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REVIEW article
Front. Genome Ed. , 18 March 2025
Sec. Genome Editing in Plants
Volume 7 - 2025 | https://doi.org/10.3389/fgeed.2025.1533197
This article is part of the Research Topic Genome Editing for Addressing the Challenges of Climate Change Adaptation in Agriculture Crops and Livestock View all articles
Climate change is a global concern for agriculture, food security, and human health. It affects several crops and causes drastic losses in yield, leading to severe disturbances in the global economy, environment, and community. The consequences on important staple crops, such as rice, maize, and wheat, will worsen and create food insecurity across the globe. Although various methods of trait improvements in crops are available and are being used, clustered regularly interspaced short palindromic repeats and CRISPR-associated protein 9 (CRISPR/Cas9) mediated genome manipulation have opened a new avenue for functional genomics and crop improvement. This review will discuss the progression in crop improvement from conventional breeding methods to advanced genome editing techniques and how the CRISPR/Cas9 technology can be applied to enhance the tolerance of the main cereal crops (wheat, rice, and maize) against any harsh climates. CRISPR/Cas endonucleases and their derived genetic engineering tools possess high accuracy, versatile, more specific, and easy to design, leading to climate-smart or resilient crops to combat food insecurity and survive harsh environments. The CRISPR/Cas9-mediated genome editing approach has been applied to various crops to make them climate resilient. This review, supported by a bibliometric analysis of recent literature, highlights the potential target genes/traits and addresses the significance of gene editing technologies in tackling the vulnerable effects of climate change on major staple crops staple such as wheat, rice, and maize.
The world is facing drastic climate change, with high global temperatures and elevated carbon dioxide (CO2) levels. This has resulted in extreme events, adversely affecting all dimensions of the world, including agriculture, biodiversity, and human community. The root cause of climate change is greenhouse gas emissions, mainly from anthropogenic activities, ascending the global surface temperature by 1.5°C since 1850 (Karavolias et al., 2021; Nunez et al., 2019). Recent reports confirmed that the year 2023 was the hottest in Earth’s history, which had several consequences such as drought, compound flooding, heavy precipitation, global sea rise, and upper sea acidification in certain regions of the planet (Li Z. et al., 2024). Rising temperatures are a warning sign for current agricultural production, although they have already started devastating effects worldwide. Climate change will mostly affect crop yield at lower latitudes more than at higher latitudes (Shukla et al., 2019). The situation of agriculture at lower latitudes might worsen with a temperature rise, whereas higher latitudes might benefit from higher temperatures, which would increase crop yield (Iizumi et al., 2018). More specifically, areas closer to the equator are more prone to desertification and eventual agricultural loss, which has already started in the Asian and African continents (Zougmoré et al., 2018). These regions are already facing high populations and unsustainable land management issues; therefore, their agricultural productivity and biodiversity are under great threat due to climate change (Viana et al., 2022). Increasing temperatures rise to over 35°C in California caused the browning of berries, reducing the yield by almost 50% (Kizildeniz et al., 2018).
The current scenario indicates that crop production in tropical areas will be most affected by high temperatures and droughts (Esquivel-Muelbert et al., 2019). It is estimated that food crop production in Africa will be reduced by 2.9% by 2050 owing to climate change. Global productivity losses disrupt other facets of the ecosystem.
The entire ecosystem relies on food supplies, occupying the highest agricultural land share. Studies have revealed that approximately 90% of food calories and 80% of the proteins and fats originate from agricultural land. Hence, agricultural land contributes to food security and various FAO sustainability goals (Avtar et al., 2020; FAO, 2017). However, most arable land has been degraded because of non-sustainable agricultural practices, including spraying chemical fertilizers, excess groundwater use, intensive farming, and deforestation. Such practices have increased greenhouse gas emissions, which are a major cause of temperature increases (Funk, 2021; Shahzad et al., 2021). Therefore, some areas of the globe might experience drought while others might be flooded owing to rising sea levels. Currently, areas suitable for crop production will soon become unsuitable (Iizumi et al., 2018). Therefore, identifying suitable areas for crop production is crucial for addressing the impacts of climate change. Several studies have focused on identifying suitable areas for agriculture in different countries (Musakwa, 2018). However, this alone is insufficient to overcome the effects of climate change.
A meta-analysis of about a hundred studies explained the impacts of climate change on biodiversity and found that a moderate rise in temperature can cause significant harm to biodiversity (Nunez et al., 2019). Owing to climate change pressure, food production needs to be enhanced, which requires more land, exacerbating biodiversity loss. For instance, the production of soybeans, palm oil, beef, and wood from 2000 to 2011 in seven countries was responsible for 40% of the deforestation of tropical forests and carbon losses (Henders et al., 2015; Ortiz et al., 2021).
Despite this, plants adapt extraordinary mechanisms to survive in the harsh climate. Such mechanisms involve root and leaf modification, stomatal regulation, osmotic adjustment, ion transport and sequestration, morphological behavior, and genetic adaptations. However, these processes require years to develop a climate-resilient plant (Krishna et al., 2023). Therefore, dealing with these issues in a short step is feasible using a genome-editing technique called clustered regularly interspaced short palindromic repeats and CRISPR-associated protein 9 (CRISPR/Cas9). Climate-smart crops in terms of increasing abiotic and biotic stress tolerance and high-yielding biofortified crops can be generated using this approach (Figure 1). Here, we summarize the progression in crop improvement from conventional breeding methods to advanced genome editing techniques and how the CRISPR/Cas9 technology can be applied to safeguard the main cereal crops (wheat, rice, and maize) from harsh climates.
Figure 1. Gene editing, a sustainable technology to reduce the Vulnerability of major Staple Crops to Climate Change (Graphical abstract).
Although climate change is progressing at an exceptional rate, it is not easy to envisage the loss it can cause to agriculture. Nevertheless, the scientific community has made tremendous efforts with conventional crop improvement techniques to combat the effects of climate change. Such approaches include breeding that produces superior varieties using donor and recipient plants with desired characteristics (Sharma et al., 2023; Van et al., 2022). This process was revolutionized in 1940–1950 when semi-dwarf wheat varieties were developed. Although hybrid varieties are superior and popular innovations, reaching the final product is time-consuming, costly, and requires intensive labor. Breeding is associated with the plant’s phenotypic trait, which is highly influenced by environmental factors, and requires several backcrosses to obtain the desired trait. Furthermore, conventional breeding methods result in selecting an inferior parent crop limiting the germplasm gene pool and causing genetic erosion (Krishna et al., 2023). Breeding can lead to the development of undesired traits because the transfer of genetic information cannot be controlled (Figure 2). Additionally, large arable land requirements with huge investments are another drawback of conventional breeding. The scientific community is making tremendous efforts using breeding to combat the effects of climate change. However, these efforts are insufficient, and more advanced strategies are required to improve agricultural techniques.
