Kartoshka navlaridagi chidamlilik genlarining viruslarga qarshi himoya mexanizmi

Ahmadjonova Sevinchbonu Ulugbek kizi; Jovliyeva D.T.; Fayziyev Vokhid Bakhramovich

doi:10.66073/refocus-conf/3010033

Kartoshka navlaridagi chidamlilik genlarining viruslarga qarshi himoya mexanizmi

Ahmadjonova Sevinchbonu Ulugbek kizi, Jovliyeva D.T., Fayziyev Vokhid Bakhramovich iDChirchiq davlat pedagogika universiteti2026

KonferensiyalarFundamental va amaliy mikrobiologiyaning bugungi holati va rivojlanish istiqbollari: yosh olimlar nigohiJild 1Son 8Sahifa 123–126

Annotatsiya

Maqolada kartoshka (Solanum tuberosum L.) navlarida uchraydigan virusga chidamlilik genlarining ishlash mexanizmlari adabiyotlar asosida tahlil qilingan. Gipersezgirlik reaksiyasi va ekstremal chidamlilikni ta‘minlovchi genlar (Rx1, Rx2, Ny, Ry), ularning virus oqsillarini tanib olishi hamda RNK-interferensiya tizimi bilan bog‘liqligi yoritilgan. Chidamlilik genlarini molekulyar markerlar yordamida aniqlashning seleksiyadagi ahamiyati ko‘rsatilgan.

Boshqa tillarda

ruschaМеханизм противовирусной защиты, обеспечиваемой генами устойчивости у сортов картофеля

В статье на основе литературных данных проанализированы механизмы действия генов вирусоустойчивости у картофеля (Solanum tuberosum L.). Рассмотрены гены (Rx1, Rx2, Ny, Ry), обеспечивающие реакцию сверхчувствительности и экстремальную устойчивость, их распознавание вирусных белков и связь с системой РНК-интерференции. Показано значение молекулярных маркеров в селекции на устойчивость.

inglizchaAntiviral defense mechanism mediated by resistance genes in potato varieties

The article analyzes, based on the literature, the mechanisms of virus resistance genes in potato (Solanum tuberosum L.). Genes (Rx1, Rx2, Ny, Ry) that provide the hypersensitive response and extreme resistance are reviewed, together with their recognition of viral proteins and their link to the RNA interference system. The importance of molecular markers in resistance breeding is shown.

Kalit so‘zlar

kartoshka
virusga chidamlilik
Rx1
NLR oqsillari
ekstremal chidamlilik
gipersezgirlik
RNK-interferensiya
KXV
KYV

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FUNDAMENTAL VA AMALIY MIKROBIOLOGIYANING BUGUNGI HOLATI VA RIVOJLANISH ISTIQBOLLARI: YOSH OLIMLAR NIGOHI Respublika ilmiy-amaliy anjumani · Toshkent, 2026-yil 30-oktabr

