سازوکارهای فیزیولوژیک و مولکولی تحمل به شوری در غلات: 2- روش‌های به‌نژادی پیشرفته و چشم‌انداز آینده

نوع مقاله : مقاله مروری

نویسندگان

1 استادیار پژوهش، مرکز ملی تحقیقات شوری، سازمان تحقیقات، آموزش و ترویج کشاورزی، یزد، ایران

2 دانشیار، گروه زراعت و اصلاح نباتات، دانشکده کشاورزی، دانشگاه یاسوج، یاسوج، ایران

3 استاد، گروه تولید و ژنتیک گیاهی، دانشکده کشاورزی، دانشگاه شیراز، شیراز، ایران

4 دانشیار پژوهش، مرکز ملی تحقیقات شوری، سازمان تحقیقات، آموزش و ترویج کشاورزی، یزد، ایران

10.22124/cr.2026.32537.1886

چکیده

مقدمه: شوری، به‌عنوان یکی از مهم‌ترین عوامل محدودکننده رشد و تولید محصولات زراعی استراتژیک مانند گندم، برنج و ذرت، امنیت غذایی جهانی را به‌خطر انداخته است. با توجه به هزینه‌بر و زمان‌بر بودن اصلاح فیزیکی خاک‌های شور، توسعه رقم‌های متحمل از طریق به‌نژادی، موثرترین و اقتصادی‌ترین راه‌کار برای مقابله با این چالش محسوب می‌شود. در این راستا، درک سازوکارهای مولکولی و ژنتیکی تحمل به شوری برای تولید ژنوتیپ‌های جدید و متحمل به شوری ضروری است. هدف از مطالعه حاضر، مرور پیشرفت‌های اخیر و بررسی کاربرد فناوری‌های نوین به‌نژادی به‌منظور  شتاب‌بخشی به توسعه رقم‌های متحمل به شوری در گیاهان زراعی به‌ویژه غلات بوده است.
 
مواد و روش‌ها: در این مطالعه، به‌طور اختصاصی کاربرد نظام‌مند زیست‌فناوری‌های نوین از جمله ادغام ابزارهای اومیکس، نقشه‌یابی ژنومی، انتخاب به‌کمک نشانگر و ویرایش ژنوم به‌منظور انتقال مؤثر ژن‌ها و QTLهای شناسایی شده به ژنوتیپ‌های برتر و تسریع برنامه‌های به‌نژادی تشریح شده است.

نتایج و بحث: گزینش به‌کمک نشانگرهای مولکولی، روشی کارآمد در به‌نژادی است که به‌جای در نظر گرفتن فقط فنوتیپ، امکان انتخاب ژنوتیپ‌های برتر را با استفاده از الگوهای نواربندی DNA در مراحل اولیه رشد موجود زنده فراهم می‌کند. این روش با کاهش تأثیرپذیری از محیط، دقت و سرعت برنامه‌های به‌نژادی را افزایش داده و دوره به‌نژادی را که در روش‌های کلاسیک ممکن است در حدود هشت تا ده سال طول بکشد، به‌طور چشم‌گیری کاهش می‌دهد. کاربرد موفق گزینش به‌کمک نشانگر در انتقال ­QTLهایی نظیر Saltol برای افزایش تحمل به شوری در برنج اثبات شده است. همچنین، نقش مؤثر این روش در محصولاتی مانند گندم و ذرت در غلبه بر تنش‌های غیرزیستی از جمله تنش شوری نشان داده است. با این‌حال، این روش برای صفات کمی پیچیده که توسط ژن‌ها یا ­QTLهای با اثرات کوچک کنترل می‌شوند، کارآیی محدودتری دارد. امروزه، رویکردهای پیشرفته‌تری مانند گزینش ژنومی (Genomic Selection) و ویرایش ژنوم مبتنی بر CRISPR/Cas9 همراه با فنوتیپ‌سازی با توان عملیاتی بالا (High-Throughput Phenotyping)، به‌عنوان راه‌کارهای مکمل برای افزایش دقت و سرعت برنامه‌های به‌نژادی به‌منظور ایجاد رقم‌های متحمل به تنش‌های محیطی نظیر شوری به‌کار گرفته می‌شوند.

نتیجه‌گیری: ابزارهای نوین به‌نژادی گیاهی مانند انتخاب به‌کمک نشانگر، مطالعات ارتباطی در سطح ژنوم و به‌ویژه فناوری‌های اُمیکس (ترانسکریپتومیکس، پروتئومیکس و متابولومیکس) و ویرایش ژنوم، انقلابی در فرآیند شناسایی و انتقال ژن‌های مطلوب به گیاهان زراعی ایجاد کرده‌اند. این فناوری‌ها امکان هرمی کردن ژن‌ها (Pyramiding Genes) و انتقال همزمان چندین ژن تحمل به شوری را با دقت و سرعت بالا فراهم می‌سازند.

