Use of Crop Wild Relatives in Wheat for Climate Change and Food Security

Anand Kumar; Laxmidas Verma; Ravindra Kumar; Sagar

doi:10.5772/intechopen.1012644

Abstract

Global food security relies on the ongoing enhancement of a limited number of key food and feed crops. The current food supply is significantly dependent on technology. To feed over 9 billion people by 2050, we must double high-quality food production while ensuring environmental and social sustainability. Current crop improvement practices are insufficient to meet future demands or address the impact of climate change. Integrating food security and climate change strategies presents a promising approach to enhance crop resilience against biotic and abiotic stresses. Wheat is a major staple crop worldwide; wheat yield gains were achieved through increased agricultural inputs and the expansion of arable land. However, future improvements hinge on genetic enhancements, which have declined due to reduced genetic diversity in wheat breeding. This lack of variability poses a challenge for further advancements in technology. Moreover, using both wild and cultivated relatives of wheat offers new opportunities for genetic improvement in wheat production. Innovations in genetic enhancements for wheat breeding are vital for addressing climate change challenges and ensuring food security. Identifying and utilizing these advancements will revolutionize wheat cultivation, enabling precise improvements in crop resilience and productivity. There are several conventional techniques used to improve wheat productivity and resistance to biotic and abiotic stress. However, these techniques are more expensive and time-consuming. Here, in this book chapter, we describe the use of crop wild relatives and advancements in technology for global food security.

Keywords

wheat
global food security
crop wild relatives (CWR)
biotic and abiotic stress and advancement of technology
climate change

Author Information

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Anand Kumar *
- Faculty of Agricultural Sciences, GLA University, Mathura, India
Laxmidas Verma
- Department of Genetics and Plant Breeding, Acharya Narendra Dev University of Agriculture and Technology, Kumarganj, India
Ravindra Kumar
- Department of Agriculture, Mata Gujri College, Fatehgarh Sahib, India
Sagar
- Faculty of Agricultural Sciences, GLA University, Mathura, India

*Address all correspondence to: anandkumar002014@gmail.com

1. Introduction

Mankind must confront the truth of global warming. Food security is the most essential need for all life on Earth, and it is significantly affected by climate change. This makes it a pressing issue that requires urgent action [1]. Climate change, fluctuations in weather during the growing season, adaptation strategies that help crops realize their genetic potential in response to a changing climate, and the creation of cultivars better suited to varying environmental conditions are all factors that will influence future food security [2]. Since our capacity to foretell future climates is uncertain, these problems may seem insurmountable. Despite these obstacles, we can learn more about the interplay between soil, plants, and the atmosphere, and how to use this information to further our aim of universally improved food security [3].

Crop wild relatives (CWR) are plant species that are closely related to field crops and their ancestors. These species can contribute beneficial traits to crop improvement, such as resistance to biotic and abiotic stresses, and they can enrich the gene pool, which ultimately leads to enhanced plant yield [4]. This helps humanity in its relentless search for food production to meet the ever-growing demands of a burgeoning global population. Actually, CWRs are well-known for their immense potential to maintain and improve global food security, which would greatly benefit humanity [5]. This underscores the vital importance of identifying, describing, and preserving these elements in crop breeding efforts. The significance of this work is further amplified in light of the recent challenges posed by global climate change [6].

Given the severe consequences, particularly the concerning environmental risks, of the agricultural practices that heavily relied on soil extraction throughout the 1960s, sometimes referred to as the “Green Revolution,” the task becomes even more important [7].

Climate change poses a serious risk to humankind’s ability to grow food. Greenhouse gas emissions from human activities are rapidly altering the environment and accelerating the process of global warming. Soil water deficiencies will worsen, and organic matter decomposition will speed up as a result of this in the future [8]. Soil erosion will be a major issue unless we put conservation measures in place to address the rising frequency of heavy rainfall, runoff, and flooding in the future [9]. Conservative agricultural practices, such as reduced or no tillage, crop residue retention, and diverse cropping systems, can help reduce atmospheric CO₂ levels, according to a new mitigation strategy that aims to transfer the gas into soil organic carbon. The reach of these policies is limited, nevertheless, due to the many interregional and international constraints that prevent them from being implemented [10].

Putting the two trends of rising populations and average global temperatures side by side paints a bleak picture. According to current predictions, the global temperature is expected to increase by 1.8–4°C, and the global population could reach 10 billion by 2100, a 142.9% increase from the present 7 billion, or around a 1.8% per year increase [5]. Two types of “greenhouse effects” exist: one, which is “natural,” makes certain high-latitude and high-altitude regions habitable for life, and the other, which is far more common and much more harmful to the environment, is caused by industrial gases like carbon dioxide and agricultural practices like the overuse and unchecked use of nitrogen fertilizers such as urea. These practices lead to huge concentrations of nitrous oxide in the air and are killing off plants and humans alike [11]. There has been a 2000% increase in the atmospheric concentration of carbon dioxide (CO₂) from its pre-industrial value of 20 ppm to 400 ppm, with a significant portion of this increase attributable to emissions from automobiles [12], an absolutely astounding growth. The most worrisome aspect, however, is the generation of nitrous oxide (N₂O), also called “laughing gas” (dinitrogen monoxide), which is a key contributor to the depletion of stratospheric ozone and, by extension, to climate change. As a “greenhouse gas,” nitrous oxide has the third-greatest impact on global warming, after carbon dioxide and methane (CH₄) [13]. Nitrous oxide (N₂O) is 310 times more effective at trapping heat than carbon dioxide (CO₂). It typically remains in the atmosphere for about 120 years. The excessive use of nitrogen-based fertilizers, like urea, is responsible for a significant portion of agricultural emissions, contributing to 30% of the N₂O in the atmosphere. During the 1960s, coinciding with the Green Revolution, there was a notable rise in atmospheric nitrogen oxides (NO_x) [14]. When nitrogen fertilizers, such as urea, are used in excess with the goal of increasing food production, soil bacteria convert the nitrogen in the fertilizer to nitrous oxide at a greater rate than usual [15]. All the environmental consequences of global warming, such as reduced precipitation, soil deterioration, groundwater depletion, etc., have their roots in this. This ecological catastrophe is on full display in the Indian state of Punjab, sometimes referred to as the “cradle” of the Green Revolution, where thousands of acres of arable land have been eradicated [16].

Preserving CWR and extracting potentially useful genes to boost agricultural output is crucial, especially given the growing concerns about the devastating impacts of climate change on biodiversity and food security, alongside the world’s rapidly increasing population [17]. To adapt domestic crops to climate change, CWRs are an important tool to overcome the restrictions of genetic variety. A lack of knowledge about their diversity, present and future value, and practical ways to conserve them is preventing their integration into conservation and encouraging greater systematic exploitation [18].

