Combining Ability and Genetic Parameters of Maize Based on S4 Diallel-Derived Progenies
Abstract
Maize is an important crop with strategic roles. The development of superior maize varieties requires genetically diverse parental lines with high combining ability. Advanced selfing generations, such as S4, provide increased homozygosity, allowing more reliable estimation of combining ability and genetic parameters for parental selection. This study aimed to evaluate general combining ability (GCA), specific combining ability (SCA), and genetic parameters using S4 progenies derived from a half-diallel mating design. The experiment was conducted from August to November 2025 at PT. Sukses Seed Sentosa, Malang Regency, East Java, Indonesia, using a randomized complete block design with two replications and involving 36 S4 cross combinations. The first two principal components of the GCA explained 82.36% of the total variation (PC1 = 63.26%; PC2 = 19.10%). P4 exhibited superior GCA for cob diameter and days to harvest, whereas P1 showed favorable GCA for multiple yield-related traits. SCA values showed that the first two principal components explained 84.54% of the total variation (PC1 = 74.46%; PC2 = 10.08%). P2 × P6 showed the highest SCA for multiple yield-related traits, while P2 × P4 was superior for days to silking and shelling percentage. High GCV was observed in AWC (25.26), SWC (25.62), and Y (23.79). High narrow-heritability was observed for DA (0.55), cob length (0.60), and NSPR (0.66). Most traits were predominantly controlled by additive gene action, whereas grain yield and number of seeds per cob were mainly governed by non-additive gene action.
Keywords
References
Acquaah, G. (2012). Principles of Plant Genetics and Breeding (Second). Wiley-Blackwell.
Akinwale, R. O., Eze, C. E., Traore, D., & Menkir, A. (2021). Detection of Non-Additive Gene Action within Elite Maize Populations Evaluated in Contrasting Environments under Rainforest Ecology in Nigeria. Crop Breeding, Genetics and Genomics, 3(1). https://doi.org/10.20900/cbgg20210003
Akinwale, R. O., Fakorede, M. A. B., Badu-Apraku, B., & Oluwaranti, A. (2014). Assessing the usefulness of GGE biplot as a statistical tool for plant breeders and agronomists. Cereal Research Communications, 42(3), 534–546. https://doi.org/10.1556/CRC.42.2014.3.16
Alam, M. A., Rahman, M., Ahmed, S., Jahan, N., Khan, M. A.-A., Islam, M. R., Alsuhaibani, A. M., Gaber, A., & Hossain, A. (2022). Genetic Variation and Genotype by Environment Interaction for Agronomic} Traits} in Maize (Zea mays L.) Hybrids. Plants, 11(11), 1522. https://doi.org/10.3390/plants11111522
Azrai, M., Aqil, M., Efendi, R., Andayani, N. N., Suwardi, Zainuddin, B., Pabendon, M. B., Sitaresmi, T., Anshori, M. F., Riadi, M., Yasin, M., Laurenze, R., Bahtiar, Suwarti, & Syam, A. (2025). Integrating multi-trait and multi-index approaches for identifying drought-tolerant tropical maize genotypes. Frontiers in Sustainable Food Systems, 9, 1608307. https://doi.org/10.3389/fsufs.2025.1608307
Bhullar, G. S., Gill, K. S., & Khehra, A. S. (1979). Combining ability analysis over F1-F5 generations in diallel crosses of bread wheat. Theoretical and Applied Genetics, 55(2), 77–80. https://doi.org/10.1007/BF00285194
BPS. (2026). Jumlah curah hujan di Kota Malang (milimeter (mm)). https://malangkota.bps.go.id/id/statistics-table/2/NTA4IzI=/jumlah-curah-hujan-di-kota-malang.html
Brown, J., Caligari, P., & Campos, H. (2014). Plant breeding. John Wiley & Sons.
Chaudhary, R. C. (2019). Introductory Principles of Plant Breeding (Second). Oxford & IBH Publishing Co. Pvt. Ltd.