کلیدواژه‌ها

موضوعات


عنوان مقاله [English]

Physiological and molecular mechanisms of salinity tolerance in cereals: II. Advanced breeding methods and future perspectives

نویسندگان [English]

  • Ahmad Majidimehr 1
  • Reza Amiri-Fahliani 2
  • Bahram Heidari 3
  • Gholamhassan Ranjbar 4
1 Research Assistant Professor, National Salinity Research Center, Agricultural Research, Education and Extension Organization (AREEO), Yazd, Iran.
2 Associate Professor, Department of Agronomy and Plant Breeding, Faculty of Agriculture, Yasouj University, Yasouj, Iran
3 Professor, Department of Plant Production and Genetics, Faculty of Agriculture, Shiraz University, Shiraz, Iran
4 Research Associate Professor, National Salinity Research Center, Agricultural Research, Education and Extension Organization (AREEO), Yazd, Iran
چکیده [English]

Introduction: Salinity, as one of the most significant limiting factors for the growth and production of strategic crops such as wheat, rice, and maize, poses a serious threat to global food security. Given the high cost and time-consuming nature of physical remediation of saline soils, developing tolerant varieties through breeding programs is the most effective and economical approach to addressing this challenge. In this regard, understanding the molecular and genetic mechanisms of salt tolerance is essential for generating new, salt-tolerant genotypes. The aim of the present study was to review recent advances and investigate the application of modern breeding technologies to accelerate the development of salt-tolerant varieties in crop plants, particularly cereals.
 
Materials and Methods: This review study systematically describes the application of novel biotechnologies including the integration of omics tools, genomic mapping, marker-assisted selection (MAS), and genome editing to efficiently transfer identified genes and QTLs into elite genotypes and to expedite breeding programs.

Results and Discussion: Marker-assisted selection is an efficient breeding method that, instead of relying solely on phenotype, enables the selection of superior genotypes using DNA banding patterns at early stages of organism development. By reducing environmental influence, this approach enhances the accuracy and speed of breeding programs and significantly shortens the breeding cycle, which in classical methods may take approximately eight to ten years. The successful application of MAS has been demonstrated in transferring QTLs such as Saltol to improve salt tolerance in rice. Furthermore, the effective role of this method has been shown in crops such as wheat and maize for overcoming abiotic stresses, including salinity. However, MAS has limited efficiency for complex quantitative traits controlled by genes or QTLs with small effects. Today, more advanced approaches such as genomic selection (GS), CRISPR/Cas9-based genome editing, and high-throughput phenotyping (HTP) are being employed as complementary strategies to enhance the precision and speed of breeding programs aimed at developing varieties tolerant to environmental stresses such as salinity.

Conclusion: Modern plant breeding tools such as marker-assisted selection, genome-wide association studies (GWAS), and particularly omics technologies (transcriptomics, proteomics, and metabolomics) as well as genome editing have revolutionized the process of identifying and transferring desirable genes into crop plants. These technologies enable gene pyramiding and the simultaneous transfer of multiple salt-tolerance genes with high precision and speed.

کلیدواژه‌ها [English]