Environmental shifts and the movement of birds: Israeli specialists have found that, due to climate change, Israel is now serving as a permanent wintering ground for some 500 million migrating birds. These birds used to make brief stops before heading to the warmer plains of Africa. However, instead of traveling to Africa, where food is becoming harder to find due to expanding deserts and frequent droughts, they are now choosing to stay longer in milder areas [19]. Ornithologist Shay Agmon, who is in charge of the wetlands sector of Agamon Hula in northern Israel, stated that “many more birds and a greater number of species can no longer cross the desert,” meaning that Israel has become more than simply a brief stopover for the birds in recent decades [20]. Over time, their duration of stay will increase, and the overall pattern of migration will undergo a transformation. Migrating birds may be fascinating to observers of birds and visitors, but they pose a threat to farmers due to their voracious appetite for crops. In the Syrian-African Rift Valley, in the Hula reserve, workers have made life more comfortable for the birds by feeding them in the marshes and luring them away from the fields [21]. A considerably wider desert is more difficult for the birds to fly across, and they just cannot do it. According to Agmon, fuel is running low, and there aren’t enough “gas stations” along the route; thus, Israel has become a major hub for them. The migratory patterns of birds can teach us something about how to deal with the problems of climate change, which threatens the entire global food chain. Worldwide, regionally, and even locally, climatic change, stress, and variability [22] are the primary factors that determine biodiversity and evolutionary change [23]. Perhaps no other environmental threat that the world confronts today compares to the severity of global warming. An assessment of this issue is given in a recent report by an intergovernmental panel on climate change [24]. Climate change has been studied by [25] in relation to agricultural and wildlife populations and by [26] in terms of bananas, barley, beans, cassava, chickpeas, pigeonpeas, potatoes, faba beans, groundnuts, lentils, maize, millet, rice, sorghum, soybeans, wheat, yam, forages, fisheries, and aquaculture. From a scientific, political, and economic perspective, the most controversial subject today is global warming and its causes, which were briefly touched upon in the introductory chapter. There is still noticeable disagreement among some global powers, especially the United States, but everyone agrees that if we don’t do something about climate change, it will have a devastating impact on farming, just like the very soil-exploitative Green Revolution had on the poor [27].

Domestication of plants and animals is the first step in the evolutionary chain that, after millennia, culminated in modern human society. Darwin postulated in 1905 that domestication is a massive evolutionary experiment that produces new species through adaptation and speciation. Like speciation in nature, it has mostly been done by humans in the last 10,000 years [28].

Adaptive syndromes that are suitable for human ecology are the result. Humans were permanently settled into cities, developed sophisticated cultural practices, and saw a massive increase in population as a result of domestication and the shift from pre-agricultural to agricultural economies. Domesticated adaptation syndromes in wheat involve higher yields and spikes that remain intact. Due to the process of domestication, these cultivars have become culturally and economically reliant on humans, thriving only in specific agricultural environments shaped by human activity [29]. One of the most detrimental outcomes of domestication, in hindsight, was the complete failure of the so-called Green Revolution and the environmental risks it introduced due to the overuse of artificial fertilizers on already fertile soils [30]. The sad commentary of this collapse is seen in the countries of South Asia, especially India and, more specifically, in the Indian state of Punjab.

2. A brief history of CWR in wheat

Crop wild relatives (CWRs) are wild plant species closely related to cultivated crops. They serve as vital reservoirs of genetic diversity that have significantly contributed to agricultural advancements. In the case of wheat (Triticum spp.), CWR have been crucial in boosting yields, improving disease resistance, and adapting to various environmental conditions [31]. From the early domestication of wheat in the Fertile Crescent to the cutting-edge genomics of the 21st century, CWR have been a cornerstone of wheat improvement. As global challenges mount, the importance of these genetic resources continues to grow, underscoring the need for their conservation and sustainable use [32].

2.1 Early domestication and role of wild relatives

The history of wheat begins around 10,000 years ago in the Fertile Crescent, where early humans first domesticated Triticum dicoccum (emmer wheat) and Triticum monococcum (einkorn wheat). These species evolved from wild relatives like Triticum urartu and Aegilops tauschii. The hybridization of these wild ancestors gave rise to hexaploid wheat (Triticum aestivum), which is the basis of modern bread wheat [33].

During this domestication process, early farmers unintentionally selected for beneficial traits. However, domestication also resulted in a genetic bottleneck, where much of the genetic diversity present in wild species was lost. This made cultivated wheat less resilient to environmental changes, diseases, and pests [34].

2.2 19th and early 20th century: Rediscovery of CWR

The systematic use of CWR in wheat breeding began in the late 19th and early 20th centuries. Botanists and plant breeders recognized the potential of wild relatives as sources of resistance to diseases such as rusts (caused by Puccinia spp.) [35].

Nikolai Vavilov’s contribution: The Russian geneticist Nikolai Vavilov was among the first to identify the importance of CWR. In the 1920s and 1930s, Vavilov’s expeditions across the world helped collect a vast array of wild wheat species and landraces. He proposed the “Centers of Origin” theory, emphasizing the significance of wild relatives in the genetic improvement of crops [36].
Aegilops tauschii in bread wheat: The discovery of Aegilops tauschii as the donor of the D-genome in hexaploid wheat highlighted its importance in breeding. This wild relative provided genetic material crucial for traits like cold tolerance and resistance to certain pathogens [37].

2.3 Mid-20th century: Green revolution and beyond

The Green Revolution of the 1940s–1960s, led by scientists like Norman Borlaug, brought about a massive increase in wheat production. While much of this success was due to semi-dwarf varieties and improved agronomic practices, the role of CWR in wheat improvement became increasingly evident [38].

Disease resistance: During this period, wild relatives were intensively used to introduce resistance genes. For example, genes from Aegilops spp. and Secale cereale (rye, another distant relative) were incorporated into wheat to combat rust diseases [39].
Synthetic wheat development: The concept of synthetic wheats emerged in the mid-20th century. By crossing diploid and tetraploid species with Aegilops tauschii, breeders recreated the hybridization events that originally gave rise to hexaploid wheat. These synthetic wheats introduced novel genetic diversity into the wheat gene pool, enhancing traits like drought tolerance and pest resistance [40].

2.4 Late 20th century: Advances in molecular biology

The advent of molecular biology and genomics in the 1980s and 1990s revolutionized the use of CWR in wheat breeding. Scientists were now able to precisely identify and transfer specific genes from wild relatives into cultivated varieties [41].

Marker-assisted selection (MAS): This technique enables breeders to track the incorporation of beneficial genes from CWR into wheat, reducing the time and effort required for traditional breeding.
Translocations and introgressions: Segments of chromosomes from wild relatives have been successfully introduced into wheat. For example, the wheat-rye translocation (1BL/1RS) has brought in resistance to various diseases and enhanced yield potential [42].
Notable genes:
- The Lr (Leaf rust) and Sr (Stem rust) gene series from CWR, such as Sr31, provide long-lasting resistance against devastating rust outbreaks.
- The Gpc-B1 gene, derived from wild emmer wheat, enhances grain protein content.

2.5 21st century: Climate change and renewed interest in CWR

The increasing challenges of climate change, a growing population, and the need for sustainable agriculture have reignited interest in CWR. Modern wheat breeding programs focus on traits like heat tolerance, water-use efficiency, and resistance to emerging pests and pathogens.