Figure 2. Conventional breeding versus Genome editing: This figure illustrates the advantages of genome editing over conventional breeding techniques. Created with BioRender.
Later, the emergence of recombinant DNA technology, where the genetic material of plants could be modified by inserting a foreign gene of interest to produce superior transgenic crops, also called genetically modified (GM) crops, changed the concept of producing superior varieties. Therefore, genetic manipulation can be performed in a controlled manner. Currently, GM crops, including BT cotton, corn, and soybean, are utilized worldwide, especially in the United States of America (USA) (Wechsler, 2018). Because this approach directly deals with transferring genetic information from one species to another, some myths addressing biodiversity and health concerns remain controversial. In addition, a long procedure that requires clinical trials and money expenditure is required to launch a GM product in the market (Van et al., 2022). Likewise, similar trials are required for genetically engineered plants produced via RNA interference (RNAi) technologies because of its several disadvantages, including, off-targeting effects that may lead to plant toxicity, development of insect resistance, incomplete or variable levels of silencing, and highly programmatic designing process (Sharma et al., 2023).
Gene-editing technologies have attracted the attention of the scientific community owing to their simplicity for designing, efficient editing, and accuracy. Genome editing technology uses molecular scissors to create double-stranded breaks in DNA, and the remaining part is undertaken by the host DNA repair machinery, which can either add or remove nucleotides randomly, leading to the formation of mutants. This modification can be achieved through site-specific insertion-deletion (indels), substitution, or epiallelic changes within the targetted DNA in a cell or organism. Genome editing is based on the principle of DNA repair, in which strand breaks are introduced using molecular scissors, such as endonucleases (Carroll, 2014), mega-nucleases (Gong and Golic, 2003), zinc finger nucleases (ZFNs) (Urnov et al., 2005), transcription activator-like effector nucleases (TALENs) (Sun and Zhao, 2013), and CRISPR/Cas9 (Jinek et al., 2012).
Among the above-mentioned methods CRISPR/Cas endonuclease based is the most popular, which has gained momentum in the last 10 years owing to its high efficiency, ease of use, and accuracy. CRISPR/Cas systems are diverse and adopted from bacteria and archaea. Currently, there are many tools such as CRISPRi, CRISPRa, base editor, gene knock-in, targeted protein tagging, and Viral mediated editing, that arose from the basic CRISPR/Cas9 system (Anzalone et al., 2020; Kampmann, 2018). Variants of Cas endonuclease, i.e., dCas9-foki, Cas9 nickase, HypaCas9, Sniper-Cas9, eSpCas9 (1.1), SpCas9-HF1, xCas9, evoCas9, SuperFi-Cas9, miCas9, evoCjCas9, SpRYc, KG, and SpdNG-QT.12j (Goldberg et al., 2021; Jeon et al., 2018; Karvelis et al., 2021; Kulcsár et al., 2022; Ma et al., 2020; Schmidheini et al., 2024; Schmidt et al., 2021; Schuler et al., 2022; Sun A. et al., 2023; Thakur et al., 2024; Wang et al., 2021a; Zhao et al., 2023). Moreover, Cas9 functional analogs such as, Cas12a-b, Cas12d-f, Cas12h-j, Cas121, Cas12n, Cas12 λ, Cas13 (C2c2), and Cas14 have been developed that extend the editing capability to the RNA and protein levels to enhance the performance of this technique (Hillary and Ceasar, 2022; Schindele et al., 2018; Yan et al., 2019). These all tools have been successfully applied to various crops for different purposes to alter the metabolic pathway (Ahmar and Gruszka, 2023; Kaur et al., 2020; Li D. et al., 2024; Ly et al., 2024; Toinga-Villafuerte et al., 2022; Wang J. D. et al., 2024; Xie Y. et al., 2024).
The detail mechanism of CRISPR/Cas9 for generating knockout events in plants is described in Figure 3. It has been applied to gene editing and transcriptional modulation of plants to improve various agronomical characteristics, such as drought tolerance, salinity tolerance, heat stress tolerance, disease resistance, nutritional enhancement, and yield improvement. Because plant phenology is affected by climate change events, this technology can also be used to control plant development-related factors for instance, flowering, male sterility, and photoperiod) (Cai et al., 2018; Liu et al., 2019; Shen et al., 2017). CRISPR/Cas9 derivatives, including prime and base editors, also provide opportunities to modify the plant genome precisely. Substantial research is currently underway to improve these techniques.
Figure 3. Mechanism of CRISPER/Cas9 in the plant: (1) The CRISPR/Cas9 system can be introduced into plant cells by Agrobacterium mediated transformation, protoplast transformation or particle bombardment. (2) The sgRNA guides the Cas9 to the target region of the genome. (3) Cas9 recognize the PAM region and creates double stranded cut. (4a) InDels are generated in the target site through the NHEJ repair system (4b) Precise corrections can be made in the DNA or directed sequences can be inserted through HDR repair system Created with BioRender.
Therefore, considering the vast potential of gene editing technologies, they may be the best solution for mitigating the effects of climate change (Singh et al., 2024a). This review describes the impact of climate change on food security and how it can affect the main cereal crops (wheat, rice, and maize) that are highly prone to climate change, and the research initiatives for their improvement via the CRISPR/Cas9 approach are also discussed in this review. The applications of this technology are extensive; however, certain limitations such as government regulations must be considered.
