieties exhibit 12% higher genetic diversity than historical ones, with 69 candidate regions within 922 genes identified as being under selection pressure (Hill et al., 2021). This genetic diversity is crucial for adapting to local environments and improving yield stability.

Hill et al. (2021) studied the genetic diversity and population structure of global barley germplasm. Through the division of the population into K=12 groups, the research revealed genetic differentiation of barley germplasm across different geographical regions, growth habits, and row types. Additionally, the neighbor-joining tree of Australian barley varieties further demonstrated significant genetic variation between varieties developed at different historical stages. This study provides foundational data for the genetic improvement of barley, promoting progress in global barley breeding efforts (Figure 1).

Figure 1 Population structure analysis of global barley germplasm (Adapted from Hill et al., 2021) Image caption: (a) Exploration of the optimal number of genetic subgroups (K) in global barley germplasm through Δ cross-validation error and standard error values, indicating that K = 12 is the most probable number of subgroups in this population; (b) Population structure inference for the entire barley panel, assuming 12 subgroups, where each color represents a different subgroup; (c) Population structure distribution of barley based on seven geographical locations; (d) Population structure distribution based on three growth habits; (e) Population structure distribution based on row type (two-row and six-row); (f) Neighbor-joining tree of barley genotypes based on geographical location, with different colors representing different regions; (g) Neighbor-joining tree based on growth habit, with different colors representing traits such as spring and winter barley; (h) Neighbor-joining tree analysis of 47 Australian barley cultivars, showing the genetic relationships between varieties developed during different periods (Adapted from Hill et al., 2021)

Additionally, evolutionary breeding programs have been employed to develop barley populations and lines specifically for organic and low-input agriculture. These programs combine natural and artificial selection to create heterogeneous populations with high grain yield and yield stability across different environments (Raggi et al., 2017). This approach has proven effective in developing barley varieties that perform well under low-input conditions, which is essential for sustainable agriculture.

4.2 Evolution of major Malting Barley growing regions

The cultivation of Malting Barley has evolved significantly, influenced by climate, soil, and management techniques. The spread of barley from its origin in the Fertile Crescent to various parts of the world involved significant ecological and environmental adaptations. For example, the transition from winter to spring barley in eastern Asia allowed barley to be cultivated in regions with different growing seasons, enhancing its adaptability (Liu et al., 2017).

Moreover, the genetic adaptation of barley to new environments involved the selection of preexisting genetic variants rather than the acquisition of new mutations. This adaptation process is exemplified by the natural variation in the HvCEN gene, which contributed to the successful environmental adaptation of spring and winter barleys (Comadran et al., 2012). The ability of barley to adapt to diverse environments has been a key factor in its global spread and cultivation.

4.3 The importance of Malting Barley in modern agriculture

Malting Barley plays a crucial role in modern agriculture, contributing to agricultural economics, food security, and the brewing industry. The differentiation of agricultural products, such as unique barley varieties for craft malting and brewing, has been adopted as a strategy to improve farm profitability and business sustainability. Studies have shown that consumers are willing to pay more for beers that support local farmers, highlighting the economic importance of barley in the craft beer value chain (Craine et al., 2022).

Furthermore, the domestication and genetic improvement of barley have provided insights into ancient human civilization and the mechanisms that converted wild species into essential food crops. The identification of domestication genes has revealed that most drastic changes during domestication resulted from functional impairments in transcription factor genes, which facilitated the development of traits such as increased seed number per spike (Pourkheirandish and Komatsuda, 2007). This genetic knowledge is vital for ongoing efforts to improve barley varieties and ensure food security.

5 Global Impact of Malting Barley and Brewing Culture

5.1 Influence of Malting Barley on global brewing cultures

The cultivation of barley and its use in brewing has significantly influenced brewing cultures worldwide. Different barley varieties and their growing conditions contribute uniquely to the flavor profiles of beers, which in turn shape regional brewing traditions. For instance, research has shown that barley genotype and growing location can affect beer flavor, with specific varieties like Golden Promise and Full Pint contributing distinct sensory descriptors such as fruity, floral, and malty notes (Figure 2) (Herb et al., 2017; Morrissy et al., 2022). Additionally, the timing of nitrogen fertilization and pest infestation can impact the quality of the malt and the final beer product, further illustrating the intricate relationship between barley cultivation practices and brewing outcomes (Marconi et al., 2011).