Global conservation efforts: Organizations like the Global Crop Diversity Trust (Crop Trust) and initiatives such as the Svalbard Global Seed Vault are safeguarding CWR for future use. The International Maize and Wheat Improvement Center (CIMMYT) and other research institutions maintain vast germplasm collections, including CWR.
Genome editing and genomics: The application of CRISPR-Cas9 and other genome-editing tools has made the precise transfer of genes from CWR into wheat a reality. Researchers are now able to directly target traits of interest without disrupting other aspects of the plant’s genome.
New frontiers: Efforts are underway to domesticate CWR themselves, effectively turning wild species into viable crops. Additionally, advanced genomic tools have revealed hidden diversity within wild species, opening new possibilities for wheat improvement.

2.6 Challenges in using CWR

Despite their potential, integrating CWR into wheat breeding is not without its challenges.

Genetic linkage drag: Transferring desirable traits often brings along unwanted characteristics.
Crossing barriers: Some CWR are difficult to hybridize with cultivated wheat due to ploidy differences or genetic incompatibilities.
Habitat loss: The natural habitats of many CWRs are under threat from human activities, reducing the availability of these valuable genetic resources.

3. Use of advance technology for biotic and abiotic resistance

The genetic variability of wheat is increasingly limited due to reliance on conventional breeding methods involving landraces and common varieties. This narrowing gene pool poses challenges in addressing emerging biotic and abiotic stresses driven by climate change, such as salinity, waterlogging, drought, heat, and pests. Broadening the genetic base is critical for enhancing productivity and yield stability. Wild wheat progenitors harbor valuable genetic variations that can strengthen breeding pools. Utilizing these resources through conventional breeding and wide-crossing techniques, including interspecific, intraspecific, and intergeneric hybridization, is essential for improving wheat resilience and adaptability [43].

3.1 Biotic stress

Biotic stress in plants arises from living organisms such as fungi, viruses, insects, nematodes, arachnids, and weeds. Unlike abiotic stresses caused by environmental factors, biotic stresses directly affect plant growth and development by depriving them of essential nutrients, which can lead to reduced vigor and even death. In agriculture, these stresses are major contributors to pre- and post-harvest losses, significantly impacting crop productivity and food security [44].

The rising global population and increasing abiotic stresses, exacerbated by climate change, are compounded by significant biotic stresses that threaten wheat production worldwide. Diseases like rusts and mildew, along with pests such as aphids, drastically reduce the yield potential of elite wheat cultivars. The complex nature of plant-parasite interactions is pivotal in determining resistance expression. Biotic stress in plants is caused by living organisms such as fungi, viruses, insects, nematodes, arachnids, and weeds. Unlike abiotic stresses (e.g., heat and drought) caused by environmental factors, biotic stress agents directly impact host growth and development by depriving plants of essential nutrients, leading to reduced vigor and, in severe cases, plant death. In agriculture, biotic stresses significantly contribute to both pre- and post-harvest losses, posing a major challenge to crop productivity and food security [45].

3.1.1 Leaf and stem rust

Wheat leaf rust, caused by Puccinia triticina Eriks., wheat stem rust by Puccinia graminis f. sp. tritici, and wheat stripe rust by Puccinia striiformis f. sp. tritici, are among the most significant diseases affecting wheat crops. Among these, leaf rust occurs more frequently and in a wider range of regions globally compared to stem rust and stripe rust. Yield losses due to P. triticina have a considerable economic impact; infections typically result in a reduced number of kernels per head and decreased kernel weight, leading to yield losses that can reach up to 40% in highly susceptible cultivars [46]. Yield losses caused by stem rust pathogens during the mid-20th century reached 20–30% in Eastern and Central Europe, as well as in many other countries, including Australia, China, and India [47, 48].

3.2 Abiotic stress (e.g., drought tolerance, heat tolerance, salinity, and heavy metals tolerance)

In recent years, global warming and climate change have significantly impacted agricultural productivity in tropical and subtropical regions, leading to the emergence of new abiotic stresses. Among these, heat, drought, moisture, and salt stress are the most prevalent. This section provides insights into the effects of various abiotic stresses on the growth and physiology of wheat [49].

3.2.1 High temperature

Heat is one of the most common abiotic stresses affecting wheat. This section explores its impact on wheat growth and physiology. Wheat is particularly vulnerable to heat stress during specific physiological growth stages [50]. Heat stress, caused by elevated temperatures, is defined as an increase in air temperature beyond a critical threshold for a specific duration, leading to injury or irreversible damage in crop plants. A temperature rise of 1°C can result in a yield loss of approximately 4.1% to 6%. Studies have shown that heat stress induces pollen sterility, reduces carbon dioxide assimilation, and enhances photorespiration in wheat. Additionally, high temperatures impair the photosynthetic process, negatively affecting wheat growth and yield [51].

3.2.2 Drought stress

Drought stress, characterized by water deficiency, triggers significant morphological, biochemical, physiological, and molecular changes, making plant growth and crop production challenging. It negatively affects wheat at all developmental stages, with the severity of its impact depending on stage-specific stress and local climate conditions [52]. More than half of the world’s wheat-growing regions experience drought regularly, leading to production losses of up to 10%. Additionally, global warming is expected to exacerbate drought frequency and intensity due to declining precipitation and increased evaporation.

3.2.3 Salinity stress

Salinity is one of the most detrimental abiotic stresses, significantly reducing agricultural productivity. It severely impacts plant morphology, physiology, and biochemistry, affecting processes such as photosynthesis, nutrient uptake, and yield. Studies have shown that wheat cultivars exhibit varying degrees of salt tolerance, with some being more resistant, while others are highly susceptible. High salinity levels hinder germination and increase Na⁺ and Cl⁻ ion concentrations, disrupting the normal metabolic processes of wheat plants.

Salt stress affects plants through osmotic imbalance, ion toxicity, and nutritional disorders. In response to salinity stress, researchers assess seedling growth, seed reserve utilization, and biomass weight. Studies on different wheat cultivars show that salinity reduces relative water content, chlorophyll and carotenoid levels, biomass, and grain yield, likely due to increased oxidative stress. Under 150 mM NaCl stress, key physiological parameters, such as net photosynthetic rate, transpiration rate, stomatal conductance, and sub-stomatal carbon dioxide concentration decline significantly [53].

When plants are exposed to environmental stresses such as drought, salinity, and heat, they generate a large number of reactive oxygen species (ROS), which are biosynthesized in the chloroplast, mitochondria, and/or peroxisomes of plant cells. Reactive oxygen species (ROS) are highly reactive, and their excessive production can disrupt normal plant metabolism. High ROS concentrations can damage plant cells by promoting lipid peroxidation, protein degradation, and DNA denaturation, leading to oxidative stress. This, in turn, can impair enzymatic activity, disrupt cellular structures, and even result in cell death. Another stress metabolite, methylglyoxal (MG), is produced in plant chloroplasts, mitochondria, and the cytosol under abiotic stress conditions. MG is toxic to plant cells, contributing to oxidative stress and disrupting cellular functions. Its impact depends on its concentration and the severity of the stress. Among abiotic stresses, drought stress disrupts the balance between antioxidant defenses and drought-induced ROS production. Malondialdehyde (MDA) production is considered a marker of ROS generation and oxidative damage. Elevated levels of H₂O₂ and MDA in wheat cells disrupt adaptive mechanisms under drought-induced oxidative stress. Similarly, salinity stress is a major environmental constraint that inhibits plant growth and physiological functions by triggering excessive ROS production. Increased MDA accumulation in wheat cultivars under salinity stress leads to oxidative damage in parenchymal cells [54].