Wheat is an important cereal crop that serves two purposes: feed for the global community and support for the nation’s economy. Average wheat production worldwide is over 700 million tons, with China, India, the United States, the Russian Federation, and France being the top five wheat producers. The main exporters of wheat are the United States, Canada, France, Australia, and Russia, whereas the main importers are Egypt, Italy, Brazil, Japan, and Algeria (http://faostat.fao.org/). The overall production and trade of wheat reflects its significant role and demand in the global population. Approximately 95% of the total wheat produced worldwide is hexaploid bread wheat (Triticum aestivum sp. Aestivum, AABBDD, 2n = 6x = 42), whereas the remaining 5% is tetraploid durum wheat (Triticum turgidum sp. Durum, AABB, 2n = 4x = 28), also known as pasta wheat. The dough-forming ability of wheat flour increases the product range of wheat into bread, pasta, noodles, and biscuits. The key components responsible for dough formation are the grain storage proteins called gluten in wheat flour, whose interactions with water upon kneading form a proteinaceous structure (Shewry, 2019). Starch is another important component of wheat that promotes dough formation and causes gelatinization. In total, mature wheat grains contain approximately 13% water, 71% carbohydrates, 11% proteins, 2% lipids, 2% minerals, and 0.1% vitamins and phytochemicals, contributing significantly to human health (Wieser et al., 2020). Wheat has undoubtedly contributed significantly to global food security; however, wheat production is at great risk owing to climate change. As a C3 crop, wheat may benefit from high CO2 concentrations in the environment by improving water-use efficiency, photosynthesis, and transpiration. However, the grain quality can be negatively affected by higher levels of CO2. At higher temperatures, wheat productivity may decline because of a shorter crop season and an increase in transpiration (Bouras et al., 2019). A report published in Nature Climate Change estimated the impact of rising temperatures on global wheat yield using three independent models. The results indicated that the per degree rise in temperature can decline wheat productivity by 4.1%–6.4% (Liu et al., 2016). Various studies have shown that increasing temperature can have a drastic effect on wheat grain quality by shortening the grain-filling period, which will affect the gluten composition of wheat. High temperatures can change the ratio of gliadin to glutenin, weakening the dough-making properties of wheat flour. Moreover, extended heat waves and high temperatures can reduce the nitrogen level and, ultimately, the protein content of the grain (Wang and Liu, 2021b). In conclusion, climate change is going to impact wheat; therefore, the development of climate-tolerant wheat, which is difficult through conventional breeding techniques due to its hexaploidy genome, is an utmost priority. CRISPR/Cas9 technology was successfully used to obtain stable inherited mutations in wheat. Agrobacterium-mediated transformation has been used to deliver CRISPR constructs into immature wheat embryos (Howells et al., 2018; Zhang Z. et al., 2019). CRISPR/Cas9 genome editing-derived transgene-free wheat plants have also been produced using biolistic and protoplast transfection methods (Liang et al., 2017; Zhang et al., 2016). These studies have successfully implemented CRISPR/Cas9 genome editing technology in wheat, providing a platform for improving wheat concerning climate change-mediated problems. Grain weight, grain size, and grain yield per plant are the main agronomic attributes highly influenced by abiotic and biotic factors. Researchers have identified several genes that function as negative regulators of grain’s weight, size, and overall yield. For example, Receptor-like protein kinase 1 (RPK1), the Brittle rachis gene (BTR1-A), GASR7, TaSPL13, GW2, and TaARE1 (Table 1). A study reported the knockout of GASR7 and GW2 genes via the ribonucleoprotein (RNP)-derived CRISPR/Cas9 approach in bread wheat and pasta wheat resulted in transgene-free mutants with higher grain weight (Zhang Y. et al., 2021). Similarly, the TaSPL13 gene, responsible for controlling grain size and number, was edited using CRISPR/Cas9 (Gupta et al., 2023). The results revealed that mutations in this gene lead to an increase in the size and number of grains in allohexaploid wheat, demonstrating the importance of TaSPL13 in the evolution of yield-related attributes in wheat. Similarly, Ta-eIF4E alone has been targeted in wheat, resulting in viral resistance and improved plant height and grain length (Kan et al., 2023). Another important agronomic factor in wheat is early heading. Recently, the role of the thioredoxin gene (TaTRXH9) is characterized and validated in wheat using CRISPR/Cas9-mediated gene editing, and a loss-of-function mutation in this gene found in early-heading wheat (Fan et al., 2023). These studies reveal the success of CRISPR/Cas9 genome editing in improving wheat yield, which can significantly mitigate the impact of climate change on yield reduction.
The unstable weather can increase gluten levels in wheat, which makes it less suitable for consumption (Mkhabela et al., 2022). However, Chinese researchers have developed a Gluten gene Enrichment and Sequencing (GlutEnSeq) system, screened thousands of prolamin genes from different wheat varieties, and low gluten wheat was produced by modifying γ- and a-gliadin genes via CRISPR/Cas9 (Jouanin et al., 2019). A study developed improved winter and spring wheat varieties with high amylose content via CRISPR/Cas9-mediated genome editing of starch-branching enzyme (SBE) II (TaSBEIIa), and multiple transgene-free mutant lines were obtained (Li J. et al., 2021). Correspondingly, the CRISPR/Cas9 approach has been utilized to modify four genes, i.e., puroindoline b (PINb), granule-bound starch synthase gene (GBSS or WAXY), polyphenol oxidase gene (PPO), and phytoene synthase gene (PSY), in wheat via the Agrobacterium-mediated transformation method of gene delivery (Zhang Y. et al., 2021; Zhang Z. et al., 2019). The PINb gene controls grain hardness, the WAXY gene is responsible for amylose synthesis, PSY is the main gene controlling the carotenoid biosynthetic pathway, and PPO controls color via the oxidation of phenolic compounds.
Micronutrient deficiency in wheat is likely to occur because of increasing temperatures and drought, which can lead to malnutrition and other health disorders. The CRISPR/Cas9 genome editing approach is a straightforward method that can be used to enhance the nutritional composition of wheat and address this problem; for instance, Ibrahim et al. edited the Inositol Pentakisphosphate 2-Kinase 1 (TaIPK1) gene, which expresses an enzyme involved in the final step of the phytate biosynthesis pathway (Ibrahim et al., 2022). Phytic acid is considered an anti-nutrient because it reduces the bioavailability of iron and zinc in humans. The phytic in wheat tissues binds to these micronutrients as phytate and decreases their bioavailability. CRISPR/Cas9 genome engineering of TaIPK1 in wheat improved iron (1.5–2.1-fold) and zinc content (1.6–1.9-fold) and lowered phytic acid accumulation in wheat grains. Several agronomic improvements via CRISPR/Cas9-mediated genome editing in wheat are presented in Table 1.
