18O'ZBEKISTONDA BUG'DOY TADQIQOTLARINING KELGUSI ISTIQBOLLARI
O. S. Turayev
O'zbekiston Respublikasi Prezidenti huzuridagi Orolbo'yi xalqaro innovatsiya markazi
A. A. Dolimov
Genetika va o'simliklar eksperimental biologiyasi instituti
B. T. Tursunmurodova
Genetika va o'simliklar eksperimental biologiyasi instituti
S. S. Baboyeva
Genetika va o'simliklar eksperimental biologiyasi instituti
P. A. Tureshov
O'zbekiston Respublikasi Prezidenti huzuridagi Orolbo'yi xalqaro innovatsiya markazi
A. I. Seytmusayev
Qoraqalpog'iston qishloq xo'jaligi va agrotexnologiyalar instituti
Z.M. Ziyayev
Genetika va o'simliklar eksperimental biologiyasi instituti
F. N. Kushanov
O'zbekiston Milliy universiteti, Biologiya fakulteti
ANNOTATSIYA
Bug'doy ekinini yetishtirish jarayonida o'simliklar biotik va abiotik stress omillar ta'sirida ko'plab to'siqlarga duch keladi, natijada hosil kamayadi. Bu kabi muammolarning yechimi genetika va genomikaning ilg'or texnologiyalarini qo'llagan xoldi chidamli navlarni yaratish hisoblanadi. Ushbu tahliliy maqolada, biz, O'zbeksiton bug'doy tadqiqotlarida qo'llanilishi boshlangan va yaqin kelajakda boshlanadigan tadqiqotlar, genom texnologiyalari borasida ma'lumotlar keltirdik.
Kalit so'zlar. Bug'doy, Triticum aestivum, MAS, GWAS, CRISPR/Cas9, poliploidiya, chidamlilik, DNK markerlari, genlar.
ABSTRACT
During wheat cultivation, plants face many challenges under the influence of biotic and abiotic stress factors, resulting in yield decreases. One of the solutions to such problems is the creation of resistant varieties using advanced technologies in genetics and genomics. In this review, we
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reported on the genomic technologies being used and will be used in the recent future in Uzbekistan wheat research.
Keywords. Wheat, Triticum aestivum, MAS, GWAS, CRISPR/Cas9, polyploidy, resistance, DNA markers, genes.
Dunyo aholisining yil sayin ortib borishi sababli don mahsulotlariga bo'lgan talab ham ortib bormoqda. 2050 yilga kelib, dunyo aholisi 9,8 milliardga qadar o'sib borishi kutilmoqda [40], bu esa oziq-ovqat mahsulotini ikki barobarga oshirishni talab etadi. Xususan, so'nggi 10 yil ichida dunyo aholisining o'sish sur'ati 1,1% atrofida bo'lgan bo'lsa, bu davrda bug'doy donini ishlab chiqarish 1,8% ni tashkil etgan. Agar aholining o'sish sur'atlarini bug'doy yetishtiriladigan maydonlarni kengaytirish imkoniyatlari bilan taqqoslansa, keyingi avlodni don bilan ta'minlashda jiddiy muammolar paydo bo'lishi mumkin [24]. Bu muammoning asosiy yechimlaridan biri, salbiy ta'sir etuvchi, biotik va abiotik omillarga bardoshli, hosildor va yuqori sifatli bug'doy navlarni yetishtirishdan iborat. Birinchi navbatda ushbu talablarga to'liq javob beradigan birlamchi manbalarni (donorlarni) aniqlab, zamonaviy usul va texnologiyalar bilan foydalanilgan holda [17], jumladan, Markerlarga asoslangan seleksiya (MAS), genlar espressiyasi tahlili, butun-genom assotsiatsiyalar tahlili (GWAS - ingl. Genome Wide Association Study), poliploidiya va CRISPR/Cas9 kabi ilg'or texnologiyalarni qo'llash selektsiya jarayonlarini qisqartirish, aniqligini va samaradorligini oshirish imkonini beradi.