3.2.4 Advances in molecular breeding for wheat improvement

While traditional breeding has played a significant role in enhancing wheat yields, modern molecular breeding techniques have transformed the creation of superior wheat varieties. Some important molecular methods include:

Genomic selection (GS): Genomic selection employs high-throughput DNA sequencing and statistical models to estimate the breeding value of wheat plants based on their genetic composition. This method enables breeders to identify the most promising candidates for breeding, thereby shortening the time needed for variety development [55].
CRISPR/Cas9 genome editing: CRISPR/Cas9 has become a powerful tool for making precise genetic changes in wheat. This technology allows researchers to modify genes that influence yield, stress tolerance, and disease resistance, thus speeding up the breeding process [56].
Transgenic approaches: Genetic engineering facilitates the incorporation of advantageous genes from other organisms into wheat. For instance, genes that provide resistance to fungal pathogens or enhance nitrogen-use efficiency have been successfully integrated into wheat using transgenic methods [57].

3.3 Current status of CWR usage for abiotic stress tolerance in wheat

The primary gene pool of wheat consists of its direct progenitors, which can be easily crossed with cultivated wheat to produce fertile offspring. These include Triticum urartu, Aegilops speltoides, and Aegilops tauschii. The secondary gene pool comprises wild relatives, such as Triticum turgidum ssp. dicoccoides and Triticum timopheevii, which can hybridize with wheat, though the resulting progeny are often infertile (Pour-Aboughadareh et al., 2021). The tertiary gene pool includes more distantly related species, such as Thinopyrum ponticum and Thinopyrum distichum, which typically require advanced breeding techniques, such as colchicine treatment or embryo rescue, to produce viable progeny [58].

Crop wild relatives (CWR) are valuable plant genetic resources (PGR) widely utilized by breeders. Sharing a close evolutionary relationship with their domesticated counterparts, CWR have retained higher genetic diversity due to their avoidance of domestication [59]. Ex situ conservation of CWR plays a crucial role in preserving this diversity and serves as a vital resource for introducing tolerance to abiotic stress factors in crop improvement programs. Climate change intensifies environmental (abiotic) stress, significantly impacting crop production. Rising global temperatures alter precipitation patterns, leading to increased rainfall in some regions and drought in others [60]. Additionally, higher temperatures accelerate evapotranspiration, further exacerbating water stress and affecting crop growth and yield. Thinopyrum distichum (Thunb.) Löve, commonly known as sea wheat, is a halophytic wild relative of wheat. Native to the coastal dunes of South Africa, this species thrives in saline environments, making it a valuable genetic resource for improving salt tolerance in wheat breeding. Extensive research has identified three key mechanisms that plants employ to tolerate high salt levels: osmotic adjustment, ion homeostasis, and ROS homeostasis. These mechanisms collectively help plants maintain cellular function and sustain growth under saline conditions [61].

Although, Thinopyrum distichum is known for its inherent tolerance to saline environments, the specific mechanisms it employs to manage high salt stress remain unidentified. Further research is needed to determine the physiological and molecular pathways underlying its salt tolerance [62]. Advances in molecular breeding techniques, including genomics, MAS, transgenic approaches, and gene editing, have greatly accelerated the development of salt-tolerant wheat varieties. Genome sequencing and the identification of quantitative trait loci (QTLs) associated with salinity tolerance enable more precise selection, while genetic engineering and CRISPR/Cas9 technology facilitate targeted gene modifications to enhance salt resistance [63]. Additionally, introgression breeding incorporating beneficial traits from wild wheat relatives further strengthens wheat’s resilience to saline conditions. A comprehensive understanding of the genetic and phenotypic diversity within local Thinopyrum distichum populations can enhance its potential for integration into wheat breeding programs, facilitating the development of salt-tolerant wheat varieties [64]. Previous research at Stellenbosch University’s Plant Breeding Laboratory on Thinopyrum distichum has involved hybridization efforts, exploring its potential for improving wheat through breeding. The salt tolerance potential of perennial grasses has been studied since the 1960s, when Agropyron elongatum was identified as the most salt-tolerant CWR species at the time [61].

Conservation and utilization of CWRs

Conservation strategies: To ensure the ongoing availability of CWRs, effective conservation efforts are essential. These efforts include:

In situ conservation: Safeguarding CWRs in their natural environments by establishing protected areas and biodiversity reserves.
Ex situ conservation: Storing CWRs in gene banks and seed vaults, such as the Svalbard Global Seed Vault and the International Center for Agricultural Research in the Dry Areas (ICARDA).
Community-based conservation: Involving local farmers and indigenous communities in the conservation of CWRs [65].

Challenges in CWR conservation and utilization

Despite their importance, several challenges impede the conservation and utilization of CWRs:

Habitat destruction: Urban development, deforestation, and changes in land use pose threats to the natural habitats of CWRs.

Limited research and funding: Many CWRs are under-researched due to a lack of funding for breeding programs and conservation efforts.

Crossbreeding barriers: Some CWRs are genetically distant from cultivated crops, necessitating advanced breeding techniques like MAS and gene editing.

Case studies: Success stories of CWR utilization

The introduction of wild wheat genes into commercial varieties: Researchers have successfully integrated genes from wild wheat species (Aegilops tauschii) to improve disease resistance and stress tolerance in modern wheat varieties. These initiatives have led to increased wheat yields in areas prone to drought [66].

4. Food and nutrition security challenge

Food security, as defined by the UN’s FAO, exists when all people, at all times, have physical, social, and economic access to sufficient, safe, and nutritious food that meets their dietary needs and preferences for an active, healthy life. This means food must not only be available but also safe to eat and nutritionally adequate. In many Western countries, where food abundance has been the norm for decades, food safety has become a greater concern than availability [67]. Plant breeders must recognize this shift, as consumer safety is paramount. Additionally, the food industry is highly sensitive to reputational risks, and companies will quickly adapt – whether by changing suppliers or raw materials – to mitigate the chances of negative publicity. The formation of the processing contaminant acrylamide during high-temperature cooking and processing of foods made from potatoes, cereals, and other crops is a significant food safety concern. The food industry is highly sensitive to such issues and seeks to minimize risks, making any increase in acrylamide levels particularly unwelcome [68].

The relationship between food security and nutrition security is complex, as evidenced by the link between food insecurity and malnutrition outcomes such as overweight and obesity. Data indicate that obesity is more prevalent in food-insecure populations for several reasons. Limited access to healthy foods can contribute to weight gain through multiple pathways. For instance, low-income families often prioritize cost-effective, energy-dense foods over more expensive, nutrient-rich options to maximize their limited resources [69].

Evidence suggests that food-insecure households often prioritize food quantity over quality or variety to prevent absolute hunger. This can lead to nutrition insecurity despite an abundance of calories, particularly if households lack access to a consistent and adequate diet. Additionally, when food is available only intermittently, individuals may cope by overeating during periods of accessibility, which can contribute to overweight and obesity [70].