Transcriptome profiling of wheat grains revealed several heat stress-associated gene, including heat shock transcription factor gene (TaHSFA6e), ascorbate peroxidase, β-amylase, γ-gliadin-2, and LMW-glutenin, were upregulated during the high-temperature stress (Rangan et al., 2020; Wen et al., 2023). Such studies can help select the potential target genes for CRISPR/Cas-mediated genome editing, which might lead abiotic stress resistant crops. Moreover, studies on different crops can be used to develop wheat that is tolerant to increasing temperatures caused by climate change. For example, CRISPR/Cas9-mediated knockout of the mitogen-activated protein kinase gene (SlMAPK 3) enhances heat tolerance in tomato plants, and the edited pyrabactin resistance 1 (PYR1)/PYR1-like (PYL) (pyl1/4/6) makes rice tolerant to hot weather (Miao et al., 2018; Yu et al., 2019). Since wheat is hexaploid and has a complex genome, there are fewer reports available on producing abiotic stress-tolerant wheat plants than rice. TaSAL1, TaMBF1c, and TaHAG1 genes have been edited to produce abiotic stress-tolerant wheat plants (Mohr et al., 2022; Tian et al., 2022; Zheng M. et al., 2021). The histone acetyltransferase (TaHAG1) gene contributes to salt tolerance by affecting free radical production in hexaploid wheat. MBF1c confers thermotolerance by regulating the translation of specific mRNA translation, whereas SAL1 negatively regulates drought tolerance (Table 1). These studies suggest that genome editing of wheat could be a promising approach for conferring climate change.
Rice (Oryza sativa L.) is a staple crop, with a global consumption rate of approximately 21% and 76% in Asian countries. Worldwide, 776.5 million tons of rice were produced in 2022, with approximately 90.5% of the average rice production taking place in Asia, in which China and India are the biggest producers (). This high demand for rice is due to its taste and versatility in a variety of international cuisines (Castanho et al., 2023). In addition, rice is enriched in nutrients, mainly complex carbohydrates, and moderate levels of vitamin B, phosphorous, iron, calcium, and protein. Most nutrients (minerals, vitamins, proteins, and antioxidants) are present in the rice brain, which has a brown outer layer. Rice contains all essential amino acids except lysine and is a great source of a balanced diet, as it does not contain cholesterol, fat, or sodium (Sasaki and Burr, 2000). However, the current climate change scenarios have adversely affected rice crops from farms to consumers in various ways (Zhao et al., 2016). Increasing temperature, carbon dioxide, drought, salinity, rainfall, pests, and diseases are the main stressors that can directly affect various rice attributes, such as grain size, quality, yield, nutritional constitution, and appearance. Rice cultivation can be successful if the optimum temperature and rainfall are provided; however, an increase of 1 °C can adversely affect rice yield. A recent study showed a decline in the nutritional content of rice due to rising atmospheric CO2 concentrations. Micronutrients, proteins, and several vitamins are diminished in rice, leading to malnutrition in infants and children of rice-dependent countries. A decline of approximately 17%–30% in vitamins, 5% in zinc, 8% in iron, and 10% in protein was observed in rice grown under high CO2 conditions (Smith and Myers, 2018). In contrast, Guo et al. found a 15% increase in the mineral content of rice grown under high CO2 and temperature conditions (Guo et al., 2022). Though high CO2 is normally considered a growth-stimulating agent along with high temperature, it acts antagonistically. Jing et al. reported reduced rice yield due to elevated temperatures under high CO2 conditions created using temperature-free air CO2 enrichment (T-FACE) systems (Jing L. et al., 2024). Another T-FACE experiment in China reported that increasing CO2 substandardized the sensory quality of rice by increasing chalkiness (Wang et al., 2024d). The chalkiness of rice is a major obstacle to rice marketing. Chalky rice forms owing to abnormal starch development inside the grain and appears as a scattered coarse material, thus making the surface turbid. This abnormal starch accumulation occurs due to the high temperature, mainly at the time of grain filling, which reduces the ability of α-amylase enzyme to degrade the starch and, hence, leads to malformed starch synthesis (Shimoyanagi et al., 2021). In addition to grain quality, other issues in rice that can occur due to climate change include decreased shoot biomass and carbohydrate content in the stem, decreased starch content, and reduced photosynthetic ability, transpiration, and leaf area (Cui et al., 2024; Huanhe et al., 2024; Yamori et al., 2025). Hence, concerning the future outcomes, researchers have been trying different strategies, one of which is the robust and versatile genetic editing technique called as CRISPR/Cas9 system for generating climate-smart rice crops. CRISPR/Cas9-mediated gene editing has been optimized in rice using Agrobacterium-mediated transformation, particle bombardment, and RNPs (Miao et al., 2013; Shan et al., 2013). Moreover, a multiplexing approach was successfully established for rice. Recently, a group of researchers accomplished ultra-multiplexing targeting 49 genes in rice using both Agrobacterium-mediated and biolistic approaches (Wu et al., 2024).
CRISPR/Cas9 mediated-knockout of two main quantitative trait loci associated with grain length and thousand-grain weight, i.e., GS3 and GL3.1 was reported in rice GS3 encodes a protein that restrains the cell division of the spikelet hull, leading to shorter grains, whereas GL3.1 encodes an enzyme that controls grain size by dephosphorylating the cell cycle-related protein Cyclin-T1,3 and inhibits cell proliferation in the hull. Mutations in this gene increase grain size in rice (Yuyu et al., 2020; Zhang Y. M. et al., 2021). Huang et al. used the CRISPR/Cas9 technique to create an Indica maintainer line, Mei1B, containing an edited GS3 allele to improve grain yield and quality (Huang et al., 2022). Another study mutated the OsSPL16 or GW8 gene via CRISPR/Cas9 technology to improve the cylindrical shape and grain yield of Basmati rice (Usman et al., 2021). Rice aroma is an important quality parameter that is affected by climate change. Nevertheless, improving and introducing aromas into rice is feasible to improve and introduce aromas into rice using genetic engineering techniques. Mutations in Betaine Aldehyde Dehydrogenase 2 (OsBADH2) via CRISPR/Cas9 added aroma to elite non-aromatic rice variety (Hui et al., 2022; Tang et al., 2021). Several reports are available on the use of CRISPR-mediated gene editing for improving various agronomic traits in rice (Table 2). Recently, loss-of-function mutants of OsCKX were found to affect various attributes including plant height, grain size, grain number, panicle size, seed shape, and starch accumulation. This gene encodes a cytokinin-degrading enzyme that inactivates cytokinins, which plays important roles in plant growth and cell proliferation (Zheng et al., 2023). Similarly, enhanced grain yield was observed by deleting a target site of the transcription factor An-1 in the cis-regulatory region of the Ideal Plant Architecture 1 (IPA1) gene (Song et al., 2022).
