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NUTRITIONAL COMPONENT ANALYSIS OF MUNG BEAN (VIGNA RADIATA L.)

S. WANG, M. WU, Q. HU, C. JIANG, S. WANG, Y. YAO, and L. WANG

Citation: Wang S, Wu M, Hu Q, Jiang C, Wang S, Yao Y, Wang L (2025). Nutritional component analysis of mung bean (Vigna radiata L.). SABRAO J. Breed. Genet. 57(3): 1050-1059. http://doi.org/10.54910/sabrao2025.57.3.17.

Summary

Mung bean (Vigna radiata L.), being rich in protein, starch, and bioactive compounds, has become more recognized as a nutritious and healthy food crop. Mining germplasm resources with higher contents of nutritional factors will further improve their efficient use. In the present study, assessing the crude protein, total starch, total polyphenol and flavonoid, and vitexin/isovitexin contents occurred in 400 accessions of mung bean core collection. The results showed that overall, the crude protein content ranged from 21.26% to 31.14% with an average of 25.91%, whereas the total starch content ranged from 34.24% to 59.82% with an average of 40.19%. The total polyphenol content (2.07–5.89 mg/g) was higher than the total flavonoid content (1.13–2.76 mg/g), observing a positive correlation between these two factors. A significant positive correlation also emerged between the contents of vitexin (0.18–2.28 mg/g) and isovitexin (0.15–1.97 mg/g). Similarly, the results also specified that the wild germplasm had higher levels of nutritional components, except for starch. This study will lay a foundation for enhancing the utilization of mung bean resources in developing new varieties with high contents of bioactive compounds.

Mung bean (V. radiata L.), crude protein, total starch, total polyphenol and flavonoid, vitexin and isovitexin

The study showed higher variations for vitexin/isovitexin and flavonoid contents in mung bean (V. radiata L.) genetic resources. The contents of crude protein, total flavonoids, total polyphenols, vitexin, and isovitexin were notable at higher rates in the wild germplasm than breeding cultivars, with no obvious relationships among the contents of nutritional components, except for vitexin and isovitexin.

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SABRAO Journal of Breeding and Genetics
57 (3) 1050-1059, 2025
http://doi.org/10.54910/sabrao2025.57.3.17
http://sabraojournal.org/
pISSN 1029-7073; eISSN 2224-8978

Date published: June 2025

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ASSESSMENT OF HEAT TOLERANCE IN WHEAT (TRITICUM AESTIVUM L.) AT THE SEEDLING STAGE

G.F. MAMATKULOVA, D.SH. ERJIGITOV, O.A. MUKHAMMADIEV, D.B. SOKIBOYEVA, M.D. KHOLOVA, B.B. ORIPOVA, D.K. ERNAZAROVA, S.K. BABOEV, O.S. TURAEV, and F.N. KUSHANOV

Citation: Mamatkulova GF, Erjigitov DSH, Mukhammadiev OA, Sokiboyeva DB, Kholova MD, Oripova BB, Ernazarova DK, Baboev SK, Turaev OS, Kushanov FN (2025). Assessment of heat tolerance in wheat (Triticum aestivum L.) at the seedling stage. SABRAO J. Breed. Genet. 57(3): 1041-1049. http://doi.org/10.54910/sabrao2025.57.3.16.

Summary

Wheat (Triticum aestivum L.) is a primary food crop globally, susceptible to environmental fluctuations, particularly the heat stress. With climate change and its increasing threats, wheat’s response evaluation to the elevated temperatures during early growth stages is crucial for securing the crop and its yield. The presented study sought to assess the heat tolerance in 15 common wheat cultivars at the seedling stage under controlled conditions. Growing wheat cultivars ensued at the optimal (25 °C) and heat-stressed (35 °C) conditions. After seven days of growth, the seedling height, root length, and root number’s measurement underwent assessment for heat stress effects. The quantification of the degree of plants damaged used the Damage Index (DI), with heat tolerance evaluated using the Heat Tolerance Index (HTI). The results showed significant variations among the wheat cultivars for heat tolerance, and the two cultivars, Tuyatish and Nodir, displayed minimal growth reduction under the highest temperature, achieving maximum HTI values (98.3% and 89.6%, respectively). The cultivar FAWWON-IRR-D-77 exhibited a substantial damage and low tolerance under high-temperature stress. This study provides essential insights into heat tolerance mechanisms in wheat, forming a foundation for future breeding and molecular studies aimed at improving heat resilience in crops.