Cruz, C. D., Regazzi, A. J., & Carneiro, P. C. S. (2012). Biometric models applied to genetic improvement. Viçosa, MG: UFV, 1, 514.
Cyplik, A., Sobiech, A., Tomkowiak, A., & Bocianowski, J. (2022). Genetic Parameters for Selected Traits of Inbred Lines of Maize (Zea mays L}.). Applied Sciences, 12(14), 6961. https://doi.org/10.3390/app12146961
Damtie, Y., Assefa, G., & Mulualem, T. (2021). Genetic variability, heritability, trait associations and path coefficient analysis of maize (Zea mays L.) inbreed lines. Journal of Current Opinion in Crop Science, 2(1), 86–94. https://doi.org/10.62773/jcocs.v2i1.22
Elayaraja, K., Gadag, R. N., Kumari, J., & Mishra, U. (2018). Combining ability and gene action in experimental hybrids of Sweet Corn ( Zea mays var. saccharata ). Indian Journal of Horticulture, 75(1), 64. https://doi.org/10.5958/0974-0112.2018.00011.7
Faheem, M., Arain, S. M., Sial, M. A., Laghari, K. A., & Qayyum, A. (2023). Genotype by yield*trait (GYT) biplot analysis: a novel approach for evaluating advance lines of durum wheat. Cereal Research Communications, 51(2), 447–456. https://doi.org/10.1007/s42976-022-00298-7
Fasahat, P. (2016). Principles and Utilization of Combining Ability in Plant Breeding. Biometrics & Biostatistics International Journal, 4(1), 1–24. https://doi.org/10.15406/bbij.2016.04.00085
Hassani, M., Mahmoudi, S. B., Saremirad, A., & Taleghani, D. (2024). Genotype by environment and genotype by yield*trait interactions in sugar beet: analyzing yield stability and determining key traits association. Scientific Reports, 13(1), 23111. https://doi.org/10.1038/s41598-023-51061-9
He, Z.-H., Xiao, Y., Lv, Y.-W., Yeh, F. C., Wang, X., & Hu, X.-S. (2023). Prediction of Genetic Gains from Selection in Tree Breeding. Forests, 14(3), 520. https://doi.org/10.3390/f14030520
Hosseini, S. M. S., Shiri, M., Mostafavi, K., Mohammadi, A., & Miri, S. M. (2025). Genetic analysis and association detection of agronomic traits in maize genotypes. Scientific Reports, 15(1), 399. https://doi.org/10.1038/s41598-024-84471-4
Kar, D. K., & Halder, S. (2020). Plant Breeding, Biometry & Biotechnology. New Central Book Agency.
Kearsey, M. J., & Pooni, H. (1996). Genetical analysis of quantitative traits. Garland Science.
Kendal, E. (2019). Comparing durum wheat cultivars by genotype × yield ×trait and genotype × trait biplot method. Chilean Journal of Agricultural Research, 79(4), 512–522. https://doi.org/10.4067/S0718-58392019000400512
Merrick, L. F., Glover, K. D., Yabwalo, D., & Byamukama, E. (2020). Use of Genotype by Yield*Trait (GYT) Analysis to Select Hard Red Spring Wheat with Elevated Performance for Agronomic and Disease Resistance Traits. Crop Breeding, Genetics and Genomics, 2(2). https://doi.org/10.20900/cbgg20200009
Mufidah, N., Sugiharto, A. N., & Waluyo, B. (2021). Assessment of combining ability in purple corn parents under line × tester mating design using GGE biplot. Biodiversitas Journal of Biological Diversity, 22(10), 4545–4554. https://doi.org/10.13057/biodiv/d221048
Mutimaamba, C., Macrobert, J., Cairns, J. E., Magorokosho, C. E., Ndhlela, T., Mukungurutse, C., Minnaar-Ontong, A., & Labuschagne, M. T. (2017). Diallel analysis of acid soil tolerant and susceptible maize inbred lines for grain yield under acid and non-acid soil conditions. Euphytica, 213(4). https://doi.org/10.1007/s10681-017-1877-5