Rice quality improvement is crucial because rice passes through various downstream processes, such as dehydration, milling, removal of bran, cleaning, and cooking after harvesting, and each process directly or indirectly decreases the nutrient content of rice. For instance, cleaning alone can reduce vitamin levels by 25%–60%, potassium by 20%–40%, and proteins by 3%–7% in rice (Müller et al., 2022). Combining this with the effect of climate change, the rice produced would not be useful. Therefore, biofortification is the only method that can sustain rice nutrients. Carotenoid-enriched marker-free rice has been developed by inserting two carotenoid biosynthetic genes, SSU-CRTI and ZmPSY30, using the CRISPR/Cas9 approach (Dong et al., 2020). Achary and Reddy produced a rice variety with an enhanced accumulation of iron, zinc, potassium, calcium, and phosphorous in endosperm via CRISPR/Cas9-mediated editing of GW2 (Grain width and grain weight) locus (Achary and Reddy, 2021). Moreover, the aleurone layer gained thickness with an enhanced protein content. Another study found decreased grain chalkiness, high ammonium cation, and phosphate ion uptake, and high photosynthetic activity under high CO2 conditions in miR166-RDD1 knocked-out rice plants (Iwamoto, 2022).
Among abiotic stressors, salinity is a serious event that can potentially tarnish rice production. The development of salt-tolerant varieties is a lifesaving approach feasible with CRISPR/Cas9 genome editing technology. High salt concentrations can negatively affect crop production by disrupting the metabolic and physiological processes. These factors can significantly affect plant development, seed germination, and productivity (Zörb et al., 2019). Plants respond to salt stress by increasing the biosynthesis of antioxidants, osmoregulators, and phytohormones that support the plant by maintaining ion homeostasis; however, this ability works to a certain extent, and not all plants can protect themselves from high salinity (Raza et al., 2022). Similarly, drought stress drastically affects the physiology of plants by inhibiting nutrient uptake and other life-dependent activities, including photosynthesis, cell division and elongation, turgor pressure, and gene expression. Several rice genes have been identified and mutated using CRISPR/Cas9 to produce abiotic stress-tolerant plants (Table 2). For example, salt-tolerant T2 homozygous mutant rice was developed by cleaving the OsRR22 gene in rice using Cas9 (Zhang A. et al., 2019). Knockout of OsRR22 improved the performance of rice plants under high-salt conditions (0.75%), with no side effects on grain size, yield, or plant biomass. The same strategy was used by Han et al. to develop novel rice germplasm at the seedling stage (Han et al., 2022). In another study, the drought and salt tolerance (OsDST) gene was edited using CRISPR/Cas9 to produce the indica mega rice cultivar MTU1010 with enhanced tolerance (Santosh et al., 2020). In rice, the transcription factor OsMADS26 plays a negative role, and mutations in this gene enhance drought tolerance (Anjala and Augustine, 2022). Ogata et al. characterized the Enhanced Response to ABA1 (ERA1) gene in rice using CRISPR/Cas9-mediated editing and found frameshift mutations in mutants with increased primary root growth, high sensitivity to abscisic acid stress, and increased drought stress tolerance (Ogata et al., 2020).
The effect of climate change on disease susceptibility in rice poses a major threat to rice productivity. Climate change-induced pest and microbial emergence can annihilate agriculture, thereby substantially threatening food security. Fortunately, this can be mitigated using gene-editing applications in crops. Various studies have been published on the production of pests and microorganisms that cause disease tolerance in crops (Liu M. et al., 2024; Oliva et al., 2019; Zhou et al., 2022). Table 2 shows some examples of CRISPR/Cas9-mediated development of disease-resistant plants. However, the current work is insufficient in comparison to the upcoming consequences of climate change because this change can also increase the potential of insects by providing them with favorable conditions for their growth and development. For example, fruit flies flourished more when the temperature increased from 20°C to 35 °C in certain mango varieties. This phenomenon is observed in all species worldwide. In particular, insects in temperate regions have become more active, whereas populations of tropical insects may decline or migrate (Bhattacharjee et al., 2022). Therefore, genetically edited plants may be a powerful solution for overcoming the effects of climate change on agriculture. Several studies have been conducted to develop rice resistance to various fungal and bacterial pathogens. Xa13 is involved in pollen development and exhibits recessive resistance to bacterial blight. Studies have shown that the complete loss of function of the coding region of this gene can lead to sterility; therefore, CRISPR-assisted modification was performed in the Xa13 promoter region to produce transgene-free bacterial blight-resistant rice with retained fertility (Li C. et al., 2020). Another study targeted three salicylic acid 5-hydroxylase (OsS5H) genes (BSR-D1, PI21, and ERF922) in rice and found resistance to both rice blast and bacterial blight (Zhou et al., 2022). Recently, rice with enhanced immunity was produced by editing the NAC transcription factor gene in rice via CRISPR/Cas9 approach (Son et al., 2024). Furthermore, broad-spectrum disease resistance was achieved by editing three salicylic acid hydroxylase (OsS5H) genes in rice (Li X. et al., 2023).