Common wheat (T. aestivum L.), heat stress, seedling stage, early growth stage, heat tolerance, damage index, heat tolerance index

This study identified significant variability among the common wheat (T. aestivum L.) genotypes for heat tolerance at the seedling stage. Cultivars Tuyatish and Nodir demonstrated the highest heat tolerance with minimal damage. The line FAWWON-IRR-D-77 exhibited substantial damage and low tolerance to high-temperature stress.

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SABRAO Journal of Breeding and Genetics
57 (3) 1041-1049, 2025
http://doi.org/10.54910/sabrao2025.57.3.16
http://sabraojournal.org/
pISSN 1029-7073; eISSN 2224-8978

Date published: June 2025

« Back to main page of SABRAO Journal of Breeding and Genetics Vol. 57 No. 3

EVALUATION OF DOUBLED HAPLOID RICE LINES DERIVED FROM ANTHER CULTURE OF INPARI-42 GSR AGRITAN MUTANT IN ADVANCED YIELD TRIALS

B.S. PURWOKO, A.P. NADIA, I.S. DEWI, R. HERAWATI, and M.F. ANSHORI

Citation: Purwoko BS, Nadia AP, Dewi IS, Herawati R, Anshori MF (2025). Evaluation of doubled haploid rice lines derived from anther culture of Inpari-42 GSR Agritan mutant in advanced yield trials. SABRAO J. Breed. Genet. 57(3): 1030-1040. http://doi.org/10.54910/sabrao2025.57.3.15.

Summary

Combining mutation and anther culture technology could accelerate the rice breeding program. This research evaluated the agronomic performance of doubled haploid (DH) lines derived from anther culture of Inpari 42 GSR Agritan mutant in advanced yield trials. The research was conducted in Bogor and Indramayu, Indonesia from December 2023 to April 2024. The material used was 18 DH lines and two check varieties (Inpari 42 GSR Agritan and Inpari 45 Agritan). The experiment used a randomized complete block design with three replications and the rice genotype as a factor. The result indicated a variation in agronomic characters, yield components, and productivity of the DH rice lines. The interaction of genotype by location was notable in the vegetative plant height, the number of vegetative tillers, the percentage of filled and empty grains, the weight of 1000 grains, and productivity. Highly significant and positive correlations to productivity were evident in total grain, percentage of filled grains, the number of filled grains, grain filling period, and days of harvesting. Selection used the index selection method based on productivity, percentage of filled grains, total grains, and days of harvesting, which indicated the 12 DH lines need further evaluation in multi-location trials for their potential yield and stability.

anther culture, correlation, green super rice, mutation, selection index, yield

The combined mutation and anther culture technology, followed by the index selection method, can accelerate in obtaining advanced rice lines with good agronomic and productivity characteristics.

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SABRAO Journal of Breeding and Genetics
57 (3) 1030-1040, 2025
http://doi.org/10.54910/sabrao2025.57.3.15
http://sabraojournal.org/
pISSN 1029-7073; eISSN 2224-8978

Date published: June 2025

« Back to main page of SABRAO Journal of Breeding and Genetics Vol. 57 No. 3

EVALUATION OF BRASSICA GENOTYPES FOR MORPHOLOGICAL AND BIOCHEMICAL ATTRIBUTES

H.R.M.W. ASLAM and A. QAYYUM

Citation: Aslam HRMW, Qayyum A (2025). Evaluation of brassica genotypes for morphological and biochemical attributes. SABRAO J. Breed. Genet. 57(3): 1021-1029. http://doi.org/10.54910/sabrao2025.57.3.14.

Summary

Brassica napus is an important oilseed crop, commonly grown worldwide for multiple purposes. A planned multi-locational experiment evaluated the morphophysiological and biochemical attributes of 80 Brassica napus genotypes. Data on various yield and oil-related parameters were evaluated for genetic variability, correlation (r), and broad-sense heritability (h2BS). The analysis of variance revealed the presence of significant variations among the genotypes for all studied traits. Plant height showed a positive correlation with erucic acid (0.290) and glucosinolate (0.264), while secondary branches revealed a positive association with seed yield (2.90). Principal component analysis indicated the first five principal components (PCs) contributed up to 73% of total variation, with the first PC representing the highest percentage of variation. The biplot suggests the genotypes Faisal canola, Diamond canola, AS-333, and HMU-1621B can be beneficial in breeding programs to develop new accessions with improved seed yield and oil-related attributes of Brassica.