  • CRISPR
  • Genome editing
  • High-throughput phenotyping
  • Marker-assisted selection
  • Metabolomics
  • Transcriptome
Abdul Aziz, M., & Masmoudi, K. (2023). Insights into the transcriptomics of crop wild relatives to unravel the salinity stress adaptive mechanisms. International Journal of Molecular Sciences, 24(12), 9813. doi: 10.3390/ijms24129813.##Afzal, M., Alghamdi, S. S., Nawaz, H., Migdadi, H. H., Altaf, M., El-Harty, E., Al-Fifi, S. A., & Sohaib, M. (2022). Genome-wide identification and expression analysis of CC-NB-ARC-LRR (NB-ARC) disease-resistant family members from soybean (Glycine max L.) reveal their response to biotic stress. Journal of King Saud University-Science, 34(2), 101758. doi: 10.1016/j.jksus.2021.101758.##Atta, K., Mondal, S., Gorai, S., Singh, A. P., Kumari, A., Ghosh, T., Roy, A., Hembram, S., Gaikwad, D. J., Mondal, S., Bhattacharya, S., Jha,U. C., & Jespersen, D. (2023). Impacts of salinity stress on crop plants: Improving salt tolerance through genetic and molecular dissection. Frontiers in Plant Science14, 1241736. doi: 10.3389/fpls.2023.1241736.##Barkla, B. J., Vera-Estrella, R., Hernández-Coronado, M., & Pantoja, O. (2009), Quantitative proteomics of the tonoplast reveals a role for glycolytic enzymes in salt tolerance. Plant & Cell, 21(12), 4044-4058. doi: 10.1105/tpc.109.069211.##Bernardo, R. (2016). Bandwagons I, too, have known. Theoretical & Applied Genetics129(12), 2323-2332.‏ doi: 10.1007/s00122-016-2772-5.##Cao, Y., Zhang, M., Liang, X., Li, F., Shi, Y., Yang, X., & Jiang, C. (2020). Natural variation of an EF-hand Ca2+-binding protein coding gene confers saline-alkaline tolerance in maize. Nature Communications, 11(1), 186. doi: 10.1038/s41467-019-14027-y.##Chen, M., Zhu, X., Liu, X., Wu, C., Yu, C., Hu, G., Chen, L., Chen, R., Bouzayen, M., Zouine, M., & Hao, Y. (2021a). Knockout of auxin response factor SlARF4 improves tomato resistance to water deficit. International Journal of Molecular Sciences, 22(7), 3347. doi: 10.3390/ijms22073347.##Chen, S., Zhang, N., Zhou, G., Hussain, S., Ahmed, S., Tian, H., & Wang, S. (2021b). Knockout of the entire family of AITR genes in Arabidopsis leads to enhanced drought and salinity tolerance without fitness costs. BMC Plant Biology, 21(1), 137. doi: 10.1186/s12870-021-02907-9.##Chinnusamy, V., Jagendorf, A., & Zhu, J. K. (2005). Understanding and improving salt tolerance in plants. Crop Science, 45, 437-448. doi: 10.2135/cropsci2005.0437.##Coppens, F., Wuyts, N., Inzé, D., & Dhondt, S. (2017). Unlocking the potential of plant phenotyping data through integration and data-driven approaches. Current Opinion in Systems Biology, 4, 58-63. doi: 10.1016/j.coisb.2017.07.002.##Cramer, G. R., Urano, K., Delrot, S., Pezzotti, M., & Shinozaki, K. (2011). Effects of abiotic stress on plants: A systems biology perspective. BMC Plant Biology11(1), 163. doi: 10.1186/1471-2229-11-163.##Deery, D. M., & Jones, H. G. (2021). Field phenomics: Will it enable crop improvement? Plant Phenomics, 2021, 9871989. doi: 10.34133/2021/9871989.##de Los Campos, G., Hickey, J. M., Pong-Wong, R., Daetwyler, H. D., & Calus, M. P. (2013). Whole-genome regression and prediction methods applied to plant and animal breeding. Genetics193(2), 327-345. doi: 10.1534/genetics.112.143313.##Dorostkar, S., Dadkhodaie, A., Ebrahimie, E., Heidari, B., & Ahmadi-Kordshooli, M. (2022). Comparative transcriptome analysis of two contrasting resistant and susceptible Aegilops tauschii accessions to wheat leaf rust (Puccinia triticina) using RNA-sequencing. Scientific Reports, 12(1), 821. doi: 10.1038/s41598-021-04329-x.##Duangjit, J., Causse, M., & Sauvage, C. (2016). Efficiency of genomic selection for tomato fruit quality. Molecular Breeding36(3), 29. doi: 10.1007/s11032-016-0453-3.##Endelman, J. B., Atlin, G. N., Beyene, Y., Semagn, K., Zhang, X., Sorrells, M. E., & Jannink, J. L. (2014). Optimal design of preliminary yield trials with genome-wide markers. Crop Science54(1), 48-59. doi: 10.2135/cropsci2013.03.0154.##Etesami, H., & Maheshwari, D. K. (2018). Use of plant growth promoting rhizobacteria (PGPRs) with multiple plant growth promoting traits in stress agriculture: Action mechanisms and future prospects. Ecotoxicology & Environmental Safety, 156, 225-246. doi: 10.1016/j.ecoenv.2018.03.013.