ADABIYOTLAR TAHLILI VA METODOLOGIYA
Markerlarga asoslangan selektsiya va genlarni piramidalash Suv tanqisligi, issiqlik va sho'rlanish kabi abiotik stresslar hamda zararkunanda hasharotlar va kasalliklar biotik stress kabi omillar, bug'doydan sifatli va yuqori hosil olishga to'sqinlik qiluvchi eng asosiy omillardir [39,41,42]. Ular genlarning o'zaro munosabati bog'lanishiga salbiy ta'sir ko'rsatib, o'simliklarda murakkab o'zgarishlarni keltirib chiqaradi [1,3,8,19,21]. Tadqiqotchilar, bug'doy hosili va sifatini yaxshilash uchun ushbu stress omillariga chidamli navlar va genotiplarni aniqlash, tavsiflash va selektsiya qilishga e'tibor qaratishmoqda. Ushbu seleksiya morfologik, fiziologik va molekulyar markerlarga asoslangan holda amalga oshiriladi. Morfologik markerlar ko'zga ko'rinadigan belgilardan ildiz yoki gul morfologiyasi, erta pishish va donlarning biomassasi hisobga olinadigan ba'zi muhim parametrlardir. Fiziologik va biokimyoviy markerlar xlorofill tarkibini, prolin, saxaroza to'planishini, stressdan himoya qiluvchi
KIRISH
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moddalarni, membrananing barqarorligini va gormonlar tarkibini aniqlaydi. Molekulyar markerlar abiotik va biotik stresslarga chidamlilik mexanizmlari, transkriptsiya omillari va metabolitik yo'l bilan bog'liq genlarni o'z ichiga oladi [34].
Bugungi kunda tadqiqot guruhimiz tomonidan, O'zbekiston Respublikasi Prezidenti huzuridagi Orolbo'yi xalqaro innovatsiya markazida markerlarga asoslangan selektsiya texnologiyasi asosida bahorgi bug'doyning sho'rga, issiqqa va qurg'oqchilikka chidamli, yuqori oqsil tarkibli yangi navlarini olish bo'yicha tadqiqotlar olib borilmoqda. Shuningdek, Genetika va o'simliklar eksperimental biologiyasi institutida X.To'raqulov va D.Erjigitovlar tomonidan, genlarni piramidalash texnologiyasi asosida kuzgi bug'doyning zang kasalligiga chidamli, yuqori harorat, sovuqqa va qurg'oqchilikka bardoshli navlarini yaratish ustida tadqiqotlar olib borishmoqda [22]. Bundan tashqari, U.Baxodirov va S.Baboev kabi olimlar tomonidan, markerlarga asoslangan selektsiya texnologiyasi asosida kuzgi bug'doyning shiraga chidamli yangi navlarini yaratish borasida izlanishlar olib borishmoqda [4].
Markerlarga asoslangan selektsiya texnologiyasi o'zining vaqt va boshqa qimmatli resurslarni tejab qolishi, tezkor va aniq tanlovni amalga oshirishi bilan O'zbekiston bug'doy selektsiyasini yangi bosqichga olib chiqadi. Kelgusida bug'doy selektsiyasida MAS texnologiyasining keng ko'lamda ko'llanilishi kasallik va zararkunandaga chidamli, atrof muhitning noqulay sharoitlariga moslashgan yangi navlarni tez va samarali selektsiya qilish imkonini beradi.