Biplot, genetic diversity, heritability, oil contents, PCA

Eighty accessions, screened for yield and related traits, indicated the genotypes Faisal canola, Diamond canola, AS-333, and HMU-1621B exhibited the highest yield and fatty acid content.

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SABRAO Journal of Breeding and Genetics
57 (3) 1021-1029, 2025
http://doi.org/10.54910/sabrao2025.57.3.14
http://sabraojournal.org/
pISSN 1029-7073; eISSN 2224-8978

Date published: June 2025

« Back to main page of SABRAO Journal of Breeding and Genetics Vol. 57 No. 3

BLACK PEPPER (PIPER NIGRUM L.) F1 HYBRIDS ASSESSMENT AGAINST THE STEM BORER (LOPHOBARIS PIPERIS) UNDER LABORATORY CONDITIONS

R. ROHIMATUN, S. SISWANTO, N. BERMAWIE, S. WAHYUNI, N.L.W. MEILAWATI, M. SUSILOWATI, D.W. UTAMI, R. HERYANTO, S.F. SYAHID, and D. DADANG

Citation: Rohimatun R, Siswanto S, Bermawie N, Wahyuni S, Meilawati NLM, Susilowati M, Utami DW, Heryanto R, Syahid SF, Dadang D (2025). Black pepper (Piper nigrum L.) F1 hybrids assessment against the stem borer (Lophobaris piperis) under laboratory conditions. SABRAO J. Breed. Genet. 57(3): 1009-1020. http://doi.org/10.54910/sabrao2025.57.3.13.

Summary

Yield losses from stem borer (Lophobaris piperis Marsh) attacks have reached up to 72%. Currently, information on the preference and intensity of stem borer attacks on the black pepper (Piper nigrum L.) is limited. This study represents the first effort to develop stem borer-resistant black pepper cultivars through inter-varietal hybridization. An assessment of stem borer preferences among 29 F1 hybrids and their five parental cultivars commenced under laboratory conditions. The F1 hybrids displayed a range of resistance levels, resulting from crosses between highly preferred and less preferred black pepper varieties. Among the 29 F1 hybrids, three were highly resistant, 12 were resistant, 10 were moderately resistant, two were moderately susceptible, one was susceptible, and one was highly susceptible. These findings provide a promising material for developing resistant black pepper cultivars that support integrated pest management strategies and promote environmental sustainability.

Black pepper (Piper nigrum L.), stem borer (Lophobaris piperis), damage intensity, inter-varietal hybridization, preference, resistance, susceptible

The F1 hybrids production with various resistant categories came through hybridization among the preferred and less preferred black peppers (P. nigrum L.) cultivars. Three and 12 F1 hybrids attained categories as highly resistant and resistant, respectively, which can undergo studies for developing resistant cultivars in black pepper.

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SABRAO Journal of Breeding and Genetics
57 (3) 1009-1020, 2025
http://doi.org/10.54910/sabrao2025.57.3.13
http://sabraojournal.org/
pISSN 1029-7073; eISSN 2224-8978

Date published: June 2025

« Back to main page of SABRAO Journal of Breeding and Genetics Vol. 57 No. 3

PHENOTYPIC VARIABILITY IN THE VEGETATIVE GENERATION OF GRAPE (V. VINIFERA L.) PROTOCLONES AND SELECTION WITH STABLE SIGNS

V.S. SALIMOV, M.A. HUSEYNOV, and A.S. HUSEYNOVA

Citation: Salimov VS, Huseynov MA, Huseynova AS (2025). Phenotypic variability in the vegetative generation of grape (V. vinifera L.) protoclones and selection with stable signs. SABRAO J. Breed. Genet. 57(3): 999-1008. http://doi.org/10.54910/sabrao2025.57.3.12.