II BIOTEXNOLOGIYA, MOLEKULYAR BIOLOGIYA VA BIOINFORMATIKA DOI 10.66073/ReFocus-Conf/3010033

KARTOSHKA NAVLARIDAGI CHIDAMLILIK GENLARINING VIRUSLARGA QARSHI HIMOYA MEXANIZMI Ahmadjonova Sevinchbonu Ulug‘bek qizi, 2Jovliyeva D.T., 3Fayziyev Voxid Baxramovich Chirchiq davlat pedagogika universiteti talabasi, Toshkent viloyati, O‘zbekiston Chirchiq davlat pedagogika universiteti Biologiya kafedrasi dotsenti Chirchiq davlat pedagogika universiteti Biologiya kafedrasi professori e-mail: sevinchahmadjonova20@gmail.com Annotatsiya: Maqolada kartoshka (Solanum tuberosum L.) navlarida uchraydigan virusga chidamlilik genlarining ishlash mexanizmlari adabiyotlar asosida tahlil qilingan. Gipersezgirlik reaksiyasi va ekstremal chidamlilikni ta’minlovchi genlar (Rx1, Rx2, Ny, Ry), ularning virus oqsillarini tanib olishi hamda RNK-interferensiya tizimi bilan bog‘liqligi yoritilgan. Chidamlilik genlarini molekulyar markerlar yordamida aniqlashning seleksiyadagi ahamiyati ko‘rsatilgan. Kalit so‘zlar: kartoshka, virusga chidamlilik, Rx1, NLR oqsillari, ekstremal chidamlilik, gipersezgirlik, RNK-interferensiya, KXV, KYV. Аннотация: В статье на основе литературных данных проанализированы механизмы действия генов вирусоустойчивости у картофеля (Solanum tuberosum L.). Рассмотрены гены (Rx1, Rx2, Ny, Ry), обеспечивающие реакцию сверхчувствительности и экстремальную устойчивость, их распознавание вирусных белков и связь с системой РНК-интерференции. Показано значение молекулярных маркеров в селекции на устойчивость. Ключевые слова: картофель, вирусоустойчивость, Rx1, NLR-белки, экстремальная устойчивость, сверхчувствительность, РНК-интерференция, ВКХ, ВКY. Abstract: The article analyzes, based on the literature, the mechanisms of virus resistance genes in potato (Solanum tuberosum L.). Genes (Rx1, Rx2, Ny, Ry) that provide the hypersensitive response and extreme resistance are reviewed, together with their recognition of viral proteins and their link to the RNA interference system. The importance of molecular markers in resistance breeding is shown. Keywords: potato, virus resistance, Rx1, NLR proteins, extreme resistance, hypersensitive response, RNA interference, PVX, PVY. Kirish Kartoshka (Solanum tuberosum L.) dunyoning eng muhim oziq-ovqat ekinlaridan biridir. FAO ma’lumotlariga ko‘ra, rivojlanayotgan mamlakatlarning ko‘pchiligida u guruch, bug‘doy va makkajo‘xoridan keyin to‘rtinchi o‘rinda turadi, dunyo bo‘yicha ishlab chiqarish esa 2007-yilda 325 million tonnaga yetgan (FAO, 2009). Shu sababli kartoshka hosildorligini saqlash oziq-ovqat xavfsizligi uchun muhim vazifadir. Kartoshka asosan vegetativ yo‘l bilan, ya’ni tuganaklar orqali ko‘paytiriladi. Bunday ekinlarda virus tuganak bilan keyingi yilga o‘tadi va nazorat qilinmasa, hosil miqdori va sifatini sezilarli kamaytiradi (Loebenstein va boshq., 2001; Valkonen, 2007). Toza urug‘lik material olish uchun to‘qima kulturasi usullaridan foydalanilsa-da, ko‘p mamlakatlarda virus bilan zararlanish hanuzgacha katta yo‘qotishlarga sabab bo‘lmoqda (Loebenstein va boshq., 2001).

IQTIBOS UCHUN Ahmadjonova S.U., Jovliyeva D.T., Fayziyev V.B. Kartoshka navlaridagi chidamlilik genlarining viruslarga qarshi himoya mexanizmi // Fundamental va amaliy mikrobiologiyaning bugungi holati va rivojlanish istiqbollari: yosh olimlar nigohi: Respublika ilmiy-amaliy anjumani materiallari. – Toshkent, 2026. – B. 123–126. DOI: https://doi.org/10.66073/ReFocus-Conf/3010033