Degidrin kodlovchi genlarning tadqiqi
Degidrin oqsillari oilasi donli ekinlarda qurg'oqchilik va sovuqqa chidamlilikda muhim rol o'ynovchi himoya oqsillari hisoblanadi [9,10,12,25,28]. Degidrinlar bo'yicha ko'p yillik tadqiqotlar amalga oshirilganligiga qaramay, bug'doy degidrinlari qurg'oqchilikka chidamlilikni oshirishi borasida ishonchli dalillar yetarlicha emas. Ilmiy nashrlar degidrinlarni kodlovchi genlarning qurg'oqchilik stressiga javob mexanizmlaridagi ahamiyati, faoliyati va ta'siri haqida to'liq ma'lumot bermaydi [28]. Kelgusida, qurg'oqchilikka chidamlilikda yumshoq bug'doy degidrinlarini kodlovchi individual genlarning rolini o'rganish maqsad qilingan. Bunday tadqiqotlar bug'doyning suv tanqisligiga chidamliligi uchun markerlarga asoslangan selektsiya texnologiyasini rivojlantirishga imkon beradi.
Hozirda, Genetika va o'simliklar eksperimental biologiyasi institutida Innovatsion rivojlanish agentligi tomonidan moliyalashtirilgan "Qurg'oqchilikka chidamli istiqbolli navlarni yaratish uchun samarali molekulyar markerlarni ishlab chiqish maqsadida degidrin-kodlovchi genlarning yumshoq bug'doyning qurg'oqchilikka
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chidamlilikdagi hissasini aniqlash" mavzusida "O'zbekiston-Belarus" xalqaro fundamental loyihasi amalga oshirilib kelmoqda. Ushbu tadqiqot natijasida, genetik jihatidan kelib chiqishi turlicha bo'lgan yumshoq bug'doy navlarining qurg'oqchilikka chidamliligida eng ko'p hissa qo'shadigan degidrin-kodlovchi genlar va ularning nukleotid ketma-ketliklari aniqlanadi. Shu bilan birga, qurg'oqchilik sharoitida mazkur genlarning ekspressiya darajasi o'rganiladi. Yumshoq bug'doy genotiplarining suv tanqisligiga chidamlilik darajasini baholash hamda bug'doy selektsiyasi uchun molekulyar usullar ishlab chiqiladi. Qurg'oqchilikka chidamli bug'doy genotiplari hamda ushbu belgi bilan assotsiatsiyalangan genetik markerlar aniqlanadi va selektsiya tadqiqotlariga yo'naltiriladi.
Poliploidiya usulida bug'doy genomini oshirish
Seleksiyada turli genomli o'simliklarni o'zaro duragaylash va ulardan avlod olish juda murakkab vazifa hisoblanadi [18,23,26,]. Shu sababli, qishloq xo'jaligi ekinlarining yovvoyi ajdodlaridagi qimmatli xo'jalik belgilarini mahalliy navlarga introgressiya qilishda qiyinchiliklarga duch kelinadi. Xususan, Aegilops tauschii diploid (D genomli) turi hozirda butun dunyoda eng ko'p yetishtiriladigan geksaploid (AABBDD genomli) yumshoq bug'doy Triticum aestivum L. turining yovvoyi ajdodi hisoblanadi [6,13]. Geksaploizidatsiya jarayoni tahminan 8 ming yil oldin sodir bo'lgan. Bunda, diploid Aegilops tauschii turi tetraploid Triticum turgidum turi bilan tasodifiy duragaylanib qolgan [13,30].
Diploid, tetraploid va geksaploid genomli organizmlarning o'zaro duragaylanishidagi murakkabliklarni hisobga olgan xolda, poliploidizatsiya usulini qo'llash eng maqbul yo'l hisoblanadi. Hozirda, Genetika va O'EB instituti olimlari tomonidan bug'doyning Aegilops tauschii turi genomini kolxitsin moddasi ta'siri asosida tetraploidga oshirish ustida izlanishlar olib borilmoqda. Kelgusida, olingan poliploidlarni boshqa tetraploid turlar bilan duragaylash, olingan duragaylar genomini gekcaploidga oshirish rejalashtirilgan. Bu esa diploid va tetraploid genomli bug'doy turlaridagi qimmatli genlar majmuini mahalliy yumshoq bug'doy navlariga introgressiya qilish imkonini beradi.