Figure 2 Correspondence analysis (CA) of beer sensory evaluation (Adapted from Morrissy et al., 2022) Image Caption: The analysis data is derived from a trained sensory panel, evaluating the aroma, taste, and mouthfeel of beers brewed from different barley varieties using the "CATA" method. Principal components F1 and F2 explain 52.03% of the total data variance. Different barley varieties are represented by abbreviations. Various sensory attributes are color-coded: blue for aroma, orange for taste, and gray for mouthfeel. Sensory attributes such as "sweet," "bitter," "grassy," "bready," etc., are positioned around the corresponding samples, indicating the specific sensory characteristics of each variety (Adapted from Morrissy et al., 2022)

Morrissy et al. (2022) presented a correspondence analysis that demonstrates the relationship between barley varieties and sensory characteristics, revealing significant differences in aroma, taste, and mouthfeel among different varieties. For example, the Thunder variety is associated with fruity and full-bodied mouthfeel, while the DH140963 line is linked to bitterness and acidity. This analysis supports the selection of barley varieties based on sensory traits, providing data for variety screening in the brewing process and facilitating the development of beers with distinct flavors and mouthfeels.

5.2 Evolution of barley varieties in cultural exchanges

Barley varieties have evolved through cultural exchanges and adaptation to local environments, leading to the selection of specific genotypes that best suit regional brewing needs. Studies have highlighted the nuanced contributions of barley variety and growing location to beer flavor, emphasizing the importance of selecting appropriate barley genotypes for desired sensory outcomes (Herb et al., 2017; Morrissy et al., 2022). The genetic characterization and selection for barley contributions to beer flavor are crucial, as different genotypes can significantly influence the sensory descriptors of beer, even with varying degrees of malt modification (Herb et al., 2017). This ongoing evolution and selection process ensure that barley varieties continue to meet the diverse demands of global brewing cultures.

5.3 Globalization of brewing and Malting Barley as a cultural symbol

The globalization of brewing has transformed beer from a local tradition to a global cultural symbol. The widespread cultivation of barley and the exchange of brewing techniques have facilitated this transformation. For example, the adaptive laboratory evolution of ale and lager yeasts has improved brewing efficiency and beer quality, catering to the global market's demand for diverse and high-quality beers (Gibson et al., 2020). Moreover, the integration of barley brewing by-products from barley into various food applications highlights the versatility and economic significance of barley in the global food industry (Gupta et al., 2010). As a result, Malting Barley has become a symbol of cultural exchange and innovation, reflecting the dynamic nature of global brewing traditions.

By examining the influence of barley variety, the evolution of barley through cultural exchanges, and the globalization of brewing, we can appreciate the profound impact of Malting Barley on global brewing cultures. This interconnectedness underscores the importance of continued research and innovation in barley cultivation and brewing practices to sustain and enhance the rich tapestry of global beer culture.

6 Adaptation and Challenges of Malting Barley in the Context of Climate Change

6.1 Impact of climate change on Malting Barley cultivation

Climate change poses significant challenges to Malting Barley cultivation, primarily through increased frequency and severity of droughts, shifts in precipitation patterns, and extreme weather events. Studies have shown that extreme drought and heat can lead to substantial decreases in barley yields, with average losses ranging from 3% to 17% depending on the severity of the conditions. These yield reductions directly impact the availability and economic accessibility of beer, as reflected by the dramatic regional decreases in beer consumption and significant price increases (Xie et al., 2018). Additionally, the Mediterranean basin, a key barley-growing region, is projected to experience hotter and drier conditions, leading to an overall 9% reduction in grain yield under future climate scenarios (Cammarano et al., 2019). The combined effects of heat and drought stress during critical growth stages, such as grain filling, further exacerbate yield losses and negatively affect malting quality traits (Mahalingam, 2017).

6.2 Climate-resilient breeding and technological advances

To address the challenges posed by climate change, significant efforts are being made in breeding climate-resilient barley varieties (Huang, 2024). The genetic diversity present in landraces, wild barley accessions, and other Hordeum species offers valuable sources of new alleles for developing climate-smart crops. Advances in genomic and analytical tools facilitate the exploration and capture of this genetic variation, enabling the development of barley varieties tailored to specific environments (Dawson et al., 2015). For instance, studies have identified specific genes and alleles associated with drought tolerance, which can be targeted in breeding programs to enhance barley's resilience to climate stresses (Sallam et al., 2019). Moreover, the use of exome sequencing and multi-environment field trials has elucidated the genetic basis of adaptation in barley, providing practical opportunities for crop improvement (Bustos-Korts et al., 2019). Technological advances, such as the Decision Support System for Agrotechnology Transfer (DSSAT) model, also play a crucial role in simulating the impacts of different climate projections and identifying effective adaptation strategies (Cammarano et al., 2019).

6.3 Future strategies to tackle climate change

Future strategies to enhance barley's resilience to climate change involve a combination of innovative breeding techniques and sustainable agricultural management practices. One promising approach is the introduction of novel germplasm into breeding programs to increase genetic diversity and improve adaptive capacity (Mahalingam, 2017). Additionally, the selection of proper cultivars for future climate conditions, including those that can withstand extreme weather events, is crucial for maintaining or improving crop productivity (Niero et al., 2015). Participatory improvement methods, which involve decentralized breeding efforts and collaboration with local farmers, can also help develop barley varieties that are well-suited to specific environmental conditions (Dawson et al., 2015). Furthermore, integrating advanced climate modeling and trait/environment association analyses can guide the development of targeted breeding strategies and inform agricultural management practices to mitigate the impacts of climate change on barley cultivation (Dawson et al., 2015; Cammarano et al., 2019).