Summary

The following study sought to determine yield stability and quality traits in the vegetative generation of highly productive protoclones chosen through clonal selection in promising grape (V. vinifera L.) genotypes. In determining the degree of variability for various quantitative and qualitative traits, growing the selected protoclones of grapevines occurred under various environmental conditions. The newly identified clonal forms proved genetically more stable and homogeneous than their parental genotypes for phenotypic traits, and the degree of variability in their vegetative populations was much lower. The results further enunciated the relative stability of yield and quality parameters of highly productive clones, showing chief association with the clonal breeding.

Grape (V. vinifera L.), promising populations, genetic variability, stability, yield and quality traits, quantitative and qualitative variables

The newly identified clonal forms of the grape (V. vinifera L.) appeared genetically more stable and homogeneous than their parental genotypes for phenotypic traits. The degree of variability in their vegetative population was much lower, and the clonal selection was successful.

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SABRAO Journal of Breeding and Genetics
57 (3) 999-1008, 2025
http://doi.org/10.54910/sabrao2025.57.3.12
http://sabraojournal.org/
pISSN 1029-7073; eISSN 2224-8978

Date published: June 2025

« Back to main page of SABRAO Journal of Breeding and Genetics Vol. 57 No. 3

ENGINEERING OF sgRNA TARGETING DREB4 ISOLATED FROM ORYZA SATIVA USING pRGEB32 IN CRISPR/Cas9 GENE EDITING

M.H. FENDIYANTO, I.R. KURNIYANTO, E. SETIAWAN, M.P. PRATAMI, F.S. FASTANTI, C. WOSONOWATI, Z. ZULKHAERIYAH, E.Q. AZZAHROH, and D.P. VARDHANA

Citation: Fendiyanto MH, Kurniyanto IR, Setiawan E, Pratami MP, Fastanti FS, Wosonowati C, Zulkhaeriyah Z, Azzahroh EQ, Vardhana DP (2025). Engineering of sgRNA targeting DREB4 isolated from Oryza sativa using pRGEB32 in CRISPR/Cas9 gene editing. SABRAO J. Breed. Genet. 57(3): 989-998. http://doi.org/10.54910/sabrao2025.57.3.11.

Summary

Rice (Oryza sativa L.) production on acidic drylands needs optimization through breeding to develop drought-resistant cultivars. The rice cultivar Inpago-5 showed moderate tolerance to drought stress. However, further enhancing this tolerance can proceed by precise gene editing of the DREB (dehydration-responsive element binding) gene, utilizing the CRISPR (clustered regularly interspaced palindromic repeats) /Cas9 system. The presented study aimed to engineer the single-guided RNA (sgRNA) targeting the DREB gene in rice cultivar Inpago-5 using the CRISPR/Cas9 approach. The use of the pRGEB32 plasmid harboring the Cas9 gene helped facilitate the cloning of sgRNA-DREB through the Golden Gate cloning technique and heat shock transformation. The sgRNA-DREB confirmation succeeded through colony PCR (polymerase chain reaction), followed by plasmid isolation and subsequent validation using PCR with specific primers and sequencing primers targeting the pRGEB32 vector. The latest analysis revealed the pRGEB32-sgDREB_sg1 and pRGEB32-sgDREB_sg2 transformed bacteria proliferated on LB+Kanamycin selection media, indicating successful insertion of sgRNA-DREB into the pRGEB32 plasmid. The verification through colony and plasmid PCR with specific primers confirmed the presence of a 425-bp amplicon. Sequencing data showed the correct insertion of sgRNA-DREB sequences at the designated site.

Rice (O. sativa L.), cultivar Inpago-5, plant breeding, CRISPR/Cas9, gene editing

The gene encoding rice (O. sativa L.) drought tolerance DREB4 succeeded integration into the pRGEB32 vector, with the sgRNA-DREB accurately inserted at the targeted site.

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SABRAO Journal of Breeding and Genetics
57 (3) 989-998, 2025
http://doi.org/10.54910/sabrao2025.57.3.11
http://sabraojournal.org/
pISSN 1029-7073; eISSN 2224-8978

Date published: June 2025

« Back to main page of SABRAO Journal of Breeding and Genetics Vol. 57 No. 3

HETEROSIS AND INBREEDING DEPRESSION IN POTATO (SOLANUM TUBEROSUM L.) HYBRIDS DEVELOPED THROUGH TRUE POTATO SEED IN NORTHERN PAKISTAN

B.A. KHAN, N. ALI, S. ALI, S. ASLAM, and A.A. MALIK

Citation: Khan BA, Ali N, Ali S, Aslam S, Malik AA (2025). Heterosis and inbreeding depression in potato (Solanum tuberosum L.) hybrids developed through true potato seed in Northern Pakistan. SABRAO J. Breed. Genet. 57(3): 977-988. http://doi.org/10.54910/sabrao2025.57.3.10.