Kartoshkada eng ko‘p o‘rganilgan viruslarga kartoshka X virusi (KXV, Potato virus X) va kartoshka Y virusi (KYV, Potato virus Y) kiradi. O‘simlik virusologlari o‘rtasida o‘tkazilgan so‘rovda ilmiy va iqtisodiy ahamiyati bo‘yicha eng muhim 10 ta o‘simlik virusi aniqlanganda, KYV beshinchi, KXV esa o‘ninchi o‘rinni egalladi (Scholthof va boshq., 2011). Virus o‘simlik hujayrasi ichida ko‘payadi, shuning uchun uni kimyoviy vositalar bilan yo‘qotish qiyin. Eng ishonchli yo‘l chidamli navlarni yaratishdir. Chidamlilik genlarining ko‘pchiligi virus oqsilini maxsus taniydigan NLR oqsillarini kodlaydi (Jones va Dangl, 2006). Genlarning ketma-ketligi ma’lum bo‘lgani sari ularni molekulyar markerlar yordamida aniqlash imkoniyati ham kengaymoqda (Ahmadvand va boshq., 2013). Shuning uchun chidamlilik genlari qanday ishlashini bilish seleksiyada to‘g‘ri gen tanlash va yangi navlarni tez baholash uchun zarur. Maqolaning maqsadi kartoshkadagi asosiy chidamlilik genlarining viruslarga qarshi ishlash mexanizmini adabiyotlar asosida tahlil qilish va ularni aniqlash usullari bilan bog‘lashdir. Metodlar Ish adabiyotlar tahlili usulida bajarildi. PubMed Central, Google Scholar va NCBI ma’lumotlar bazalaridagi ilmiy maqolalar o‘rganildi. Chidamlilik genlari qaysi turdan olingani, qaysi virus oqsilini tanishi va qanday javob reaksiyasi berishi bo‘yicha solishtirildi. Natijalar 1-jadvalda jamlandi. Natijalar 1. Chidamlilik genlarining turlari Jones va Dangl (2006) o‘simlik immunitetini ikki bosqichga bo‘ladi: birinchisida o‘simlik patogenning umumiy belgilarini tanisa, ikkinchisida patogenning aynan o‘ziga xos oqsilini taniydi. Kartoshkadagi virusga qarshi genlar ko‘pincha ikkinchi bosqichga kiradi. Ular NLR oqsillarini (NB-LRR) kodlaydi. Bu oqsil virusning ma’lum bir oqsilini taniganda himoya javobini ishga tushiradi. Asosiy genlar 1-jadvalda berilgan. Javob ikki xil bo‘lishi mumkin. Birinchisi gipersezgirlik reaksiyasi (GR): virus kirgan hujayralar nobud bo‘ladi va virus bargdagi kichik nekrotik dog‘ ichida qolib ketadi. Ikkinchisi ekstremal chidamlilik (EC): virus juda erta to‘xtatiladi va o‘simlikda deyarli hech qanday belgi ko‘rinmaydi. Rx genida ikkala javob ham mavjud. Bendahmane va boshq. (1999) ko‘rsatishicha, Rx hujayra nobud bo‘lishini ham boshlay oladi, lekin ekstremal chidamlilik nekrozdan alohida ishlaydi va undan ustun turadi. 1-jadval. Kartoshkadagi asosiy virusga chidamlilik genlari Gen Manba Virus Reaksiya turi Virus omili Rx1 S. tuberosum ssp. KXV Ekstremal Qobiq oqsili (CP) andigena chidamlilik Rx2 S. acaule KXV Ekstremal Qobiq oqsili (CP) chidamlilik Nb Madaniy kartoshka KXV (ayrim Gipersezgirlik 25K harakat oqsili navlari shtammlari) Ny-1, Ny-2, Madaniy kartoshka KYV (ayrim Gipersezgirlik To‘liq Ny-tbr navlari shtammlari) aniqlanmagan Ry-adg, Ry- S. tuberosum ssp. KYV (ko‘p Ekstremal To‘liq sto, Ry-chc andigena, S. shtammlari) chidamlilik aniqlanmagan stoloniferum, S. chacoense