Food security and nutrition security are interdependent; one cannot be achieved without the other. Nutrition security is a crucial component of food security, as adequate nutrition requires more than just sufficient caloric intake – it necessitates a diverse range of macro and micronutrients to support good health and disease prevention. Recognizing this essential connection, experts increasingly use the term "food and nutrition security" to highlight the integration of both food availability and nutritional well-being in addressing population health needs [71].

5. Conclusion

Crop wild relatives (CWR) are invaluable genetic resources for improving wheat resilience against both biotic and abiotic stresses. They possess diverse traits that can enhance resistance to diseases, pests, drought, salinity, heat, and other environmental challenges. With climate change and the emergence of new pathogens posing significant threats to global wheat production, harnessing the genetic diversity of CWR has become increasingly important for ensuring food security and sustainable agriculture.

Through conventional breeding, wide hybridization, and molecular techniques such as marker-assisted selection and gene editing, key resistance genes from CWR have been successfully introgressed into cultivated wheat varieties. Notable successes include the incorporation of genes for resistance to rusts, powdery mildew, and wheat blast, as well as tolerance to extreme climatic conditions. These advancements not only enhance yield stability but also reduce reliance on chemical inputs, promoting eco-friendly agricultural practices.

However, despite their potential, the utilization of CWR in wheat breeding faces several challenges, including difficulties in crossbreeding, linkage drag, and the loss of genetic diversity due to habitat destruction. Therefore, conservation efforts, both in situ (natural habitats) and ex situ (gene banks), are critical for preserving these genetic resources for future breeding programs.

In conclusion, the integration of CWR into wheat breeding programs is essential for developing climate-resilient, high-yielding, and disease-resistant wheat varieties. Continued research, investment in genetic conservation, and advanced breeding strategies will be key to unlocking the full potential of CWR in addressing global food security challenges.

Acknowledgments

All authors contribute equally.

Conflict of interest

The authors declare no conflict of interest.

Notes/thanks/other declarations

The authors thank the eminent reviewers.