Maize (Zea mays) is the most important cereal crop in the world, with the highest production after rice and wheat, and fulfills the needs of human food, animal feed, and biofuels (Chávez-Arias et al., 2021). Maize is sensitive to heat stress during seed germination and vegetative growth. It significantly affects maize plant germination and seedling emergence. Heat stress causes the formation of abscisic acid and affects the activity of enzymes responsible for breaking down starch (Chandra et al., 2023). Additionally, it inhibits the synthesis of proteins in the embryo, which reduces the germination of maize seeds at over 37°C, resulting in a decrease in plant density (Buriro et al., 2011). Increased oxidative stress, altered membrane permeability, decreased stomatal conductance, and other signs occur regularly in plants under heat stress. The rate of photosynthesis was negatively affected by a reduction in stomatal conductance. Moreover, it induces the production of reactive oxygen species and causes oxidative stress (Soengas et al., 2018; Zandalinas et al., 2017). Heat stress negatively affects the number of florets, silk number, fertilization, filling, development, and final grain yield during flowering (Lizaso et al., 2018). Recently, the continuous span of heat waves has affected maize yield and productivity. They alter the morphology, physiology, genomic expression, and biochemical metabolism of crops. In response to these alterations, plants activate tolerance mechanisms via heat shock transcription factors and proteins essential for reducing and preventing heat-related damage (Li and Zhang, 2022d). Heat stress affects the integrity of the plasma membrane and the accumulation of reactive oxygen species (Dogra and Kim, 2020). Moreover, proteins are misfolded or unfolded, which disrupts cell metabolism and physiology and eventually leads to cell death. The development of climate resilience in maize is urgently needed because heat-induced decline is high in this crop (Chandra et al., 2023).
Currently, a decrease in yield resulting from drought stress is estimated to affect over 20% of the annual maize area, and at high temperatures, an average of 7.4% is lost for every 1°C increase (Boyer et al., 2013; Malenica et al., 2021). Notably, Brazil, the third largest producer of maize worldwide, showed a decrease in yield in the 2015–16 and 2020–21 growing seasons of approximately 18 and 23 Mt, correspondingly to losses of approximately 21% compared to the 2014–15 and 2019–20 seasons (Lopes Filho et al., 2023). These crop failures occurred in years marked by extreme drought conditions, resulting in diminished yields in many of the largest producer geographies. Similarly, crop yield reductions and spiking prices were affected by the 2012 drought in the US (the world’s largest producer) (Boyer et al., 2013). Therefore, it is essential to reduce the potential losses caused by the increased frequency, severity, and duration of stresses associated with global climate change by continuously developing new maize cultivars that target better genetic adaptation and using improved agricultural practices.
Abiotic stress tolerance and yield are two complex traits strongly influenced by environmental factors and linked to small-effect genetic loci. Using genomic engineering techniques to develop superior cultivars for these traits is more difficult because of such complexity, which makes it challenging to reliably evaluate the molecular mechanisms behind gene activities and measure phenotypes. Considering the few instances developed for complex traits, transgenic maize cultivars with enhanced herbicide and insect resistance have been on the market for decades (Yassitepe et al., 2021). The difficulty in applying a transgenic approach to control complex traits that are stable in multiple environments has limited the development of biotech cultivars that could be widely used (Simmons et al., 2021).
Gene editing technology has enabled an efficient and consistent way to understand the role of key genes and develop new germplasm in maize (Doll et al., 2019; Wang Y. et al., 2022).
Potential putative functions of numerous genes involved in maize development programs and stress responses have been investigated thoroughly using CRISPR/Cas9 technology (Wang Y. et al., 2022). CRISPR has been widely used to enhance numerous agronomic traits in maize, yield, nutrition, improved pollen characteristics, drought tolerance, and disease resistance (Jiang L. et al., 2024; Kaul et al., 2024; Lv et al., 2024; Wang G. et al., 2024; Xie Z. et al., 2024). ZmGDIɑ was specifically edited using CRISPR/Cas9 to significantly increase maize resistance to the maize rough dwarf virus without negative agronomic effects (Liu Y. et al., 2023). ZmCOIɑ interacts inversely with ZmJAZ15 to alter maize immunity against Gibberella stalk rot (GSR, a teleomorph of Gibberella zeae), and further downregulation of ZmCOIɑ can increase maize resistance to GSR (Ma et al., 2021). MMS21 maintains the activity and integrity of the maize genome, resulting in improved root and vegetative growth, pollen germination, and seed development (Zhang et al., 2021b). ZmCLCg positively regulates sodium chloride stress and chloride transport in maize as a stress response (Luo et al., 2021). ZmSRL5 is essential for sustaining cuticular wax structure and drought tolerance in maize (Pan Y. et al., 2020).
Maize yield is severely affected by several environmental factors, including drought, high temperatures, floods, and unsuitable soil conditions. The breeding of stress-tolerant variants has shown great potential when applying genome editing tools compared with conventional breeding methods (Chávez-Arias et al., 2021; Chennakesavulu et al., 2021; Prasanna et al., 2021). The development of maize lines with high stalk strength has become considerably important to breeders for maintaining high and constant production, as stalk lodging caused by different environmental factors poses a significant threat to maize quality and production. STIFF1 is a negative regulator of maize stalk length, its altered allele with a 2bp deletion caused a frameshift and an early slowdown translation, conferring CRISPR-edited plants with a stronger stalk, which contributed to high-density planting and avoided stalk lodging (Armarego-Marriott, 2020). Furthermore, ZmGA20OX3 has been modified to develop semi-dwarf maize plants using CRISPR/Cas9 technology, which may be useful for developing a novel genotype that is more resilient to lodging and suitable for high-density planting (Liu Y. et al., 2023). For drought tolerance, precisely editing the promoter sequence of ARGOS8 leads to an increase in its expression and enhances maize grain yield under drought stress (Shi et al., 2017). By inserting the GOS2 promoter from maize plants at the 5ʹ untranslated regions that remain of the ARGOS8 gene’s native promoter, which acts as a negative regulator of ethylene responses (Zafar et al., 2020). Targeted alteration of the native maize promoter using CRISPR/Cas9 enhanced ARGOS8 expression and improved grain production under drought conditions. In addition, CRISPR/Cas9 has been used to target Slagamous-Like 6 (SIAGL6) to achieve heat tolerance (Doll et al., 2019). Therefore, developing new germplasm sources for breeding stress-tolerant maize has been achieved using CRISPR/Cas9 technology. Table 3 illustrates several improvements in the agronomic traits of maize using CRISPR/Cas9 gene editing.
A bibliometric analysis was conducted to examine global trends in CRISPR/Cas9 research, focusing on cereal crops and their applications for stress tolerance and yield improvement. The search used terms like “CRISPR,” “Genome editing,” “Gene editing,” and “Gene silencing,” along with crop-related terms such as “Cereal crops” and stress-related topics like “Abiotic stress,” “Biotic stress,” and “Drought tolerance.” The dataset, initially containing 1,232 articles from 2019 to 2025, was narrowed to 597 after excluding reviews, book chapters, and non-English publications. The citation analysis and other visualizations were conducted using Web of Science (WoS) and VOSviewer.