Bug'doy tadqiqotlarida GWAS va CRISPR/Cas9 texnologiyalarini qo'llash
Butun-genom assotsiatsiyalar tahlili (GWAS). GWAS ko'plab qishloq xo'jaligi ekinlarida agronomik muhim belgilar bilan bog'liq bo'lgan miqdoriy belgilar lokuslarini (QTL) aniqlash uchun keng qo'llaniladi [11,17,32,35] . Yuqori samarali yagona nukleotid polimorfizmi (SNP)ga asoslangan mikrochip texnologiyasi genotiplash platformalarini ishlab chiqish o'simliklarda GWAS tahlilini amalga oshirish imkonini berdi va
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murakkab belgilar uchun QTL va allel o'zgarishini aniqlash uchun samarali yondashuvga aylandi. GWAS bug'doyda abiotik stressga chidamlilik [27,29], hosil komponentlari, don sifati yoki kasalliklarga chidamlilik [5,31] bilan bog'liq QTL hududlarini aniqlash uchun muvaffaqiyatli qo'llanildi. Biroq, hozirgi kunga qadar, O'zbeksitonda GWAS texnologiyasini qo'llash bo'yicha genetik tadqiqotlari amalg a oshirilmagan.
Genlarni tahrirlashning CRISPR/Cas9 texnologiyasi. Genomni tahrirlash molekulyar biologiya tadqiqotlari uchun, ayniqsa murakkab genomlarga ega bo'lgan o'simliklar uchun uzoq muddatli muammo bo'lib kelgan [33]. So'ngi yigirma yil davomida kashf etilgan CRISPR/Cas9 (ingl. Clustered Regularly Interspaced Short Palindromic Repeats/CRISPR-associated protein 9) tizimi kichik RNKlarni boshqarish asosida bir nechta genlarni tahrirlash imkonini beruvchi genomni tahrirlash uchun ko'p qirrali vositadir [14]. CRISPR/Cas9 tizimining samaradorligi diploid o'simliklarda olib borilgan bir nechta tadqiqotlarda ko'rsatilgan bo'lsada, uni qo'llash poliploid va murakkab genomli o'simliklar uchun qiyinligicha qolmoqda [16]. Shunga qaramasdan, mukrakkab genomli geksaploid bug'doyda CRISPR/Cas9 texnologiyasini muvaffaqiyatli qo'llash borasida tadqiqot natijalari mavjud [2,7,15,20,36,37,38]. Nufuzli ilmiy jurnallarda O'zbeksitonda CRISPR/Cas9 texnologiyasining muvaffaqiyatli qo'llanilishi bo'yicha birorta ham ilmiy tadqiqot natijalari keltirilmagan. Ushbu texnologiyaning O'zbekiston bug'doy tadqiqotlarida qo'llanilishi bug'doy seleksiyasidagi mavjud muammolarni gen muhandisligi yo'li bilan tez va samarali hal etish imkonini beradi.
XULOSALAR
O'zbekistonda bug'doyning genetik va genom tadqiqotlarida GWAS, MAS, poliploidiya va CRISPR/Cas9 texnologiyalarini qo'llash, qimmatli xo'jalik belgilarga aloqador genlarni yuqori aniqlikda identifikatsiya qilish, ikkita va undan ortiq genlarni bir genotipga jamlash, genetik to'siqlarni bartaraf etish hamda genlarni maqsadli boshqarish imkonini beradi.
REFERENCES
1. Bhatta, M., Morgounov, A., Belamkar, V., Wegulo, S.N., Dababat, A.A., Erginbas-Orakci, G., Bouhssini, M.E., Gautam, P., Poland, J., Akci, N. and Demir, L., 2019. Genome-wide association study for multiple biotic stress resistance in synthetic hexaploid wheat. International Journal of Molecular Sciences, 20(15), p.3667.
2. Bhowmik, P., Ellison, E., Polley, B., Bollina, V., Kulkarni, M., Ghanbarnia, K., Song, H., Gao, C., Voytas, D.F. and Kagale, S., 2018. Targeted mutagenesis in wheat microspores using CRISPR/Cas9. Scientific reports, 8(1), pp.1-10.