2. Rx1 va Rx2 genlari: KXVga qarshi himoya Rx1 geni S. tuberosum ssp. andigena shaklidan, Rx2 geni esa yovvoyi S. acaule turidan olingan. Rx1 XII xromosomada, Rx2 esa V xromosomada joylashgan. Ikkala gen ham CC-NB-LRR oqsilini kodlaydi va KXVning qobiq oqsilini taniydi (Bendahmane va boshq., 1995, 1999, 2000). Bu ikki gen bir-biriga juda o‘xshash bo‘lgani uchun ularni ajratib aniqlash uchun alohida molekulyar markerlar ishlab chiqilgan (Ahmadvand va boshq., 2013). Rx oqsilining qismlari bir-biri bilan o‘zaro ta’sirlashadi: CC qismi NB-LRR qismi bilan, LRR qismi esa CC-NB qismi bilan bog‘lanadi. Bu qismlarni alohida ifodalab ham ishlaydigan oqsil hosil qilish mumkin (Moffett va boshq., 2002). Tameling va Baulcombe (2007) Rx oqsili hujayrada RanGAP2 oqsili bilan bog‘lanishini aniqladi. RanGAP2 kamaytirilganda Rx orqali chidamlilik susayadi, ya’ni bu oqsil KXVga qarshi himoya uchun kerak. 3. Nb, Ny va Ry genlari Nb geni ham KXVga qarshi ishlaydi, lekin gipersezgirlik orqali. U virusning 25K harakat oqsilini taniydi, bunda oqsilning 6-o‘rnidagi izoleysin aminokislotasi muhim (Malcuit va boshq., 1999). KYVga qarshi Ny genlari gipersezgirlik, Ry genlari esa ekstremal chidamlilik beradi. Ry-adg (S. andigenadan) XI xromosomada, Ry-sto (S. stoloniferumdan) XII xromosomada joylashgan, Ry-chc esa S. chacoense turidan olingan. Grech-Baran va boshq. (2020) Ry-sto geni TIR-NLR oqsilini kodlashini, uning ishlashi haroratga bog‘liq emasligini va EDS1 hamda NRG1 oqsillarini talab qilishini ko‘rsatdi. 4. RNK-interferensiya Genlardan tashqari, o‘simlikda RNK-interferensiya tizimi ham virusga qarshi ishlaydi: u virus RNKsini parchalaydi. Viruslar bunga qarshi susaytirish supressorlarini hosil qiladi. KXVda bu p25 oqsili, KYVda HC-Pro oqsilidir. Muhokama Ekstremal chidamlilik beradigan genlar (Rx1, Rx2, Ry) seleksiya uchun qulayroq, chunki ular virusni nekroz hosil qilmasdan erta to‘xtatadi. Lekin bu genlarning hammasi virus oqsilini tanishga asoslangan. Agar virus oqsili o‘zgarsa, gen uni tanimay qo‘yishi mumkin. Bendahmane va boshq. (1995) qobiq oqsilini KXV shtammlariga xos elitsitor deb ko‘rsatgan, ya’ni shtammlar orasida farq bor. Shu sababli bitta genga tayanish xavfli. Bitta navda turli mexanizmli genlarni birlashtirish (piramidalash) va virus shtammlarini kuzatib borish maqsadga muvofiq. Genlarning ketma-ketligi ma’lum bo‘lgani uchun ularni PCR markerlari bilan aniqlash mumkin. Rx1 geni kartoshka sista nematodasiga (Globodera pallida) qarshi Gpa2 geniga juda o‘xshash. Ikkala gen XII xromosomada bir-biridan atigi 0,02 cM masofada joylashgan va bitta yovvoyi shakldan (S. tuberosum ssp. andigena CPC 1673) o‘tgan (Rouppe van der Voort va boshq., 1999; Van der Vossen va boshq., 2000). Shuning uchun praymerlar Rx1ga xos bo‘lishi va Gpa2 bilan chalkashmasligi kerak. Buni dastlab kompyuterda (Primer-BLAST, in silico PCR) tekshirish vaqt va mablag‘ni tejaydi. Bu ishning cheklovi shuki, mexanizmlar asosan xorijiy navlar va model o‘simliklarda o‘rganilgan. Mahalliy navlarda genlarning bor-yo‘qligini alohida tekshirish kerak. Xulosa 1. Kartoshkadagi virusga chidamlilik genlarining ko‘pchiligi NLR oqsillarini kodlaydi va virus oqsilini tanib, gipersezgirlik yoki ekstremal chidamlilikni ishga tushiradi. 2. Rx1 va Rx2 KXV qobiq oqsilini taniydi, Nb esa 25K harakat oqsilini taniydi. Ry genlari KYVga qarshi ekstremal chidamlilik beradi. 3. Viruslar o‘zgarib chidamlilikni yengib o‘tishi mumkin, shuning uchun turli genlarni birlashtirish zarur. 4. Gen-spetsifik praymerlar chidamli navlarni tez va aniq tanlash imkonini beradi.