##Fernandez‑Gallego, J. A., Kefauver, S. C., Gutiérrez, N. A., Nieto‑Taladriz, M. T., & Araus, J. L. (2018). Wheat ear counting in‑field conditions: High throughput and low‑cost approach using RGB images. Plant Methods, 14(22), 1-12. doi: 10.1186/s13007-018-0289-4.##FAO (2023). FAOSTATE. Agricultural Statistics. Food and Agriculture Organization of the United Nations. Rome, Italy. https://www.fao.org/faostat/en/#home.##Formentin, E., Sudiro, C., Perin, G., Riccadonna, S., Barizza, E., Baldoni, E., Lavezzo, E., Stevanato, P., Sacchi, G. A., Fontana, P., Toppo, S., Morosinotto, T., Zottini, M., & Lo Schiavo, F. (2018). Transcriptome and cell physiological analyses in different rice cultivars provide new insights into adaptive and salinity stress responses. Frontiers in Plant Science, 9, 204. doi: 10.3389/fpls.2018.00204.##Gaj, T., Gersbach, C. A., & Barbas, C. F. (2013). ZFN, TALEN, and CRISPR/Cas-based methods for genome engineering. Trends in Biotechnology, 31(7), 397-405. doi: 10.1016/j.tibtech.2013.04.004.##Gharaghanipor, N., Arzani, A., Rahimmalek, M., & Ravash, R. (2022). Physiological and transcriptome indicators of salt tolerance in wild and cultivated barley. Frontiers in Plant Science13, 819282. doi: 10.3389/fpls.2022.819282.##Gilliham, M., Able, J. A., & Roy, S. J. (2017). Translating knowledge about abiotic stress tolerance to breeding programmes. The Plant Journal90(5), 898-917. doi: 10.1111/tpj.13456.##Guo, G., Ge, P., Ma, C., Li, X., Lv, D., Wang, S., Ma, W., & Yan, Y. (2012). Comparative proteomic analysis of salt response proteins in seedling roots of two wheat varieties. Journal of Proteomics, 75(6), 1867-1885. doi: 10.1016/j.jprot.2011.12.032.##Haque, M. A., Rafii, M. Y., Yusoff, M. M., Ali, N. S., Yusuff, O., Datta, D. R., Anisuzzaman, M., & Ikbal, M. F. (2021). Advanced breeding strategies and future perspectives of salinity tolerance in rice. Agronomy11(8), 1631. doi: 10.3390/agronomy11081631.##Hong, M. J., Ko, C. S., Kim, J. B., & Kim, D. Y. (2024). Identification and transcriptomic profiling of salinity stress response genes in colored wheat mutant. PeerJ, 12, e17043. doi: 10.7717/peerj.17043.##Hu, P., Zheng, Q., Luo, Q., Teng, W., Li, H., Li, B., & Li, Z. (2021). Genome-wide association study of yield and related traits in common wheat under salt-stress conditions. BMC Plant Biology21(1), 27. doi: 10.1186/s12870-020-02799-1.##Hu, Y., Knapp, S., & Schmidhalter, U. (2020). Advancing high-throughput phenotyping of wheat in early selection cycles. Remote Sensing12(3), 574. doi: 10.3390/rs12030574.##Huang, L., Wu, D. Z., & Zhang, G. P. (2020). Advances in studies on ion transporters involved in salt tolerance and breeding crop cultivars with high salt tolerance. Journal of Zhejiang University-Science B21(6), 426-441. doi: 10.1631/jzus.B1900510.##Huang, Q. N., Shi, Y. F., Zhang, X. B., Song, L. X., Feng, B. H., Wang, H. M., Xu, X., Li, X.H., Guo, D., & Wu, J. L. (2016). Single base substitution in OsCDC48 is responsible for premature senescence and death phenotype in rice. Journal of Integrative Plant Biology, 58(1), 12-28. doi: 10.1111/jipb.12372.##Jaganathan, D., Ramasamy, K., Sellamuthu, G., Jayabalan, S., & Venkataraman, G. (2018). CRISPR for crop improvement: An update review. Frontiers in Plant Science, 9, 985. doi: 10.3389/fpls.2018.00985.##Jahan, N., Raihan, M. S., Islam, M. M., Era, F. M., Alalawy, A. I., Omran, A. M., Alanazi, Y. F., Sakran, M., Alasmari, A., Alzuaibr, F., El Sabagh, A., Kahrizi, D., & Islam, A. K. M. A. (2024). Genome-wide association studies of salinity tolerance in local aman rice. Cellular & Molecular Biology, 70(2), 10-17. doi: 10.14715/cmb/2024.70.2.2.##Jha, S., Maity, S., Singh, J., Chouhan, C., Tak, N., & Ambatipudi, K. (2022). Integrated physiological and comparative proteomics analysis of contrasting genotypes of pearl millet reveals underlying salt‐responsive mechanisms. Physiologia Plantarum, 174(1), e13605. doi: 10.1111/ppl.13605.##Joseph, B., & Jini, D. (2010). Proteomic analysis of salinity stress-responsive proteins in plants. Asian Journal of Plant Sciences9(6), 307. doi: 10.3923/ajps.2010.307.313.##Kamal, A. H., Kihyun, K., Kwang-Hyun, S., Jong-Soon, C., Byung-Kee, B., Tsujimoto, H., Hwa-Young, H., Chul-Soo, P., & Sun-Hee, W. (2010). Abiotic stress responsive proteins of wheat grain determined using proteomics technique. Australian Journal of Crop Science, 4, 196-208.