7 Future Globalization Trends and Sustainable Development of Malting Barley

7.1 Sustainable practices in Malting Barley cultivation

Sustainable practices in Malting Barley cultivation are essential for reducing the environmental impact and ensuring long-term agricultural viability. One promising approach is the use of low-input, organically grown cereals, which can lead to a lower environmental footprint. This is particularly relevant for old wheat varieties, which, despite their lower yield, offer higher nutritional value and can be economically sustainable when integrated into brewing processes (Albanese et al., 2017). Additionally, the application of biostimulants such as seaweed extract has shown to improve nitrogen use efficiency and increase yield, thereby reducing the need for excessive nitrogen fertilization and minimizing environmental damage (Cozzolino et al., 2021). The integration of these practices can significantly contribute to ecological agriculture and carbon footprint reduction.

7.2 Impact of the global beer industry on barley cultivation

The global beer industry exerts considerable influence on barley cultivation, driven by consumer demand and market trends. Beer is the most popular alcoholic beverage worldwide, and its production heavily relies on barley. However, extreme weather events such as drought and heat, exacerbated by climate change, pose significant threats to barley yields, leading to potential decreases in beer supply and increases in prices (Xie et al., 2018). Moreover, the brewing industry's focus on sustainability is evident in efforts to valorize by-products like brewers' spent grain, which can be repurposed for human consumption, thus reducing waste and promoting resource efficiency (Jaeger et al., 2021). Regional barley-malt-beer value chains, as seen in Bavaria, Germany, also highlight the potential ecological benefits and the importance of local sourcing to mitigate environmental impacts (Maier et al., 2020).

7.3 Future directions for the global Malting Barley industry

The future of the global Malting Barley industry lies in balancing technological innovation with sustainable agricultural practices and cultural preservation. Advances in genomics and molecular markers have paved the way for better understanding and improving barley traits, which can enhance yield and resilience to environmental stresses (Haas et al., 2019). However, it is crucial to maintain the cultural heritage associated with traditional barley varieties and brewing methods. Craft breweries, despite facing economic sustainability challenges, play a vital role in preserving brewing culture and promoting biodiversity (Albanese et al., 2017). By integrating innovative technologies with sustainable practices and cultural values, the Malting Barley industry can achieve a harmonious balance that supports both economic and environmental sustainability.

8 Concluding Remarks

The domestication of barley, one of the earliest crops cultivated by humans, marks a significant milestone in the transition from hunter-gatherer societies to agrarian civilizations. Approximately 12 000 years ago, in the Near East, barley was domesticated through critical genetic mutations that enabled grain retention on the inflorescence, facilitating effective harvesting. This process involved the selection of non-brittle rachis variants, which were independently chosen by early farmers in different regions of the Levant, contributing to the rise of early agrarian societies.

Barley, along with wheat, was a cornerstone of the agricultural revolution in the Fertile Crescent around 10 000 years ago. The domestication process involved the evolution of traits beneficial to human use, such as improved grain retention, threshability, and yield, as well as changes in photoperiod sensitivity and nutritional value. Advances in genomic technologies have furthered our understanding of these domestication traits, revealing the complex genetic underpinnings of barley's evolution. The spread of barley from its origins in the Fertile Crescent to other parts of the world, including Tibet, highlights its adaptability and the role of cultural exchanges, such as those facilitated by the Silk Road, in its dissemination. The genetic diversity observed in barley populations across different regions underscores the crop's evolutionary history and its adaptation to various environmental conditions.

Barley holds a unique position in global agriculture and culture. It is one of the oldest and most versatile crops, capable of thriving in diverse climates and environments. Historically, barley has been integral to various cultural practices, including baking, cooking, and beer brewing, and has played a significant role in the diets and economies of ancient civilizations such as those in Mesopotamia and Egypt.

The future prospects for barley are promising, particularly in the realm of sustainable agriculture. Its genetic diversity and adaptability make it a valuable resource for developing resilient crop varieties that can withstand the challenges posed by changing environmental conditions. As research continues to uncover the genetic mechanisms behind barley's domestication and evolution, there is potential for further improvements in yield, nutritional quality, and stress resistance, ensuring that barley remains a staple crop in global agriculture and culture for generations to come.

Acknowledgments

We thank the anonymous reviewers for their insightful comments and suggestions that greatly improved the manuscript.

Conflict of Interest Disclosure

The authors affirm that this research was conducted without any commercial or financial relationships that could be construed as a potential conflict of interest.

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