Summary

Heterosis and inbreeding depression are key considerations in potato breeding, particularly in regions with limited resources. Heterosis (F1) and inbreeding depression (F2) investigations ensued in 2022–2024 in eight different potato (Solanum tuberosum L.) hybrids generated from true potato seeds in Abbottabad and Battakundi, Pakistan. A randomized complete block design experiment transpired with three replications. The analysis of variance revealed significant differences among the potato hybrids for all traits, except the tuber germination percentage, signifying a substantial level of genetic variability. The Cardinal × Roko hybrid has demonstrated exceptional hybrid vigor for the number of medium tubers plot-1, with high estimates of relative heterosis (61.78%), heterobeltiosis (61.78%), and standard heterosis (86.01%). The negative inbreeding depression observed for all studied traits, ranging from –0.364% to –187.67%, indicates inbreeding has significantly impacted the performance of hybrids. The noted positive correlations among potato traits provide valuable insights for breeding programs. Results concluded the cross combinations, Cardinal × Roko and Kuroda × Burna, exhibited significant and high heterotic values for the number and weight of medium tubers plot-1, while desirable negative heterosis for days to maturity. It confirms these crosses have strong potential for release as new potato hybrids. Further evaluation of these hybrids in future breeding strategies can probe ultimate potential for resilience, yield, and other important traits.

Potato, true potato seed, heterosis, inbreeding depression, Cardinal × Roko, Kuroda × Burna

Given the heterotic effects, the hybrids’ cross combinations demonstrated superior mean performance for the evaluated traits compared with their parental cultivars. Cardinal × Roko and Kuroda × Burna were the best crosses due to the high number and weight of medium tubers plot-1, positive heterosis, and desirable negative heterosis for days to maturity. Additionally, Battakundi has become a valid ideal environment for potato cultivation and future breeding programs.

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SABRAO Journal of Breeding and Genetics
57 (3) 977-988, 2025
http://doi.org/10.54910/sabrao2025.57.3.10
http://sabraojournal.org/
pISSN 1029-7073; eISSN 2224-8978

Date published: June 2025

« Back to main page of SABRAO Journal of Breeding and Genetics Vol. 57 No. 3

COMBINING ABILITY AND HETEROTIC EFFECTS IN YELLOW MAIZE FOR MORPHOLOGICAL AND YIELD COMPONENTS

A. ALI, S.A. KHAN, N. ALI, S. ALI, I. HUSSAIN, S.M. KHAN, H. RAZA, and M.Y. KHAN

Citation: Ali A, Khan SA, Ali N, Ali S, Hussain I, Khan SM, Raza H, and Khan MY (2025). Combining ability and heterotic effects in yellow maize for morphological and yield components. SABRAO J. Breed. Genet. 57(3): 968-976. http://doi.org/10.54910/sabrao2025.57.3.9.

Summary

The success of hybrid maize (Zea mays L.) breeding depends on understanding the combining ability effects of the inbred lines. The study aimed to evaluate the general combining ability (GCA), specific combining ability (SCA), and heterotic effects in yellow maize for various traits. Ten inbred lines’ crossing with three testers used the line-by-tester mating scheme. The resulting 30 F1 hybrids underwent evaluation against two check genotypes in a randomized complete block design (RCBD) with three replications during the kharif growing season of 2021 at the Cereal Crop Research Institute (CCRI), Nowshehra, Pakistan. Significant heterotic and combining effects were evident for the studied traits. Inbred lines YL-02 and YL-06 showed significant positive GCA for ear length, 100-grain weight, and grain yield. The F1 hybrids YL-02 × YD-04 (8209.6 kg ha-1) and YL-01 × YL-07 RC (6979.6 kg ha-1) revealed maximum grain yield and significant positive SCA effects, indicating potential for yield improvement. Estimates of SCA were greater than GCA for all the studied traits, signifying the importance of the non-additive gene action in the inheritance of studied traits. Based on combing ability effects and yield performance, the F1 crosses YL-02 × YD-04 and YL-01 × YL-07 RC are desirable for future breeding programs.