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12. Magadza CH. Climate change: Some likely multiple impacts in Southern Africa. Climate Change and World Food Security. 1996;37:449–483
13. Bréchet LM, Salomόn RL, Machacova K, Stahl C, Burban B, Goret J-Y, Steppe K, Bonal D, Janssens IA. Insights into the subdaily variations in methane, nitrous oxide and carbon dioxide fluxes from upland tropical tree stems. New Phytologist. 2025;245:2451–2466
14. Ali AM. Environmental pollution and climate change implications of agricultural fertilizer use. Agricultural Nutrient Pollution and Climate Change: Challenges and Opportunities. 2025;11:1–28
15. Sarika, Anand A, Meena R, Mina U, Shukla A, Sharma A. Adoption of the green energy technology for the mitigation of greenhouse gas emission: Embracing the goals of the Paris agreement. Nanomaterials and Nanoliquids: Applications in Energy and Environment. 2023;16:47–72
16. Jatav HS, Minkina T, Singh SK, Singh B, Rajput VD, editors. Environmental Nexus for Resource Management, Vol. 12. Boca Raton: CRC Press. 2024. 01–416
17. Aziz A. Climate change: A growing challenge for food security in Pakistan. Social Science Review Archives. 2025;3:1390–1402
18. Telch A. Unravelling the Nexus between Food Systems and Climate Change: A Legal Analisys. A Plea for Smart Agriculture, A” New” Organic Agriculture and A Wiser Use of Biotechnologies in the Name of Human Rights Protection. 2019
19. Singh G, Sharma S. Desert ecology and functional aspects of desert ecosystems. Textbook of Forest Science. 2025;19:227–251
20. Friedemann G, Leshem Y, Kerem L, Shacham B, Bar-Massada A, McClain KM, Bohrer G, Izhaki I. Multidimensional differentiation in foraging resource use during breeding of two sympatric top predators. Scientific Reports. 2016;6:01–13
21. Dunne P. The Courage of Birds: And the Often Surprising Ways They Survive Winter. London, England: Chelsea Green Publishing; 2024
22. Root TL, Price JT, Hall KR, Schneider SH, Rosenzweig C, Pounds JA. Fingerprints of global warming on wild animals and plants. Nature. 2003;421:57–60
23. Nevo A. Measuring market power in the ready-to-eat cereal industry. Econometrica. 2001;69:307–342
24. Intergovernmental Panel on Climate Change. Key dates in the AR5 schedule. Available. 2013
25. Mercer KL, Perales HR. Evolutionary response of landraces to climate change in centers of crop diversity. Evolutionary Applications. 2010;3:480–493
26. Thornton PK, Kristjanson P, Förch W, Barahona C, Cramer L, Pradhan S. Is agricultural adaptation to global change in lower-income countries on track to meet the future food production challenge? Global Environmental Change. 2018;52:37–48
27. Howe JP. Behind the Curve: Science and the Politics of Global Warming. United States: University of Washington Press; 2014
28. Zohary D, Hopf M, Weiss E. Domestication of Plants in the Old World: The Origin and Spread of Domesticated Plants in Southwest Asia, Europe, and the Mediterranean Basin. Oxford, England: Oxford University Press; 2012
29. Yeomans L. Zooarchaeology of managed, captive, tame, and domestic birds: Shifts in human–avian relationships. Journal of Archaeological Research. 2025;33:1–42
30. He Y. Toward a Relational Foodscape in Chinese and Japanese North American Literature. Sale, England: Transnational Press London; 2025
31. Peters Haugrud AR, Achilli AL, Martínez‐Peña R, Klymiuk V. Future of durum wheat research and breeding: Insights from early career researchers. The Plant Genome. 2025;18:01–25
32. Zair W Conservation of Crop Wild Relatives’ Diversity in the Fertile Crescent (Doctoral dissertation, University of Birmingham). 2020
33. Giles RJ. The Origins of Triticum Aestivum L. England: The University of Manchester (United Kingdom); 2004
34. Turcotte MM, Araki H, Karp DS, Poveda K, Whitehead SR. The eco-evolutionary impacts of domestication and agricultural practices on wild species. Philosophical Transactions of the Royal Society B: Biological Sciences. 2017;372:01–09
35. Tirnaz S, Zandberg J, Thomas WJ, Marsh J, Edwards D, Batley J. Application of crop wild relatives in modern breeding: An overview of resources, experimental and computational methodologies. Frontiers in Plant Science. 2022;13:01–17
36. Endresen DTF Utilization of Plant Genetic Resources: a Lifeboat to the Gene Pool (P. 3). PhD Thesis, Department of Agriculture and Ecology, Faculty of Life Sciences, Copenhagen University, Denmark. 2011
37. Kishii M. An update of recent use of Aegilops species in wheat breeding. Frontiers in Plant Science. 2019;10:585
38. Cotter J. Troubled Harvest: Agronomy and Revolution in Mexico, 1880-2002. Bloomsbury Publishing USA; 2003;22
39. Kaur S, Jindal S, Kaur M, Chhuneja P. Utilization of wild species for wheat improvement using genomic approaches. In: Biotechnologies of crop improvement, volume 3. Genomic Approaches. Cham: Springer International Publishing; 2018. p. 105–150
40. Saville RJ Understanding DELLA in Wheat Linking Genotype to Phenotype (Doctoral dissertation, University of East Anglia). 2011
41. Jovovic Z, Andjelkovic V, Przulj N, Mandic D. Untapped genetic diversity of wild relatives for crop improvement. Rediscovery of Genetic and Genomic Resources for Future Food Security. 2020;13:25–65
42. Ning S, Wang X, Ma C, Liao J, Ye F, Li S, Fan C, Cui Q, Wang X, Song D, Hao M. Development and characterization of small translocations between the Pm13 introgression from Aegilops longissima and wheat homoeologous A, B and D chromosomes. BMC Plant Biology. 2025;25:179
43. Pramanik S, Debnath S. Development of rural landraces through mutation breeding approaches. Plant Mutagenesis: Sustainable Agriculture and Rural Landscapes. 2024;8:169–186
44. Choudhary V, Kumar KK, Sahu BHIMESHWARI, Muthappa SK, Das ADA. Agronomic innovations in biotic stress management and its combined effect with abiotic stresses in crop production. Indian Journal of Agronomy. 2021;66:237–257
45. Afzal F, Chaudhari SK, Gul A, Farooq A, Ali H, Nisar S, Sarfraz B, Shehzadi KJ, Mujeeb-Kazi A. Bread wheat (Triticum aestivum L.) under biotic and abiotic stresses: An overview. Crop Production and Global Environmental Issues 13. 2015;293–317
46. Knott DR. The Wheat Rusts—breeding for Resistance, Vol.12. Berlin and Heidelberg, Germany: Springer Science & Business Media; 2012
47. Leonard KJ, Szabo LJ. Stem rust of small grains and grasses caused by Puccinia graminis. Molecular Plant Pathology. 2005;6:99–111
48. Zadoks TC. Epidemiology of wheat rust in Europe. International Journal of Pest Management B. 1967;13:29–46
49. Oshunsanya SO, Nwosu NJ, Li Y. Abiotic stress in agricultural crops under climatic conditions. Sustainable Agriculture, Forest and Environmental Management. 2019;9;71–100
50. Akter N, Rafiqul Islam M. Heat stress effects and management in wheat. A review. Agronomy for Sustainable Development. 2017;37:1–17
51. Wahid A, Gelani S, Ashraf M, Foolad MR. Heat tolerance in plants: An overview. Environmental and Experimental Botany. 2007;61:199–223
52. Ahmad Z, Waraich EA, Akhtar S, Anjum S, Ahmad T, Mahboob W, Hafeez OBA, Tapera T, Labuschagne M, Rizwan M. Physiological responses of wheat to drought stress and its mitigation approaches. Acta Physiologiae Plantarum. 2018;40:1–13
53. Dhakal A, Adhikari C, Manandhar D, Bhattarai S, Shrestha S. Effect of abiotic stress in wheat: A review. Reviews in Food and Agriculture. 2021;2:69–72
54. Hossain A, Skalicky M, Brestic M, Maitra S, Ashraful Alam M, Syed MA, Hossain J, Sarkar S, Saha S, Bhadra P, Shankar T. Consequences and mitigation strategies of abiotic stresses in wheat (Triticum aestivum L.) under the changing climate. Agronomy. 2021;11:241.
55. Yunus MH, Firdaus A, Khan Z, Ansari MYK. Genomics-assisted breeding (GAB) for trait improvement: Unveiling genomic strategies for accelerated crop enhancement. Plant Breeding Technology: Future Trends and Challenges. 2025;64:138
56. Abdallah NA, Elsharawy H, Abulela HA, Thilmony R, Abdelhadi AA, Elarabi NI. Multiplex CRISPR/Cas9-mediated genome editing to address drought tolerance in wheat. GM Crops & Food. 2025;16:1–17
57. Kaya C. Plant metabolic engineering for enhanced nutrition and food security during climate uncertainty. Food and Energy Security. 2025;14:01–21
58. Mujeeb-Kazi A, Kazi AG, Dundas I, Rasheed A, Ogbonnaya F, Kishii M, Bonnett D, Wang RRC, Xu S, Chen P, Mahmood T. Genetic diversity for wheat improvement as a conduit to food security. Advances in Agronomy. 2013;122:179–257
59. Renzi JP, Coyne CJ, Berger J, von Wettberg E, Nelson M, Ureta S, Hernández F, Smýkal P, Brus J. How could the use of crop wild relatives in breeding increase the adaptation of crops to marginal environments. Frontiers in Plant Science. 2022;13:01–21
60. Engels JM, Thormann I. Main challenges and actions needed to improve conservation and sustainable use of our crop wild relatives. Plants. 2020;9:968
61. Haridass K Exploring the Abiotic Resistance Traits of the Local Wild Wheat Relative, Thinopyrum Distichum (Doctoral dissertation, Stellenbosch University). 2024
62. Neupane B Evaluation of Winter Wheat and Tritipyrum Germplasm to Identify Useful Variation for Biotic and Abiotic Stress Tolerance (Master’s thesis, North Dakota State University). 2023
63. Ashraf M, Munns R. Evolution of approaches to increase the salt tolerance of crops. Critical Reviews in Plant Sciences. 2022;41:128–160
64. Pour-Aboughadareh A, Kianersi F, Poczai P, Moradkhani H. Potential of wild relatives of wheat: Ideal genetic resources for future breeding programs. Agronomy. 2021;11:1656
65. Rouichi S, Ghanem ME, Amri M. In-situ and ex-situ conservation priorities and distribution of lentil wild relatives under climate change: A modelling approach. Journal of Applied Ecology. 2025;62:414–428
66. Yao Y, Guo W, Gou J, Hu Z, Liu J, Ma J, Zong Y, Xin M, Chen W, Li Q, Wang Z. Wheat2035: Integrating pan-omics and advanced biotechnology for future wheat design. Molecular Plant. 2025;18:272–297
67. Saint Ville A, Po JYT, Sen A, Bui A, Melgar-Quiñonez H. Food security and the Food Insecurity Experience Scale (FIES): Ensuring progress by 2030. Food Security. 2019;1:483–491
68. Curtis T, Halford NG. Food security: The challenge of increasing wheat yield and the importance of not compromising food safety. Annals of Applied Biology. 2014;164:354–372
69. Bhattacharya J, Currie J, Haider S. Poverty, food insecurity, and nutritional outcomes in children and adults. Journal of Health Economics. 2004;23:839–862
70. Timmer CP. Food Security and Scarcity: Why Ending Hunger Is so Hard. Philadelphia: University of Pennsylvania Press; 2015
71. Pérez-Escamilla R. Food and nutrition security definitions, constructs, frameworks, measurements, and applications: Global lessons. Frontiers in Public Health. 2024;12:01–10

[1] 1. Khan M, Zhang Y, Khan S. Echoes of Survival: Climate change impact & typologies of adaptation among vulnerable communities toward climate-induced food insecurity in Pakistan. Research on World Agricultural Economy. 2025;6:290–318

[2] 2. Muhammad M, Wahab A, Waheed A, Hakeem KR, Mohamed HI, Basit A, Toor MD, Liu YH, Li L, Li WJ. Navigating climate change: Exploring the dynamics between plant–soil microbiomes and their impact on plant growth and productivity. Global Change Biology. 2025;31:01–22

[3] 3. Raza A, Razzaq A, Mehmood SS, Zou X, Zhang X, Lv Y, Xu J. Impact of climate change on crops adaptation and strategies to tackle its outcome: A review. Plants. 2019;8(34):01–29