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Ma’lumot topilmadi

ma’lumot kutilmoqda

Adabiyotlar15 ta adabiyot

Ahmadvand R., Wolf I., Gorji A.M., Polgár Z., Taller Y. Development of molecular tools for distinguishing between the highly similar Rx1 and Rx2 PVX extreme resistance genes in tetraploid potato // Potato Research. 2013. Vol. 56. P. 277–291.
Bendahmane A., Köhm B.A., Dedi C., Baulcombe D.C. The coat protein of potato virus X is a strain-specific elicitor of Rx1-mediated virus resistance in potato // Plant Journal. 1995. Vol. 8(6). P. 933–941.
Bendahmane A., Kanyuka K., Baulcombe D.C. The Rx gene from potato controls separate virus resistance and cell death responses // Plant Cell. 1999. Vol. 11(5). P. 781–792.
Bendahmane A., Querci M., Kanyuka K., Baulcombe D.C. Agrobacterium transient expression system as a tool for the isolation of disease resistance genes: application to the Rx2 locus in potato // Plant Journal. 2000. Vol. 21(1). P. 73–81.
FAO. International Year of the Potato 2008: New light on a hidden treasure. An end-of-year review. Rome: FAO, 2009.
Grech-Baran M., Witek K., Szajko K., et al. Extreme resistance to Potato virus Y in potato carrying the Rysto gene is mediated by a TIR-NLR immune receptor // Plant Biotechnology Journal. 2020. Vol. 18(3). P. 655–667.
Jones J.D.G., Dangl J.L. The plant immune system // Nature. 2006. Vol. 444. P. 323–329.
Loebenstein G., Berger P.H., Brunt A.A., Lawson R.H. (eds.). Virus and virus-like diseases of potatoes and production of seed-potatoes. Dordrecht: Kluwer Academic Publishers, 2001.
Malcuit I., Marano M.R., Kavanagh T.A., De Jong W., Forsyth A., Baulcombe D.C. The 25- kDa movement protein of PVX elicits Nb-mediated hypersensitive cell death in potato // Molecular Plant-Microbe Interactions. 1999. Vol. 12(6). P. 536–543.
Moffett P., Farnham G., Peart J., Baulcombe D.C. Interaction between domains of a plant NBS-LRR protein in disease resistance-related cell death // EMBO Journal. 2002. Vol. 21(17). P. 4511–4519.
Rouppe van der Voort J., Kanyuka K., van der Vossen E., Bendahmane A., Mooijman P., Klein-Lankhorst R., Stiekema W., Baulcombe D., Bakker J. Tight physical linkage of the nematode resistance gene Gpa2 and the virus resistance gene Rx on a single segment introgressed from the wild species Solanum tuberosum subsp. andigena CPC 1673 into cultivated potato // Molecular Plant-Microbe Interactions. 1999. Vol. 12(3). P. 197–206.
Scholthof K.-B.G., Adkins S., Czosnek H., Palukaitis P., Jacquot E., Hohn T., Hohn B., Saunders K., Candresse T., Ahlquist P., Hemenway C., Foster G.D. Top 10 plant viruses in molecular plant pathology // Molecular Plant Pathology. 2011. Vol. 12(9). P. 938–954.
Tameling W.I.L., Baulcombe D.C. Physical association of the NB-LRR resistance protein Rx with a Ran GTPase-activating protein is required for extreme resistance to Potato virus X // Plant Cell. 2007. Vol. 19(5). P. 1682–1694.
Valkonen J.P.T. Potato viruses: economical losses and biotechnological potential // Potato biology and biotechnology: advances and perspectives / eds. D. Vreugdenhil et al. Amsterdam: Elsevier, 2007. P. 619–641.
Van der Vossen E.A.G., Rouppe van der Voort J.N.A.M., Kanyuka K., Bendahmane A., Sandbrink H. et al. Homologues of a single resistance-gene cluster in potato confer resistance to distinct pathogens: a virus and a nematode // Plant Journal. 2000. Vol. 23(5). P. 567–576.

Metama’lumot

Identifikator: REL-C-014719
Nashr sanasi: 2026-10-30
Til: o‘zbek
Konferensiyalar: Fundamental va amaliy mikrobiologiyaning bugungi holati va rivojlanish istiqbollari: yosh olimlar nigohi
Jild: 1
Son: 8
DOI: 10.66073/refocus-conf/3010033
Nashriyot: LLC Ilm-fan va innovatsiyalar akademiyasi
Sahifa: 123–126
ORCID: Fayziyev Vokhid Bakhramovich0000-0001-5141-6952
Litsenziya: Ushbu ilmiy ish CC BY 4.0 litsenziyasi asosida ochiq.
Metadata manbasi: Partner conferences of the International Scientific Journal Research Focus
Oxirgi yangilanish: 2026-11-03

[1] Ahmadvand R., Wolf I., Gorji A.M., Polgár Z., Taller Y. Development of molecular tools for distinguishing between the highly similar Rx1 and Rx2 PVX extreme resistance genes in tetraploid potato // Potato Research. 2013. Vol. 56. P. 277–291.