Maize (Z. mays L.), hybrids, mean performance, combining ability, heterosis, heterobeltiosis, grain yield, gene action.

The maize (Z. mays L.) inbred lines YL-01, YL-02, and YL-06 emerged with desirable and significant GCA effects. The F1 hybrids YL-02 × YD-04 and YL-01 × YL-07 RC exhibited with significant SCA effects and maximum grain yield. These parents and hybrids could be beneficial in future hybridization programs of yellow maize.

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SABRAO Journal of Breeding and Genetics
57 (3) 967-976, 2025
http://doi.org/10.54910/sabrao2025.57.3.9
http://sabraojournal.org/
pISSN 1029-7073; eISSN 2224-8978

Date published: June 2025

« Back to main page of SABRAO Journal of Breeding and Genetics Vol. 57 No. 3

MOLECULAR-GENETIC ANALYSIS OF CEREAL YELLOW DWARF VIRUS-RPV (CYDV-RPV) AFFECTING WHEAT (TRITICUM AESTIVUM L.) IN UZBEKISTAN

T.KH. MAKHMUDOV, Z.N. KADIROVA, Z.M. ZIYAEV, B.SH. ADILOV, A.G. SHERIMBETOV, N.E. CHORSHANBIEV, O.R. ERGASHEV, Z.A. UMAROV, S.U.NOROVA, M.R. TOKHIROVA, K. ASTANAKULOV, R. EGAMBERDIEV, U.X. YULDASHOV, and V.B. FAYZIEV

Citation: Makhmudov TKH, Kadirova ZN, Ziyaev ZM, Adilov BSH, Sherimbetov AG, Chorshanbiev NE, Ergashev OR, Umarov ZA, Norova SU, Tokhirova MR, Astanakulov K, Egamberdiev R, Yuldashov UX, Fayziev VB (2025). Molecular-genetic analysis of cereal yellow dwarf virus-RPV (CYDV-RPV) affecting wheat (Triticum aestivum L.) in Uzbekistan. SABRAO J. Breed. Genet. 57(3): 957-967. http://doi.org/10.54910/sabrao2025.57.3.8.

Summary

Wheat (Triticum aestivum L.) is the most widely cultivated cereal worldwide, a staple food for 40% of the world’s population that contributes 20% of total dietary requirements. Yellow dwarf viruses are viruses affecting the grain yield and its quality and damaging the cereal crops economically. Yellow dwarf viruses are the most economically important plant viruses impacting cereal production worldwide, which include viruses from the genus Luteovirus BYDV-PAV, MAV, and Polerovirus CYDV-RPV. The Cereal Yellow Dwarf Virus (CYDV-RPV) is one of the most dangerous YDV viral species. Barley/cereal yellow dwarf viruses cause yellow dwarf disease, which is a continuous risk to cereal crop production globally. These viruses cause leaf yellowing and stunting, resulting in yield reduction of up to 80%. They are the most critical viral diseases of cereals worldwide. They have a wide host range, which includes wheat, barley, oats, and over 150 grass species. These viral infections have the potential to spread epidemics in Uzbekistan’s wheat fields during specific seasons and play a crucial role in the epidemiological process. Using the KibrayT1 isolate of the CYDV-RPV strain, the following study aimed to determine the prevalence of CYDV-RPV in wheat fields in the District of Kibray, Tashkent Region, Uzbekistan. The use of reverse transcription-polymerase chain reaction succeeded in making the diagnosis.

Wheat (T. aestivum L.), cereal yellow dwarf virus (CYDV), RT-PCR, CP gene, phylogenetic analysis, KibrayT1

The CYDV-RPV isolate detection was successful by RT-PCR in wheat (T. aestivum L.). The determination of the nucleotide sequence of the coat protein gene prevailed in this isolate. The phylogenetic analysis revealed its close relatedness to the Jordan CYDV-RPV isolate ‘Jordon’ (HQ206716.1).

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SABRAO Journal of Breeding and Genetics
57 (3) 957-967, 2025
http://doi.org/10.54910/sabrao2025.57.3.8
http://sabraojournal.org/
pISSN 1029-7073; eISSN 2224-8978

Date published: June 2025

« Back to main page of SABRAO Journal of Breeding and Genetics Vol. 57 No. 3