According to the database, increasing trend in publications and citations from 2019 to 2025 can be seen, demonstrating a significant increase in scholarly interest in CRISPR/Cas9 for enhancing crop stress tolerance and yield (Figure 4). After 2021, both publications and citations remained steady, suggesting the field’s maturation or a shift in research focus, with sustained interest continuing in the subsequent years (Francis et al., 2024). The tree map in Figure 5 organizes research across different subject areas, with Plant Sciences comprising the largest share, followed by biochemistry, molecular biology and agronomy. This distribution emphasizes the central focus on plant traits, particularly in cereals, while also highlighting the broader applications of CRISPR/Cas9 in fields like biochemistry and biotechnology. Smaller categories, such as Horticulture and Environmental Sciences, reflect the interdisciplinary nature of genome editing research (AlRyalat et al., 2019).
Figure 4. Annual publication trends from 2019 to 2025, showing the number of papers published each: The graph illustrates a surge in publications and citations from 2019 to 2025 in the area of CRISPR/Cas9 for enhancing crop stress tolerance and yield Created with BioRender.
Figure 5. The tree map illustrates research across different subject areas: This map focuses on research across different subject areas, with Plant Sciences comprising the largest share, followed by biochemistry, molecular biology and agronomy Created with BioRender.
The network visualization in Figure 6 further shows the relationships between key research themes. The central nodes CRISPR and genome editing are closely connected to other significant areas such as drought tolerance, yield improvement, and stress resistance, with a primary focus on rice, a key cereal crop. This network highlights the broad applications of CRISPR/Cas9 in enhancing abiotic stress tolerance and improving crop yield. The strong links between these themes suggest that improving crop resilience is a major goal in CRISPR/Cas9 research (Altaf et al., 2024).
Figure 6. Network visualization of keyword co-occurrence related to CRISPR/Cas9 research in cereals: The central nodes in the figure illustrates the relationship between CRISPR and genome editing specially in the research areas such as drought tolerance, yield improvement, and stress resistance, with a primary focus on rice. Visualization uses different colors to distinguish between research areas, with red highlighting drought tolerance, green representing stress resistance, and blue focusing on gene editing Created with BioRender.
The overlay visualization, tracks how research trends have changed over time, especially focusing on drought tolerance, salinity tolerance, and abiotic stress (Figure 7). From 2021 onward, the research has increasingly focused on these topics, particularly in cereal crops like rice, wheat, and maize. This shift reflects the growing need for developing climate-resilient crops, with more research being dedicated to improving stress resistance and water use efficiency in cereals. The overlay emphasizes the growing interest in improving cereals to make them more resilient and higher yielding under extreme environmental conditions. While Figure 6 shows the connections between topics, Figure 7 illustrates the evolution of these topics over time, especially highlighting cereals as a key focus in the search for more resilient crops to face climate change (Altaf et al., 2024).
Figure 7. The overlay visualization of research trends on CRISPR/Cas9 over time: The figure shows the research trends over time, especially focusing on drought tolerance, salinity tolerance, and abiotic stress. Since 2021, the research has increasingly focused on these topics, particularly in cereal crops like rice, wheat, and maize Created with BioRender.
Furthermore, the country distribution map, highlights the geographic spread of research on CRISPR/Cas9 applications in cereal crops (Figure 8). The heatmap reveals that countries such as China, United States, and India are at the forefront of this research, with other countries like South Korea, Brazil, and France contributing significantly (Martins et al., 2022). However, there is a noticeable gap in regions that rely heavily on cereal crops, such as the Middle East. This suggests an opportunity for further research in these areas, particularly to address local agricultural challenges related to climate stress and water scarcity.
Figure 8. Country-wise distribution of publications on CRISPR/Cas9 research in cereals from 2019 to 2025: The heatmap reveals that countries such as China, United States, and India are at the forefront of this research, with other countries like South Korea, Brazil, and France contributing significantly Created with BioRender.
Gene editing has been successfully used to generate climate-resilient crops for various climatic conditions. However, several limitations are descending its overall potential remains limited (Singh et al., 2024a; Singh et al., 2024b). The major drawback of gene-editing technologies is off-target, which can cause unwanted editing of other genes, which hinders their wide applicability for crop trait improvement.
Another disadvantage of, the lack of efficient tissue culture methods for regeneration, transformation and generation of gene edited crops. The use of a de novo meristem induction technique can be more useful and easier for recalcitrant crop species (Maher et al., 2020). Research is being conducted to increase the transformation ability of recalcitrant varieties using advanced tools to produce climate-resilient crops.
Apart from these technical drawbacks, policymakers and regulatory authorities must take the initiative to overcome the lack of clarity regarding genome-edited crops among the population. Altogether, these factors can provide ultimate success in applying these technologies to address the impact of climate change.
Intragenic, transgenic, and cisgenic (ICT) approaches have been used to improve plant characteristics using foreign genes which leverage its applications (Karavolias et al., 2021; Klümper and Qaim, 2014; Steinwand and Ronald, 2020).
Therefore, merging genome editing with ICT approaches would be the best solution for raising climate-smart crops and this can be possible using targeted gene integration using CRISPR/Cas9 technology. This strategy has been applied to maize varieties by successfully inserting a novel promoter upstream of a gene responsible for ethylene regulation to improve drought tolerance. Moreover, the entire gene can be replaced using these approaches; for example, the replacement of the japonica NRT1.1B allele with the indica allele improves nitrogen use efficiency in rice (Li et al., 2018). Although these approaches have great potential, regulations regarding these technologies have decreased their feasibility. Therefore, new advancements and technical improvements are required to overcome these limitations.