Nguyen, H. T. H., Chen, Z.-Q., Fries, A., Berlin, M., Hallingbäck, H. R., & Wu, H. X. (2022). Effect of additive, dominant and epistatic variances on breeding and deployment strategy in Norway spruce. Forestry: An International Journal of Forest Research, 95(3), 416–427. https://doi.org/10.1093/forestry/cpab052
Oliveira, T. R. A. de, Gravina, G. de A., Oliveira, G. H. F. de, Araújo, K. C., Araújo, L. C. de, Daher, R. F., Vivas, M., Gravina, L. M., & Cruz, D. P. da. (2018). The GT biplot analysis of green bean traits. Ciência Rural, 48(6), e20170757. https://doi.org/10.1590/0103-8478cr20170757
Peixoto, M. A., Evangelista, J. S. P. C., Coelho, I. F., Carvalho, L. P., Farias, F. J. C., Teodoro, P. E., & Bhering, L. L. (2022). Genotype selection based on multiple traits in cotton crops: The application of genotype by yield*trait biplot. Acta Scientiarum. Agronomy, 44, e54136. https://doi.org/10.4025/actasciagron.v44i1.54136
Putri, L. D. N., Saptadi, D., & Waluyo, B. (2022). Analisis Daya Gabung dan Aksi Gen Jagung (Zea mays L) menggunakan Rancangan Perkawinan Line x Tester. Agriprima : Journal of Applied Agricultural Sciences, 6(2), 191–201. https://doi.org/10.25047/agriprima.v6i2.492
Rouf Shah, T., Prasad, K., & Kumar, P. (2016). Maize A potential source of human nutrition and health: A review. Cogent Food & Agriculture, 2(1), 1166995. https://doi.org/10.1080/23311932.2016.1166995
Sary, D. N., Badriyah, L., Sihombing, R. D., Syauqy, T. A., Mustikarini, E. D., Prayoga, G. I., Santi, R., & Waluyo, B. (2022). Estimation of Heritability and Association Analysis of Agronomic Traits Contributing to Yield on Upland Rice ( Oryza sativa L.). Plant Breeding and Biotechnology, 10(4), 232–243. https://doi.org/10.9787/PBB.2022.10.4.232
Shirinpour, M., Atazadeh, E., Bybordi, A., Monirifar, H., Amini, A., Hossain, M. A., Aharizad, S., & Asghari, A. (2023). Gene action and inheritance of grain yield and root morphological traits in hybrid maize grown under water deficit conditions. South African Journal of Botany, 161, 180–191. https://doi.org/10.1016/j.sajb.2023.08.016
Shojaei, S., Mostafavi, K., Ansarifard, I., Bihamta, M., Zeinalzadeh-Tabrizi, H., Omrani, A., Göre, M., & Mousavi, S. M. N. (2023). Comparison of genotype × trait and genotype × yield-trait biplots in Sunflower cultivars. International Journal of Agriculture Environment and Food Sciences, 7(1), 136–147. https://doi.org/10.31015/jaefs.2023.1.17
Singh, D. P., Singh, A. K., & Singh, A. (2021). Plant breeding and cultivar development. Academic Press.
Singh, R. K., & Chaudhary, B. D. (1981). Biometrical methods in quantitative genetic analysis.
Stansfield, R. (1983). Genetika. Terjemahan oleh: Mohidin A, Apandi, Lanny T. 1991. Erlangga. Jakarta, 182.
Stansluos, A. A. L., Öztürk, A., Niedbała, G., Türkoğlu, A., Haliloğlu, K., Szulc, P., Omrani, A., Wojciechowski, T., & Piekutowska, M. (2023). Genotype–Trait (GT) Biplot Analysis for Yield and Quality Stability in Some Sweet Corn (Zea mays L. saccharata Sturt.) Genotypes. Agronomy, 13(6), 1538. https://doi.org/10.3390/agronomy13061538
Sumanth, V., Bg, S., Ram, J., & Srujana, G. (2017). Estimation of genetic variability, heritability and genetic advance for grain yield components in rice (Oryza sativa L.). Journal of Pharmacognosy and Phytochemistry, 6(4), 1437–1439.