April, 2023
263
3. Blum, A., 2014. The abiotic stress response and adaptation of triticale—A review. Cereal research communications, 42, pp.359-375.
4. Bohodirov U.Sh., Adilova Sh.Sh., Turaev O.S., Erjigitov D.Sh., Kushanov F.N., Baboev S.K. Determination of aphid resistance genes in local bread wheat (Triticum aestivum L.) varieties of Uzbekistan using DNA markers. international scientific and practical conference "The time of scientific progress" 2022 Warsaw, (Poland). https://doi.org/10.5281/zenodo.7198391
5. Choudhury, S., Larkin, P., Xu, R., Hayden, M., Forrest, K., Meinke, H., Hu, H., Zhou, M. and Fan, Y., 2019. Genome wide association study reveals novel QTL for barley yellow dwarf virus resistance in wheat. BMC genomics, 20(1), pp.1-8.
6. Cox, T.S., Wu, J., Wang, S., Cai, J., Zhong, Q. and Fu, B., 2017. Comparing two approaches for introgression of germplasm from Aegilops tauschii into common wheat. The Crop Journal, 5(5), pp.355-362.
7. Gil-Humanes, J., Wang, Y., Liang, Z., Shan, Q., Ozuna, C.V., Sánchez-León, S., Baltes, N.J., Starker, C., Barro, F., Gao, C. and Voytas, D.F., 2017. High-efficiency gene targeting in hexaploid wheat using DNA replicons and CRISPR/Cas9. The Plant Journal, 89(6), pp.1251-1262.
8. Golebiowska-Paluch, G. and Dyda, M., 2023. The Genome Regions Associated with Abiotic and Biotic Stress Tolerance, as Well as Other Important Breeding Traits in Triticale. Plants, 12(3), p.619.
9. Graether, S.P. and Boddington, K.F., 2014. Disorder and function: a review of the dehydrin protein family. Frontiers in plant science, 5, p.576.
10. Hara, M., Terashima, S. and Kuboi, T., 2001. Characterization and cryoprotective activity of cold-responsive dehydrin from Citrus unshiu. Journal of Plant Physiology, 158(10), pp.1333-1339.
11. He, X., Fuller, C.K., Song, Y., Meng, Q., Zhang, B., Yang, X. and Li, H., 2013. Sherlock: detecting gene-disease associations by matching patterns of expression QTL and GWAS. The American Journal of Human Genetics, 92(5), pp.667-680.
12. Hughes, S. and Graether, S.P., 2011. Cryoprotective mechanism of a small intrinsically disordered dehydrin protein. Protein Science, 20(1), pp.42-50.
13. Jia, J., Zhao, S., Kong, X. et al. Aegilops tauschii draft genome sequence reveals a gene repertoire for wheat adaptation. Nature 496, 91-95 (2013). https://doi.org/10.1038/nature12028
14. Jiang, F. and Doudna, J.A., 2017. CRISPR-Cas9 structures and mechanisms. Annual review of biophysics, 46, pp.505-529.
15. Jouanin, A., Gilissen, L.J., Schaart, J.G., Leigh, F.J., Cockram, J., Wallington, E.J., Boyd, L.A., van den Broeck, H.C., van der Meer, I.M., America, A.H.P. and Visser, R.G.F., 2020. CRISPR/Cas9 gene editing of gluten in wheat to reduce gluten content and exposure—Reviewing methods to screen for coeliac safety. Frontiers in nutrition, 7, p.51.
16. Kim, D., Alptekin, B. and Budak, H., 2018. CRISPR/Cas9 genome editing in wheat. Functional & integrative genomics, 18, pp.31-41.