For example, base editors and prime editors are second-generation CRISPR-based genome modification tools that mediate precise editing without relying on double-stranded break formation and homology direct repair. These editors are more precise in terms of single-nucleotide modification and integration (Lin et al., 2020). Base editors involve the direct conversion of a single-nucleotide base into another (A-to-G or C-to-T, and A-to-C or C-to-G), without forming double-strand breaks, introducing specific point mutations with utmost precision. Base editing has been applied to several crops including Arabidopsis, cotton, rice, tomato, maize, tobacco, and soyabean (Li X. et al., 2024; Luo et al., 2023; Wang G. et al., 2024; Wang et al., 2024f; Wei et al., 2023; Zhong D. et al., 2024). On the other hand, prime editors integrate Cas9 nickase and a reverse transcriptase, prime editing guide RNA (pegRNA) which is a combination of Cas9 sgRNA, a reverse transcriptase template, and a primer-binding site (PBS). The pegRNA guides the nCas9 to the target site, where it makes a nick in the non-target DNA strand. Then the reverse transcriptase extends the nicked strand by utilizing the reverse transcriptase template (RTT) from the pegRNA, thereby incorporating the intended modifications. With prime editors, large deletion, replacement, and inversion of larger DNA fragments can be performed in plants with high precision. Researchers have achieved DNA inversions of up to 205.4 kb in wheat plants with 51.5% by using dual prime editors. They have also been applied to edit large DNA fragments in tobacco and tomato (Zhao et al., 2025). Similarly, a recent study utilized high-efficiency prime-editing tools to knockin a 10-bp heat-shock element (HSE) into promoters of cell-wall-invertase genes (CWINs) in rice and tomato cultivars (Lou et al., 2025). These modified CRISPR tools can leverage the gene editing efficiency of manipulating chromosomes and larger DNA segments for crop improvement.
Gene editing could be a powerful solution for the present and future anticipation of climate change consequences. With the emergence of advanced genome editing techniques, including CRISPR/Cas9, base editing, and prime editing, various agronomic traits such as disease resistance, abiotic stress tolerance, and nutritional enhancement have emerged. Despite this, most gene editing technologies are still under laboratory research and have not yet been translated into the real world. This is due to technical limitations and restrictions imposed by regulatory authorities and policymakers. However, technological innovations are rapidly expanding owing to the ongoing efforts of public and private institutions. The potential of gene editing in offering solutions for climate change in agriculture is not overlooked, even though it is not the only solution to improve agriculture. Numerous studies show that gene editing can be used to enhance agriculture and combat climate change effects greatly. Nevertheless, as indicated by bibliometric analysis, significant research gaps remain, particularly in applying CRISPR/Cas9 to underexplored crops like rice, wheat and maize for comprehensive climate resilience.
NK: Formal Analysis, Investigation, Methodology, Software, Writing–original draft. MQ: Data curation, Investigation, Methodology, Writing–original draft. DF: Writing–original draft, Formal Analysis, Software. AA: Software, Validation, Writing–review and editing. ST: Conceptualization, Supervision, Validation, Writing–review and editing. ZA: Conceptualization, Data curation, Formal Analysis, Funding acquisition, Investigation, Methodology, Project administration, Software, Supervision, Writing–review and editing.
The author(s) declare that financial support was received for the research, authorship, and/or publication of this article. The authors are grateful to United Arab Emirates University (UAEU) for providing funding through the UAEU Program for Advanced Research (UPAR) program, grant number 12F059.
The authors are thankful to the UAEU University facilities and library for supporting the research and collection of the most recent literature.
The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.
The author(s) declared that they were an editorial board member of Frontiers, at the time of submission. This had no impact on the peer review process and the final decision.
The author(s) declare that no Generative AI was used in the creation of this manuscript.
All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.
The Supplementary Material for this article can be found online at: https://www.frontiersin.org/articles/10.3389/fgeed.2025.1533197/full#supplementary-material
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BADH2 Betaine Aldehyde Dehydrogenase 2
BE Base editing
CO2 Carbon dioxide
CRISPR/Cas Clustered Regularly Interspaced Short Palindromic Repeats and CRISPR-Associated System
DST Drought and salt tolerance gene
ERA1 Enhanced Response to ABA1 gene
FAO Food and Agriculture Organization
GBSS Granule-bound starch synthase gene
GlutEnSeq Gluten gene Enrichment and Sequencing
GM Genetically Modified
GMOs Genetically Modified Organisms
gRNA Guide RNA
GSR Gibberella stalk rot
HAG1 Histone acetyltransferase gene
HSFA6e Heat shock transcription factor gene
ICT Intragenic, transgenic, and cisgenic approaches
IPA Ideal Plant Architecture 1 gene
IPK1 Inositol Pentakisphosphate 2-Kinase 1 gene
IUCN International Union for the Conservation of Nature
MLO Mildew resistance locus O
PAM Protospacer Adjacent Motif
PFT1 Phytochrome and flowering time 1 gene
PINb Puroindoline b gene
PPO Polyphenol oxidase gene
PSY Phytoene synthase gene
PYR1 Pyrabactin resistance 1 gene
RNAi RNA interference
RNP Ribonucleoprotein
S genes Susceptible genes
S5H Salicylic acid 5-hydroxylase genes
SBEIIa Starch-branching enzyme (SBE) II gene
SIAGL6 Slagamous-Like 6 gene
SlMAPK 3 Mitogen-activated protein kinase gene
TALENs Transcription activator-like effector nucleases
T-FACE Temperature-free air CO2 enrichment systems
TFs Transcription factors
TMT3B Tonoplast monosaccharide transporter 3 gene
TRM TONNEAU 1-recruiting motif
TRXH9 Thioredoxin gene
USA United States of America
WSSMV Wheat spindle streak mosaic virus
WYMV Wheat yellow mosaic virus
ZFNs Zinc finger nucleases
Keywords: climate-smart crops, food security, genome editing, maize, rice, wheat
Citation: Kaur N, Qadir M, Francis DV, Alok A, Tiwari S and Ahmed ZFR (2025) CRISPR/Cas9: a sustainable technology to enhance climate resilience in major Staple Crops. Front. Genome Ed. 7:1533197. doi: 10.3389/fgeed.2025.1533197
Received: 23 November 2024; Accepted: 27 February 2025;
Published: 18 March 2025.
Edited by:
Koppolu Raja Rajesh Kumar, Indira Gandhi National Tribal University, IndiaReviewed by:
Rushil Ramesh Mandlik, Tamil Nadu Agricultural University, IndiaCopyright © 2025 Kaur, Qadir, Francis, Alok, Tiwari and Ahmed. This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.
*Correspondence: Zienab F. R. Ahmed, zienab.ahmed@uaeu.ac.ae
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