[4] 4. Bohra A, Kilian B, Sivasankar S, Caccamo M, Mba C, McCouch SR, Varshney RK. Reap the crop wild relatives for breeding future crops. Trends in Biotechnology. 2022;40:412–431

[5] 5. Nair KP. Utilizing crop wild relatives to combat global warming. Advances in Agronomy. 2019;153:175–258

[6] 6. Shyamala G, Gobinath R, Hemalatha B, Akkalla D, Ezhil SS, Sathya V. Global research trends in environmental adaptation techniques focusing on climate change through scientometric lens. Discover Sustainability. 2025;6:1–19

[7] 7. Farooq MS, Uzair M, Raza A, Habib M, Xu Y, Yousuf M, Yang SH, Ramzan Khan M. Uncovering the research gaps to alleviate the negative impacts of climate change on food security: A review. Frontiers in Plant Science. 2022;13:01–39

[8] 8. Muluneh MG. Impact of climate change on biodiversity and food security: A global perspective—a review article. Agriculture & Food Security. 2021;10:1–25

[9] 9. Hafeez A, Batool R, Arshad A, Khan MN, Ali S, Singh N, Garhwal V, Javed MA, Fatima E, Suleman F, Ali B. Soil and water conservation under changing climate. Challenges and Solutions of Climate Impact on Agriculture. 2025;13:307–328

[10] 10. Maetens W, Poesen J, Vanmaercke M. How effective are soil conservation techniques in reducing plot runoff and soil loss in Europe and the Mediterranean? Earth-Science Reviews. 2012;115:21–36

[11] 11. Sabina R, Paul J, Sharma S, Hussain N. Synthetic nitrogen fertilizer pollution: Global concerns and sustainable mitigating approaches. Agricultural Nutrient Pollution and Climate Change: Challenges and Opportunities. 2025;11:57–101

[12] 12. Magadza CH. Climate change: Some likely multiple impacts in Southern Africa. Climate Change and World Food Security. 1996;37:449–483

[13] 13. Bréchet LM, Salomόn RL, Machacova K, Stahl C, Burban B, Goret J-Y, Steppe K, Bonal D, Janssens IA. Insights into the subdaily variations in methane, nitrous oxide and carbon dioxide fluxes from upland tropical tree stems. New Phytologist. 2025;245:2451–2466

[14] 14. Ali AM. Environmental pollution and climate change implications of agricultural fertilizer use. Agricultural Nutrient Pollution and Climate Change: Challenges and Opportunities. 2025;11:1–28

[15] 15. Sarika, Anand A, Meena R, Mina U, Shukla A, Sharma A. Adoption of the green energy technology for the mitigation of greenhouse gas emission: Embracing the goals of the Paris agreement. Nanomaterials and Nanoliquids: Applications in Energy and Environment. 2023;16:47–72

[16] 16. Jatav HS, Minkina T, Singh SK, Singh B, Rajput VD, editors. Environmental Nexus for Resource Management, Vol. 12. Boca Raton: CRC Press. 2024. 01–416

[17] 17. Aziz A. Climate change: A growing challenge for food security in Pakistan. Social Science Review Archives. 2025;3:1390–1402

[18] 18. Telch A. Unravelling the Nexus between Food Systems and Climate Change: A Legal Analisys. A Plea for Smart Agriculture, A” New” Organic Agriculture and A Wiser Use of Biotechnologies in the Name of Human Rights Protection. 2019

[19] 19. Singh G, Sharma S. Desert ecology and functional aspects of desert ecosystems. Textbook of Forest Science. 2025;19:227–251

[20] 20. Friedemann G, Leshem Y, Kerem L, Shacham B, Bar-Massada A, McClain KM, Bohrer G, Izhaki I. Multidimensional differentiation in foraging resource use during breeding of two sympatric top predators. Scientific Reports. 2016;6:01–13

[21] 21. Dunne P. The Courage of Birds: And the Often Surprising Ways They Survive Winter. London, England: Chelsea Green Publishing; 2024

[22] 22. Root TL, Price JT, Hall KR, Schneider SH, Rosenzweig C, Pounds JA. Fingerprints of global warming on wild animals and plants. Nature. 2003;421:57–60

[23] 23. Nevo A. Measuring market power in the ready-to-eat cereal industry. Econometrica. 2001;69:307–342

[24] 24. Intergovernmental Panel on Climate Change. Key dates in the AR5 schedule. Available. 2013

[25] 25. Mercer KL, Perales HR. Evolutionary response of landraces to climate change in centers of crop diversity. Evolutionary Applications. 2010;3:480–493

[26] 26. Thornton PK, Kristjanson P, Förch W, Barahona C, Cramer L, Pradhan S. Is agricultural adaptation to global change in lower-income countries on track to meet the future food production challenge? Global Environmental Change. 2018;52:37–48

[27] 27. Howe JP. Behind the Curve: Science and the Politics of Global Warming. United States: University of Washington Press; 2014

[28] 28. Zohary D, Hopf M, Weiss E. Domestication of Plants in the Old World: The Origin and Spread of Domesticated Plants in Southwest Asia, Europe, and the Mediterranean Basin. Oxford, England: Oxford University Press; 2012

[29] 29. Yeomans L. Zooarchaeology of managed, captive, tame, and domestic birds: Shifts in human–avian relationships. Journal of Archaeological Research. 2025;33:1–42

[30] 30. He Y. Toward a Relational Foodscape in Chinese and Japanese North American Literature. Sale, England: Transnational Press London; 2025

[31] 31. Peters Haugrud AR, Achilli AL, Martínez‐Peña R, Klymiuk V. Future of durum wheat research and breeding: Insights from early career researchers. The Plant Genome. 2025;18:01–25

[32] 32. Zair W Conservation of Crop Wild Relatives’ Diversity in the Fertile Crescent (Doctoral dissertation, University of Birmingham). 2020

[33] 33. Giles RJ. The Origins of Triticum Aestivum L. England: The University of Manchester (United Kingdom); 2004

[34] 34. Turcotte MM, Araki H, Karp DS, Poveda K, Whitehead SR. The eco-evolutionary impacts of domestication and agricultural practices on wild species. Philosophical Transactions of the Royal Society B: Biological Sciences. 2017;372:01–09

[35] 35. Tirnaz S, Zandberg J, Thomas WJ, Marsh J, Edwards D, Batley J. Application of crop wild relatives in modern breeding: An overview of resources, experimental and computational methodologies. Frontiers in Plant Science. 2022;13:01–17

[36] 36. Endresen DTF Utilization of Plant Genetic Resources: a Lifeboat to the Gene Pool (P. 3). PhD Thesis, Department of Agriculture and Ecology, Faculty of Life Sciences, Copenhagen University, Denmark. 2011

[37] 37. Kishii M. An update of recent use of Aegilops species in wheat breeding. Frontiers in Plant Science. 2019;10:585

[38] 38. Cotter J. Troubled Harvest: Agronomy and Revolution in Mexico, 1880-2002. Bloomsbury Publishing USA; 2003;22