Supriadi, D., Bimantara, Y. M., Zendrato, Y. M., Widaryanto, E., Kuswanto, K., & Waluyo, B. (2024). Assessment of genotype by environment and yield performance of tropical maize hybrids using stability statistics and graphical biplots. PeerJ, 12, e18624. https://doi.org/10.7717/peerj.18624
Supriadi, D., Bimantara, Y. M., Zendrato, Y. M., Widaryanto, E., Kuswanto, K., & Waluyo, B. (2025). Associations and Multi-Traits Selection for Identifying Superior and Stable Maize Hybrids (Zea mays L.) Under Tropical Regions. Caraka Tani: Journal of Sustainable Agriculture, 41(1), 17. https://doi.org/10.20961/carakatani.v41i1.107896
Syukur, M., Sujiprihati, S., & Yunianti, R. (2012). Teknik Pemuliaan Tanaman. Penebar Swadaya Grup.
Tesfaye, D., Abakemal, D., & Habte, E. (2021). Genetic variability, heritability and genetic advance estimation of highland adapted maize (Zea mays L.) genotypes in Ethiopia. Journal of Current Opinion in Crop Science, 2(2), 184–191. https://doi.org/10.62773/jcocs.v2i2.57
Viana, J. M. S. (2023). The impact of epistasis in the heterosis and combining ability analyses. Frontiers in Plant Science, 14(April), 1–11. https://doi.org/10.3389/fpls.2023.1168419
Waghmare, P., Dahat, D., Amolic, V., Shinde, G., Patil, M., Shinde, S., & Dhonde, S. (2025). Combining ability for maize inbreds lines for yield and yield traits. International Journal of Advanced Biochemistry Research, 9(3), 213–223. https://doi.org/10.33545/26174693.2025.v9.i3c.3920
Welderufael, S., Abay, F., Ayana, A., & Amede, T. (2023). Genotype by Trait (GT) and Genotype by Yield*Traits (GYT) Analysis of Sorghum Landraces in Tigray, Northern Ethiopia. Crop Breeding, Genetics and Genomics, 5(2). https://doi.org/10.20900/cbgg20230002
Yan, W., & Frégeau-Reid, J. (2018). Genotype by Yield*Trait (GYT) Biplot: a Novel Approach for Genotype Selection based on Multiple Traits. Scientific Reports, 8(1), 8242. https://doi.org/10.1038/s41598-018-26688-8
Yan, W., Frégeau-Reid, J., Mountain, N., & Kobler, J. (2019). Genotype and Management Evaluation Based on Genotype by Yield*Trait (GYT) Analysis. Crop Breeding, Genetics and Genomics, 1(2). https://doi.org/10.20900/cbgg20190002
YUE, H.-W., HAN, X., WEI, J.-W., ZHENG, S.-H., XIE, J.-L., CHEN, S.-P., PENG, H.-C., & BU, J.-Z. (2023). Comprehensive evaluation of maize hybrids tested in Huang-Huai-Hai summer maize regional trial based on GYT biplot analysis. Acta Agronomica Sinica, 49(5), 1231–1248. https://doi.org/10.3724/SP.J.1006.2023.23035
Zanetta, C. U., Y. Rafii, M., Jaafar, J. N., Warkentin, T. D., Waluyo, B., & Ramlee, S. I. (2025). Variability and assessment of interrelationships among yield and yield‐related characters of pea accessions under the influence of high temperature. New Zealand Journal of Crop and Horticultural Science, 53(4), 870–888. https://doi.org/10.1080/01140671.2023.2180760
Zeyad A. Abdulhamed, Nihad.M. Abood, & Abdulsamad. H. Noaman. (2024). Recurrent selection for general and specific combining ability in maize. IRAQI JOURNAL OF AGRICULTURAL SCIENCES, 55(Special), 99–110. https://doi.org/10.36103/ijas.v55iSpecial.1889
Refbacks
- There are currently no refbacks.

