##Kamburova, V. S., Nikitina, E. V., Shermatov, S. E., Buriev, Z. T., Kumpatla, S. P., Emani, C., & Abdurakhmonov, I. Y. (2017). Genome editing in plants: An overview of tools and applications. International Journal of Agronomy2017(1), 7315351. doi: 10.1155/2017/7315351.##Katori, T., Ikeda, A., Iuchi, S., Kobayashi, M., Shinozaki, K., Maehashi, K., Sakata, Y., Tanaka, S., & Taji, T. (2010). Dissecting the genetic control of natural variation in salt tolerance of Arabidopsis thaliana accessions. Journal of Experimental Botany, 61(4), 1125-1138. doi: 10.1093/jxb/erp376.##Kausar, R., & Komatsu, S. (2022). Proteomic approaches to uncover salt stress response mechanisms in crops. International Journal of Molecular Sciences, 24(1), 518. doi: 10.3390/ijms24010518.##Khalid, M., Kausar, R., Shahzad, A., Ali, G. M., & Begum, S. (2023). Screening and validation of salt-stress responsive cg-SSR markers in wheat (Triticum aestivum L.) germplasm of Pakistan. Molecular Biology Reports50(7), 5931-5940. doi: 10.1007/s11033-023-08519-w.##Kordrostami, M., Rabiei, B., & Hassani Kumleh, H. (2017). Biochemical, physiological and molecular evaluation of rice cultivars differing in salt tolerance at the seedling stage. Physiology & Molecular Biology of Plants, 23(3), 529-544. doi: 10.1007/s12298-017-0440-0.##Koyama, M. L., Levesley, A., Koebner, R. M., Flowers, T. J., & Yeo, A. R. (2001). Quantitative trait loci for component physiological traits determining salt tolerance in rice. Plant Physiology125(1), 406-422. doi: 10.1104/pp.125.1.406.##Kumar, P., Choudhary, M., Halder, T., Prakash, N. R., Singh, V. V. T., Sheoran, S., Ravikiran, K. T., Longmei, N., Rakshit, S., & Siddique, K. H. M. (2022). Salinity stress tolerance and omics approaches: Revisiting the progress and achievements in major cereal crops. Heredity128(6), 497-518. doi: 10.1038/s41437-022-00516-2.##Kumar, V., Singh, A., Mithra, S. A., Krishnamurthy, S. L., Parida, S. K., Jain, S., Tiwari, K., Kumar, P., Rao, A. R., Sharma, S. K., Khurana, J. P., Singh, N. K., & Mohapatra, T. (2015). Genome-wide association mapping of salinity tolerance in rice (Oryza sativa). DNA Research22(2), 133-145. doi: 10.1093/dnares/dsu046.##Li, Y., Wu, X., Zhang, Y., & Zhang, Q. (2022). CRISPR/Cas genome editing improves abiotic and biotic stress tolerance of crops. Frontiers in Genome Editing, 4, 987817. doi: 10.3389/fgeed.2022.987817.##Lowe, R., Shirley, N., Bleackley, M., Dolan, S., & Shafee, T. (2017). Transcriptomics technologies. PLoS Computational Biology, 13(5), e1005457. doi: 10.1371/journal.pcbi.1005457.##Lu, Y., Li, M., Gao, Z., Ma, H., Chong, Y., Hong, J., Wu, J., Wu, D., Xi, D., & Deng, W. (2025). Advances in whole genome sequencing: Methods, tools, and applications in population genomics. International Journal of Molecular Sciences26(1), 372. doi: 10.3390/ijms26010372.##Luo, X., Wang, B., Gao, S., Zhang, F., Terzaghi, W., & Dai, M. (2019). Genome‐wide association study dissects the genetic bases of salt tolerance in maize seedlings. Journal of Integrative Plant Biology61(6), 658-674. doi: 10.1111/jipb.12797.##Malzahn, A., Lowder, L., & Qi, Y. (2017). Plant genome editing with TALEN and CRISPR. Cell & Bioscience7(1), 21. doi: 10.1186/s13578-017-0148-4.##Mani, B. R., Kumar, B. M., & Krishnamurthy, S. L. (2014). Genetic variability and diversity of rice landraces of South Western India based on morphological traits. ORYZA-An International Journal of Rice51(4), 261-266. doi: 10.1038/s41598-025-03547-x.##Mansoor, S., Karunathilake, E. M., Tuan, T. T., & Chung, Y. S. (2025). Genomics, phenomics, and machine learning in transforming plant research: Advancements and challenges. Horticultural Plant Journal11(2), 486-503. doi: 10.1016/j.hpj.2023.09.005.##Marcos, M., Sharifi, H., Grattan, S. R., & Linquist, B. A. (2018). Spatio-temporal salinity dynamics and yield response of rice in water-seeded rice fields. Agricultural Water Management195, 37-46. doi: 10.1016/j.agwat.2017.09.016.##Mardani, Z., Rabiei, B., Sabouri, H., & Sabouri, A. (2014). Identification of molecular markers linked to salt-tolerant genes at germination stage of rice. Plant Breeding, 133(2), 196-202. doi: 10.1111/pbr.12136.