17. Kushanov FN, Turaev OS, Ernazarova DK, Gapparov BM, Oripova BB, Kudratova MK, Rafieva FU, Khalikov KK, Erjigitov
April, 2023
264
ISSN: 2181-1385 ISI: 0,967 | Cite-Factor: 0,89 | SIS: 1,9 | ASI: 1,3 | SJIF: 5,771 | UIF: 6,1
DS, Khidirov MT, Kholova MD, Khusenov NN, Amanboyeva RS, Saha S, Yu JZ and Abdurakhmonov IY (2021) Genetic Diversity, QTL Mapping, and Marker-Assisted Selection Technology in Cotton (Gossypium spp.). Front. Plant Sci. 12:779386. doi: 10.3389/fpls.2021.779386
18. Lee, T.H., Tang, H., Wang, X. and Paterson, A.H., 2012. PGDD: a database of gene and genome duplication in plants. Nucleic acids research, 41(D1), pp.D1152-D1158.
19. Li, H., Tong, Y., Li, B., Jing, R., Lu, C. and Li, Z., 2010. Genetic analysis of tolerance to photo-oxidative stress induced by high light in winter wheat (Triticum aestivum L.). Journal of Genetics and Genomics, 37(6), pp.399-412.
20. Li, J., Li, Y. and Ma, L., 2021. Recent advances in CRISPR/Cas9 and applications for wheat functional genomics and breeding. Abiotech, pp.1-11.
21. Maulana, F., Huang, W., Anderson, J.D. and Ma, X.F., 2020. Genome-wide association mapping of seedling drought tolerance in winter wheat. Frontiers in Plant Science, 11, p.573786.
22. Mullaev D., Turakulov Kh., Chinikulov B., Erjigitov D., Isokulov S. (2020). Study of virulent characteristics of wheat yellow rust (Puccina Striiformis f.sp. tritica) in Uzbekistan. The American Journal of Applied Sciences. 02. 51-59. 10.37547/tajas/Volume02Issue08-07.
23. Panchy N, Lehti-Shiu M, Shiu SH. Evolution of Gene Duplication in Plants. Plant Physiol. 2016 Aug;171(4):2294-316. doi: 10.1104/pp.16.00523. Epub 2016 Jun 10. PMID: 27288366; PMCID: PMC4972278.
24. Pardey, P.G. "A Strategic Look at Global Wheat Production, Productivity and R&D Developments." Czech Journal of Genetics and Plant Breeding, vol. 47, no. Special Issue, 2011, https://doi.org/10.17221/3248-cjgpb
25. Puhakainen, T., Hess, M.W., Makela, P., Svensson, J., Heino, P. and Palva, E.T., 2004. Overexpression of multiple dehydrin genes enhances tolerance to freezing stress in Arabidopsis. Plant molecular biology, 54, pp.743-753.
26. Qiao, X., Li, Q., Yin, H., Qi, K., Li, L., Wang, R., Zhang, S. and Paterson, A.H., 2019. Gene duplication and evolution in recurring polyploidization-diploidization cycles in plants. Genome biology, 20(1), pp.1-23.
27. Sallam, A., Eltaher, S., Alqudah, A.M., Belamkar, V. and Baenziger, P.S., 2022. Combined GWAS and QTL mapping revealed candidate genes and SNP network controlling recovery and tolerance traits associated with drought tolerance in seedling winter wheat. Genomics, 114(3), p.110358.
28. Smith, M.A.; Graether, S.P. The Disordered Dehydrin and Its Role in Plant Protection: A Biochemical Perspective. Biomolecules 2022, 12, 294. https://doi.org/10.3390/biom12020294
29. Suliman, S., Alemu, A., Abdelmula, A.A., Badawi, G.H., Al-Abdallat, A. and Tadesse, W., 2021. Genome-wide association analysis uncovers stable QTLs for yield and quality traits of spring bread wheat (Triticum aestivum) across contrasting environments. Plant gene, 25, p.100269.