[39] 39. Kaur S, Jindal S, Kaur M, Chhuneja P. Utilization of wild species for wheat improvement using genomic approaches. In: Biotechnologies of crop improvement, volume 3. Genomic Approaches. Cham: Springer International Publishing; 2018. p. 105–150

[40] 40. Saville RJ Understanding DELLA in Wheat Linking Genotype to Phenotype (Doctoral dissertation, University of East Anglia). 2011

[41] 41. Jovovic Z, Andjelkovic V, Przulj N, Mandic D. Untapped genetic diversity of wild relatives for crop improvement. Rediscovery of Genetic and Genomic Resources for Future Food Security. 2020;13:25–65

[42] 42. Ning S, Wang X, Ma C, Liao J, Ye F, Li S, Fan C, Cui Q, Wang X, Song D, Hao M. Development and characterization of small translocations between the Pm13 introgression from Aegilops longissima and wheat homoeologous A, B and D chromosomes. BMC Plant Biology. 2025;25:179

[43] 43. Pramanik S, Debnath S. Development of rural landraces through mutation breeding approaches. Plant Mutagenesis: Sustainable Agriculture and Rural Landscapes. 2024;8:169–186

[44] 44. Choudhary V, Kumar KK, Sahu BHIMESHWARI, Muthappa SK, Das ADA. Agronomic innovations in biotic stress management and its combined effect with abiotic stresses in crop production. Indian Journal of Agronomy. 2021;66:237–257

[45] 45. Afzal F, Chaudhari SK, Gul A, Farooq A, Ali H, Nisar S, Sarfraz B, Shehzadi KJ, Mujeeb-Kazi A. Bread wheat (Triticum aestivum L.) under biotic and abiotic stresses: An overview. Crop Production and Global Environmental Issues 13. 2015;293–317

[46] 46. Knott DR. The Wheat Rusts—breeding for Resistance, Vol.12. Berlin and Heidelberg, Germany: Springer Science & Business Media; 2012

[47] 47. Leonard KJ, Szabo LJ. Stem rust of small grains and grasses caused by Puccinia graminis. Molecular Plant Pathology. 2005;6:99–111

[48] 48. Zadoks TC. Epidemiology of wheat rust in Europe. International Journal of Pest Management B. 1967;13:29–46

[49] 49. Oshunsanya SO, Nwosu NJ, Li Y. Abiotic stress in agricultural crops under climatic conditions. Sustainable Agriculture, Forest and Environmental Management. 2019;9;71–100

[50] 50. Akter N, Rafiqul Islam M. Heat stress effects and management in wheat. A review. Agronomy for Sustainable Development. 2017;37:1–17

[51] 51. Wahid A, Gelani S, Ashraf M, Foolad MR. Heat tolerance in plants: An overview. Environmental and Experimental Botany. 2007;61:199–223

[52] 52. Ahmad Z, Waraich EA, Akhtar S, Anjum S, Ahmad T, Mahboob W, Hafeez OBA, Tapera T, Labuschagne M, Rizwan M. Physiological responses of wheat to drought stress and its mitigation approaches. Acta Physiologiae Plantarum. 2018;40:1–13

[53] 53. Dhakal A, Adhikari C, Manandhar D, Bhattarai S, Shrestha S. Effect of abiotic stress in wheat: A review. Reviews in Food and Agriculture. 2021;2:69–72

[54] 54. Hossain A, Skalicky M, Brestic M, Maitra S, Ashraful Alam M, Syed MA, Hossain J, Sarkar S, Saha S, Bhadra P, Shankar T. Consequences and mitigation strategies of abiotic stresses in wheat (Triticum aestivum L.) under the changing climate. Agronomy. 2021;11:241.

[55] 55. Yunus MH, Firdaus A, Khan Z, Ansari MYK. Genomics-assisted breeding (GAB) for trait improvement: Unveiling genomic strategies for accelerated crop enhancement. Plant Breeding Technology: Future Trends and Challenges. 2025;64:138

[56] 56. Abdallah NA, Elsharawy H, Abulela HA, Thilmony R, Abdelhadi AA, Elarabi NI. Multiplex CRISPR/Cas9-mediated genome editing to address drought tolerance in wheat. GM Crops & Food. 2025;16:1–17

[57] 57. Kaya C. Plant metabolic engineering for enhanced nutrition and food security during climate uncertainty. Food and Energy Security. 2025;14:01–21

[58] 58. Mujeeb-Kazi A, Kazi AG, Dundas I, Rasheed A, Ogbonnaya F, Kishii M, Bonnett D, Wang RRC, Xu S, Chen P, Mahmood T. Genetic diversity for wheat improvement as a conduit to food security. Advances in Agronomy. 2013;122:179–257

[59] 59. Renzi JP, Coyne CJ, Berger J, von Wettberg E, Nelson M, Ureta S, Hernández F, Smýkal P, Brus J. How could the use of crop wild relatives in breeding increase the adaptation of crops to marginal environments. Frontiers in Plant Science. 2022;13:01–21

[60] 60. Engels JM, Thormann I. Main challenges and actions needed to improve conservation and sustainable use of our crop wild relatives. Plants. 2020;9:968

[61] 61. Haridass K Exploring the Abiotic Resistance Traits of the Local Wild Wheat Relative, Thinopyrum Distichum (Doctoral dissertation, Stellenbosch University). 2024

[62] 62. Neupane B Evaluation of Winter Wheat and Tritipyrum Germplasm to Identify Useful Variation for Biotic and Abiotic Stress Tolerance (Master’s thesis, North Dakota State University). 2023

[63] 63. Ashraf M, Munns R. Evolution of approaches to increase the salt tolerance of crops. Critical Reviews in Plant Sciences. 2022;41:128–160

[64] 64. Pour-Aboughadareh A, Kianersi F, Poczai P, Moradkhani H. Potential of wild relatives of wheat: Ideal genetic resources for future breeding programs. Agronomy. 2021;11:1656

[65] 65. Rouichi S, Ghanem ME, Amri M. In-situ and ex-situ conservation priorities and distribution of lentil wild relatives under climate change: A modelling approach. Journal of Applied Ecology. 2025;62:414–428

[66] 66. Yao Y, Guo W, Gou J, Hu Z, Liu J, Ma J, Zong Y, Xin M, Chen W, Li Q, Wang Z. Wheat2035: Integrating pan-omics and advanced biotechnology for future wheat design. Molecular Plant. 2025;18:272–297

[67] 67. Saint Ville A, Po JYT, Sen A, Bui A, Melgar-Quiñonez H. Food security and the Food Insecurity Experience Scale (FIES): Ensuring progress by 2030. Food Security. 2019;1:483–491

[68] 68. Curtis T, Halford NG. Food security: The challenge of increasing wheat yield and the importance of not compromising food safety. Annals of Applied Biology. 2014;164:354–372

[69] 69. Bhattacharya J, Currie J, Haider S. Poverty, food insecurity, and nutritional outcomes in children and adults. Journal of Health Economics. 2004;23:839–862

[70] 70. Timmer CP. Food Security and Scarcity: Why Ending Hunger Is so Hard. Philadelphia: University of Pennsylvania Press; 2015

[71] 71. Pérez-Escamilla R. Food and nutrition security definitions, constructs, frameworks, measurements, and applications: Global lessons. Frontiers in Public Health. 2024;12:01–10