##Marè, C., Zampieri, E., Cavallaro, V., Frouin, J., Grenier, C., Courtois, B., Brorrier, L., Tacconi, G., Finocchiaro, F., Serrat, X., Nogues, S., Bundo, M., Segundo, B. S., Negrini, N., Pesenti, M., Sacchi, G. A., Gavina, G., Bovina, R., Monacon, S., Tondelli, A., Cattivelli, L., & Valè, G. (2023). Marker-assisted introgression of the salinity tolerance locus Saltol in temperate Japonica rice. Rice16(1), 2.‏ 10.1186/s12284-023-00619-2.##Meuwissen, T.  H., Hayes, B. J., & Goddard, M. (2001). Prediction of total genetic value using genome-wide dense marker maps. Genetics, 157(4), 1819-1829. doi: 10.1093/genetics/157.4.1819.##Mohamed, H. I., Khan, A., & Basit, A. (2024). CRISPR-Cas9 system mediated genome editing technology: an ultimate tool to enhance abiotic stress in crop plants. Journal of Soil Science & Plant Nutrition, 24(2), 1799-1822. doi: 10.1007/s42729-024-01778-x.##Muthu, V., Abbai, R., Nallathambi, J., Rahman, H., Ramasamy, S., Kambale, R., Thulasinathan, T., Ayyenar, B.,  & Muthurajan, R. (2020). Pyramiding QTLs controlling tolerance against drought, salinity, and submergence in rice through marker assisted breeding. PloS One15(1), e0227421. doi: 10.1371/journal.pone.0227421.##Mutz, K. O., Heilkenbrinker, A., Lönne, M., Walter, J. G., & Stahl, F. (2013). Transcriptome analysis using next-generation sequencing. Current Opinion in Biotechnology, 24(1), 22-30. doi: 10.1016/j.copbio.2012.09.004.##Mwando, E., Han, Y., Angessa, T. T., Zhou, G., Hill, C. B., Zhang, X. Q., & Li, C. (2020). Genome-wide association study of salinity tolerance during germination in barley (Hordeum vulgare L.). Frontiers in Plant Science11, 00118. doi: 10.3389/fpls.2020.00118.##Naveed, S. A., Zhang, F., Zhang, J., Zheng, T. Q., Meng, L. J., Pang, Y. L., Xu, J. L., & Li, Z. K. (2018). Identification of QTN and candidate genes for salinity tolerance at the germination and seedling stages in rice by genome-wide association analyses. Scientific Reports, 8(1), 6505. doi: 10.1038/s41598-018-24946-3.##Nayyeripasand, L., Garoosi, G. A., & Ahmadikhah, A. (2021). Genome-wide association study (GWAS) to identify salt-tolerance QTLs carrying novel candidate genes in rice during early vegetative stage. Rice, 14(1), 9. doi: 10.1186/s12284-020-00433-0.##Nematzadeh, Gh. A., & Kiani, Gh. (2011). Plant Breeding. Volume 2. Molecular Methods. Publication of Sari Agricultural Sciences and Natural Resources University, Sari, Iran. 336 p. [In Persian].##Ndunge, M. M. (2021). Evaluation of agronomic traits among tropical maize under salt stress and identification of responsible saltol quantitative trait locus. Ph. D.  Dissertation. Kenyatta University, The Kenya.##Nongpiur, R. C., Singla-Pareek, S. L., & Pareek, A. (2016). Genomics approaches for improving salinity stress tolerance in crop plants. Current Genomics, 17(4), 343-357. doi: 10.1104/pp.011445.##Nongpiur, R. C., Singla-Pareek, S. L., & Pareek, A. (2020). The quest for osmosensors in plants. Journal of Experimental Botan, 71, 595-607. doi: 10.1093/jxb/erz263.##Nutan, K. K., Singla-Pareek, S. L., & Pareek, A. (2020). The Saltol QTL-localized transcription factor OsGATA8 plays an important role in stress tolerance and seed development in Arabidopsis and rice. Journal of Experimental Botany, 71(2), 684-698. doi: 10.1093/jxb/erz368.##Pang, Y., Chen, K., Wang, X., Wang, W., Xu, J., Ali, J., & Li, Z. (2017). Simultaneous improvement and genetic dissection of salt tolerance of rice (Oryza sativa L.) by designed QTL pyramiding. Frontiers in Plant Science8, 1275. doi: 10.3389/fpls.2017.01275.##Poland, J., Endelman, J., Dawson, J., Rutkoski, J., Wu, S., Manes, Y., Dreisigacker, S., Crossa, J., Sanchez-Villeda, H., Sorrells, M., & Jannink, J. L. (2012). Genomic selection in wheat breeding using genotyping-by-sequencing. The Plant Genome, 5(3), 10.3835/plantgenome2012.06.0006.##Poole, N., Donovan, J., & Erenstein, O. (2022). Continuing cereals research for sustainable health and well-being. International Journal of Agricultural Sustainability, 20(5), 693-704. doi: 10.1080/14735903.2021.1975437.##Pruthi, R. (2024). Leveraging the genomic tools to explore the molecular basis of salinity tolerance in rice and soybean. LSU Doctoral Dissertations. 6548. doi: 10.31390/gradschool_dissertations.6548.##Pythoud, F. (2004). The cartagena protocol and GMOs. Nature Biotechnology22(11), 1347-1348.##Roitsch, T., Cabrera-Bosquet, L., Fournier, A., Ghamkhar, K., Jiménez-Berni, J., Pinto, F., & Ober, E. S.  (2019). Review: New sensors and data-driven approaches-A path to next generation phenomics. Plant Science, 282, 2-10. doi: 10.1016/j.plantsci.2019.01.011.##Saini, H., Devrani, A., Synrem, G., & Priyanka. (2025). Application of CRISPR technology in plant improvement: An update review. Advances in Agriculture, 2025, 4578877. 10.1155/aia/4578877.##Satyavathi, C. T., Tomar, R. S., Ambawat, S., Kheni, J., Padhiyar, S. M., Desai, H., Bhatt, S. B., Shitap, M. S., Meena, R. C., Singhal, T., Sankar, M., Singh, S. P., & Khandelwal, V. (2022). Stage specific comparative transcriptomic analysis to reveal gene networks regulating iron and zinc content in pearl millet [Pennisetum glaucum (L.) R. Br.]. Scientific Reports, 12(1), 276. doi: 10.1038/s41598-021-04388-0.##Shan, Q., Wang, Y., Li, J., Zhang, Y., Chen, K., Liang, Z., Zhang, K., Liu, J., Xi, J. J., Qiu, J. L., & Gao, C. (2013). Targeted genome modification of crop plants using a CRISPR-Cas system. Nature Biotechnology, 31(8), 686-688. doi: 10.1038/nbt.2650.##Sheng, X., Ai, Z., Tan, Y., Hu, Y., Guo, X., Liu, X., Sun, Z., Yu, D., Chen, J., Tang, N., Duan, M., & Yuan, D. (2023). Novel salinity-tolerant third-generation hybrid rice developed via CRISPR/Cas9-mediated gene editing. International Journal of Molecular Sciences, 24(9), 8025. doi: 10.3390/ijms24098025.##Singh, A. K., Pal, P., Sahoo, U. K., Sharma, L., Pandey, B., Prakash, A., Sarangi, P. K., Prus, P., Pașcalău, R., & Imbrea, F. (2024). Enhancing crop resilience: The role of plant genetics, transcription factors, and next-generation sequencing in addressing salt stress. International Journal of Molecular Sciences, 25(23), 12537. doi: 10.3390/ijms252312537.##Singh, P., Kumar, A., & Borthakur, A. (2019). Abatement of Environmental Pollutants: Trends and Strategies. First Edition. Elsevier. doi: 10.1016/C2018-0- 03174-6.##Singh, V., Krause, M., Sandhu, D., Sekhon, R. S., & Kaundal, A. (2023). Salinity stress tolerance prediction for biomass-related traits in maize (Zea mays L.) using genome-wide markers. The Plant Genome, 16(4), e20385. doi: 10.1002/tpg2.20385.##Sobhanian, H., Aghaei, K., & Komatsu, S. (2011). Changes in the plant proteome resulting from salt stress: Toward the creation of salt-tolerant crops? Journal of Proteomics74(8), 1323-1337. doi: 10.1016/j.jprot.2011.03.018.##Song, P., Wang, J., Guo, X., Yang, W., & Zhao, C. (2021). High-throughput phenotyping: Breaking through the bottleneck in future crop breeding. The Crop Journal, 9(3), 633-645. doi: 10.1016/j.cj.2021.03.015.##Soares, A. L., Geilfus, C. M., & Carpentier, S. C. (2018). Genotype-specific growth and proteomic responses of maize toward salt stress. Frontiers in Plant Science, 9, 661. doi: 10.3389/fpls.2018.00661.##Sukumaran, S., Jarquin, D., Crossa, J., & Reynolds, M. (2018). Genomic-enabled prediction accuracies increased by modeling genotype × environment interaction in durum wheat. Plant Genome, 11(2), 30025014. doi: 10.3835/plantgenome2017.12.0112.##Supplitt, S., Karpinski, P., Sasiadek, M., & Laczmanska, I. (2021). Current achievements and applications of transcriptomics in personalized cancer medicine. International Journal of Molecular Sciences, 22(3), 1422. doi: 10.3390/ijms22031422.##Tak, Y. G., & Farnham, P. J. (2015). Making sense of GWAS: Using epigenomics and genome engineering to understand the functional relevance of SNPs in non-coding regions of the human genome. Epigenetics & Chromatin8(1), 57. doi: 10.1186/s13072-015-0050-4.##Trono, D., & Pecchioni, N. (2022). Candidate genes associated with abiotic stress response in plants as tools to engineer tolerance to drought, salinity and extreme temperatures in wheat: An overview. Plants, 11(23), 3358. doi: 10.3390/plants11233358.##Turan, S., Cornish, K., & Kumar, S. (2012). Salinity tolerance in plants: breeding and genetic engineering. Australian Journal of Crop Science6(9), 1337-1348.##Walsh, J. J., Mangina, E., & Negrão, S. (2024). Advancements in imaging sensors and AI for plant stress detection: A systematic literature review. Plant Phenomics, 6, 0153. doi: 10.34133/plantphenomics.0153.##Wang, M., Wang, Y., Zhang, Y., Li, C., Gong, S., Yan, S., Li, G., Hu, G., Ren, H., Yang, J., Yu, T., & Yang, K. (2019). Comparative transcriptome analysis of salt-sensitive and salt-tolerant maize reveals potential mechanisms to enhance salt resistance. Genes & Genomics, 41(7), 781-801. doi: 10.1007/s13258-019-00793-y.