30. Tadesse, W., Hsam, S.L., Wenzel, G. and Zeller, F.J., 2006.
Identification and monosomic analysis of tan spot resistance genes
April, 2023
ISSN: 2181-1385 ISI: 0,967 | Cite-Factor: 0,89 | SIS: 1,9 | ASI: 1,3 | SJIF: 5,771 | UIF: 6,1
in synthetic wheat lines (Triticum turgidum L.x Aegilops tauschii Coss.). Crop science, 46(3), pp.1212-1217.
31. Tomar, V., Singh, D., Dhillon, G.S., Singh, R.P., Poland, J., Joshi, A.K., Singh, P.K., Bhati, P.K., Kumar, S., Rahman, M. and Tiwari, B.S., 2021. New QTLs for spot blotch disease resistance in wheat (Triticum aestivum L.) using genome-wide association mapping. Frontiers in Genetics, 11, p.613217.
32. Tsai, H.Y., Janss, L.L., Andersen, J.R., Orabi, J., Jensen, J.D., Jahoor, A. and Jensen, J., 2020. Genomic prediction and GWAS of yield, quality and disease-related traits in spring barley and winter wheat. Scientific reports, 10(1), p.3347.
33. Vats, S.; Kumawat, S.; Kumar, V.; Patil, G.B.; Joshi, T.; Sonah, H.; Sharma, T.R.; Deshmukh, R. Genome Editing in Plants: Exploration of Technological Advancements and Challenges. Cells 2019, 8, 1386. https://doi.org/10.3390/cells8111386
34. Younis A, Ramzan F, Ramzan Y, Zulfiqar F, Ahsan M, Lim KB. Molecular Markers Improve Abiotic Stress Tolerance in Crops: A Review. Plants (Basel). 2020 Oct 15;9(10):1374. doi: 10.3390/plants9101374. PMID: 33076554; PMCID: PMC7602808.
35. Zargar, S.M., Raatz, B., Sonah, H., Bhat, J.A., Dar, Z.A., Agrawal, G.K. and Rakwal, R., 2015. Recent advances in molecular marker techniques: insight into QTL mapping, GWAS and genomic selection in plants. Journal of crop science and biotechnology, 18, pp.293-308.
36. Zhang, J., Zhang, H., Li, S., Li, J., Yan, L. and Xia, L., 2021. Increasing yield potential through manipulating of an ARE1 ortholog related to nitrogen use efficiency in wheat by CRISPR/Cas9. Journal of integrative plant biology, 63(9), pp.1649-1663.
37. Zhang, S., Zhang, R., Gao, J., Song, G., Li, J., Li, W., Qi, Y., Li, Y. and Li, G., 2021. CRISPR/Cas9-mediated genome editing for wheat grain quality improvement. Plant Biotechnology Journal, 19(9), p.1684.
38. Zhang, Z., Hua, L., Gupta, A., Tricoli, D., Edwards, K.J., Yang, B. and Li, W., 2019. Development of an Agrobacterium-delivered CRISPR/Cas9 system for wheat genome editing. Plant biotechnology journal, 17(8), pp.1623-1635.
39. Zhao, W., Liu, L., Shen, Q., Yang, J., Han, X., Tian, F. and Wu, J., 2020. Effects of water stress on photosynthesis, yield, and water use efficiency in winter wheat. Water, 12(8), p.2127.
40. БМТ. Дуне ах,олисининг истщболлари: асосий топилмалар ва жадваллар. Ишчи № ESA/P/WP/248. Бирлашган миллатлар ташкилоти, Нью-Йорк, Нью-Йорк
41. Лиховидова, В.А., Казакова, А.С. and Самофалова, Н.Е., 2019. Влияние водного и температурного стрессов на всхожесть семян сортов твердой озимой пшеницы, полученных в контрастные по погодным условиям годы. Зерновое хозяйство России, (5), pp.34-39.
42. Манукян, И.Р., 2021. Физиологические аспекты стрессоустойчивости озимой пшеницы. Нива Поволжья, (1 (58)), pp.30-36.
(2017)
April, 2023