[2] Bendahmane A., Köhm B.A., Dedi C., Baulcombe D.C. The coat protein of potato virus X is a strain-specific elicitor of Rx1-mediated virus resistance in potato // Plant Journal. 1995. Vol. 8(6). P. 933–941.

[3] Bendahmane A., Kanyuka K., Baulcombe D.C. The Rx gene from potato controls separate virus resistance and cell death responses // Plant Cell. 1999. Vol. 11(5). P. 781–792.

[4] Bendahmane A., Querci M., Kanyuka K., Baulcombe D.C. Agrobacterium transient expression system as a tool for the isolation of disease resistance genes: application to the Rx2 locus in potato // Plant Journal. 2000. Vol. 21(1). P. 73–81.

[5] FAO. International Year of the Potato 2008: New light on a hidden treasure. An end-of-year review. Rome: FAO, 2009.

[6] Grech-Baran M., Witek K., Szajko K., et al. Extreme resistance to Potato virus Y in potato carrying the Rysto gene is mediated by a TIR-NLR immune receptor // Plant Biotechnology Journal. 2020. Vol. 18(3). P. 655–667.

[7] Jones J.D.G., Dangl J.L. The plant immune system // Nature. 2006. Vol. 444. P. 323–329.

[8] Loebenstein G., Berger P.H., Brunt A.A., Lawson R.H. (eds.). Virus and virus-like diseases of potatoes and production of seed-potatoes. Dordrecht: Kluwer Academic Publishers, 2001.

[9] Malcuit I., Marano M.R., Kavanagh T.A., De Jong W., Forsyth A., Baulcombe D.C. The 25- kDa movement protein of PVX elicits Nb-mediated hypersensitive cell death in potato // Molecular Plant-Microbe Interactions. 1999. Vol. 12(6). P. 536–543.

[10] Moffett P., Farnham G., Peart J., Baulcombe D.C. Interaction between domains of a plant NBS-LRR protein in disease resistance-related cell death // EMBO Journal. 2002. Vol. 21(17). P. 4511–4519.

[11] Rouppe van der Voort J., Kanyuka K., van der Vossen E., Bendahmane A., Mooijman P., Klein-Lankhorst R., Stiekema W., Baulcombe D., Bakker J. Tight physical linkage of the nematode resistance gene Gpa2 and the virus resistance gene Rx on a single segment introgressed from the wild species Solanum tuberosum subsp. andigena CPC 1673 into cultivated potato // Molecular Plant-Microbe Interactions. 1999. Vol. 12(3). P. 197–206.

[12] Scholthof K.-B.G., Adkins S., Czosnek H., Palukaitis P., Jacquot E., Hohn T., Hohn B., Saunders K., Candresse T., Ahlquist P., Hemenway C., Foster G.D. Top 10 plant viruses in molecular plant pathology // Molecular Plant Pathology. 2011. Vol. 12(9). P. 938–954.

[13] Tameling W.I.L., Baulcombe D.C. Physical association of the NB-LRR resistance protein Rx with a Ran GTPase-activating protein is required for extreme resistance to Potato virus X // Plant Cell. 2007. Vol. 19(5). P. 1682–1694.

[14] Valkonen J.P.T. Potato viruses: economical losses and biotechnological potential // Potato biology and biotechnology: advances and perspectives / eds. D. Vreugdenhil et al. Amsterdam: Elsevier, 2007. P. 619–641.

[15] Van der Vossen E.A.G., Rouppe van der Voort J.N.A.M., Kanyuka K., Bendahmane A., Sandbrink H. et al. Homologues of a single resistance-gene cluster in potato confer resistance to distinct pathogens: a virus and a nematode // Plant Journal. 2000. Vol. 23(5). P. 567–576.