##Wang, X., Ren, P., Ji, L., Zhu, B., & Xie, G. (2022a). OsVDE, a xanthophyll cycle key enzyme, mediates abscisic acid biosynthesis and negatively regulates salinity tolerance in rice. Planta, 255(1), 6. doi: 10.1007/s00425-021-03802-1.##Wang, Y., Zafar, N., Ali, Q., Manghwar, H., Wang, G., Yu, L., Ding, X., Ding, F., Hong, N., Wang, G., & Jin, S. (2022b). CRISPR/Cas genome editing technologies for plant improvement against biotic and abiotic stresses: Advances, limitations, and future perspectives. Cells, 11(23), 3928. doi: 10.3390/cells11233928.##Witzel, K., & Mock, H. P. (2016). A proteomic View of the Cereal and Vegetable Crop Response to Salinity Stress. In: Hosseini Salekdeh, G. (Ed.). Agricultural Proteomics. Volume 2. Environmental Stresses. Springer, Cham. pp. 53-69. doi: 10.1007/978-3-319-43278-6_3.##Würschum, T. (2019). Modern Field Phenotyping Opens New Avenues for Selection. In: Miedaner, T., & Korzun, V. (Eds.). Applications of Genetic and Genomic Research in Cereals. Woodhead Publishing. pp. 233-250. doi: 10.1016/B978-0-08-102163- 7.00011-9.##Xu, C., Tang, X., Shao, H., & Wang, H. (2016). Salinity tolerance mechanism of economic halophytes from physiological to molecular hierarchy for improving food quality. Current Genomics17(3), 207-214. doi: 10.2174/1389202917666160202215548.##Xu, N., Lu, B., Wang, Y., Yu, X., Yao, N., Lin, Q., Xu, X., & Lu, B. (2023). Effects of salt stress on seed germination and respiratory metabolism in different Flueggea suffruticosa genotypes. PeerJ11, e15668. doi: 10.7717/peerj.15668.##Yin, K., Gao, C., & Qiu, J. L. (2017). Progress and prospects in plant genome editing. Nature Plants3(8), 1-6. doi: 10.1038/nplants.2017.107.##Yang, W., Feng, H., Zhang, X., Zhang, J., Doonan, J. H., Batchelor, W. D., Xiong, L., Yan, J., & Yan, J. (2020). Crop phenomics and high-throughput phenotyping: Past decades, current challenges, and future perspectives. Molecular Plant, 13(2), 187-214. doi: 10.1016/j.molp.2020.01.008.##Yousaf, M. S., Ahmad, I., Anwar-ul-Haq, M., Siddiqui, M. T., Khaliq, T., & Berlyn, G. P. (2020). Morphophysiological response and reclamation potential of two agroforestry tree species (Syzygium cumini and Vachellia nilotica) against salinity. Pakistan Journal of Agricultural Sciences57(5), 10026. doi: 10.21162/PAKJAS/20.10026.##Yu, J., Zhu, C., Xuan, W., An, H., Tian, Y., Wang, B., Chi, W., Chen, G.,  Ge, Y., Li, J., Dai, Z., Liu, Y., Sun, Z., Xu, D., Wang, C., & Wan, J. (2023). Genome-wide association studies identify OsWRKY53 as a key regulator of salt tolerance in rice. Nature Communications, 14(1), 3550. doi: 10.1038/s41467-023-39167-0.##Zeng, Y., Wen, J., Zhao, W., Wang, Q., & Huang, W. (2020). Rational improvement of rice yield and cold tolerance by editing the three genes OsPIN5b, GS3, and OsMYB30 with the CRISPR-Cas9 system. Frontiers in Plant Science10, 1663. doi: 10.3389/fpls.2019.01663.##Zhang, A., Liu, Y., Wang, F., Li, T., Chen, Z., Kong, D., Bi, J., Zhang, F., Luo, X., Wang, J., Tang, J., Yu, X., Liu, G., & Luo, L. (2019). Enhanced rice salinity tolerance via CRISPR/Cas9-targeted mutagenesis of the OsRR22 gene. Molecular Breeding, 39(3), 47. doi: 10.1007/s11032-019-0954-y.##Zhang, A., Sun, H., Wang, P., Han, Y., & Wang, X. (2012). Modern analytical techniques in metabolomics analysis. Analyst137(2), 293-300.##Zhang, X., Pérez-Rodríguez, P., Semagn, K., Beyene, Y., Babu, R., López-Cruz, M. A., Vicente, F. S., Olsen, M., Buckler, E., Jannink, J. L., Prasanna, B. M., & Crossa, J. (2015). Genomic prediction in biparental tropical maize populations in water-stressed and well-watered environments using low-density and GBS SNPs. Heredity114(3), 291-299. doi: 10.1038/hdy.2014.99.##Zheng, M., Lin, J., Liu, X., Chu, W., Li, J., Gao, Y., An, K., Song, W., Xin, M.,Yao, Y., Peng, H., Ni, Z., Sun, Q., & Hu, Z. (2021). Histone acetyltransferase TaHAG1 acts as a crucial regulator to strengthen salt tolerance of hexaploid wheat. Plant Physiology, 186(4), 1951-1969. doi: 10.1093/plphys/kiab187.##Zong, Y., Wang, Y., Li, C., Zhang, R., Chen, K., Ran, Y., Qiu,J. L., Wang, D., & Gao, C. (2017). Precise base editing in rice, wheat and maize with a Cas9-cytidine deaminase fusion. Nature Biotechnology, 35(5), 438-440. doi: 10.1038/nbt.3811.##