Morphological diversity, productivity, and grain quality of wheat morphotypes obtained by Triticum aestivum × Triticum spelta hybridization
Abstract
Spike shape and morphology in wheat are key phenotypic traits determining the potential grain set per spike and influencing yield. Hybridization between common wheat and spelt generated a wide range of new forms differing in morphobiological traits, particularly in spike architecture. Determining productivity indicators of different wheat morphotypes based on spike shape remains an important approach for identifying genotypes that combine desirable morphological characteristics with high agronomic performance. In this context, the study of variation among morphotypes contributes to a better understanding of the relationship between spike architecture and yield formation. In this study, wheat samples were classified into distinct morphotypes according to spike morphology, and their productivity and grain quality parameters were comprehensively analyzed. Comparative evaluation of morphotypes revealed significant differences in yield components, grain weight per spike, protein content, and gluten content, allowing for the assessment of their breeding value and potential use in selection programs. Based on spike morphology, the obtained genetic diversity was classified into six morphotypes: spelts, speltoids, forms with a typical common wheat spike, squareheads, subcompactoids, and compactoids. It was established that common wheat and squareheads significantly outperformed other morphotypes: T heir yield was on average 42–51% higher than that of speltoids and spelts and about 67% higher than that of subcompactoids and compactoids. A similar trend was observed for grain weight per spike, common wheat and squareheads outperformed speltoids by ~14%, spelts by ~19%, and subcompactoids and compactoids by ~34%. By contrast, the ranking by grain protein and gluten content was reversed: S pelts exhibited the highest values (around 19–20% protein and 42–44% gluten), followed by speltoids (about 16–17% and 35–38%), while common wheat and squareheads occupied an intermediate position (approximately 14.5–15.5% and 32–34%), and compactoids and subcompactoids showed the lowest levels (about 14–15% and 31–33%). The analysis of morphological diversity, productivity, and grain quality in wheat morphotypes derived from Triticum aestivum × Triticum spelta hybridization is important for breeding and genetic studies, enabling assessment of breeding value and identification of genotypes with optimal spike architecture and stable expression of economically valuable traits. This approach provides a basis for differentiating morphotypes according to yield components and grain quality parameters, including protein and gluten content. The revealed variability among morphotypes reflects the effectiveness of interspecific hybridization in generating genetically diverse material for further sele c tion.References
Curzon, A. Y., Kottakota, C., Nashef, K., Abbo, S., Bonfl, D. J., Reifen, R., Bar-El, S., Rabinovich, O., Avneri, A., & Ben-David, R. (2021). Assessing adaptive requirements and breeding potential of spelt under Mediterranean environment. Scientifc Reports, 11, 7208.
Diordiieva, I. P., Riabovol, I. S., Riabovol, L. O., Babii, M. M., Fedorenko, S. V., Serzhuk, O. P., Maslovata, S. A., Liubchenko, A. I., Novak, Z. M., & Liubchenko, I. O. (2024). Breeding and genetic improvement of spelt wheat (Triticum spelta L.) by interspecific hybridization. Regulatory Mechanisms in Biosystems, 15(3), 463–469.
Diordiieva, I. P., Riabovol, L. O., Riabovol, Y. S., Serzhuk, O. P., Nakloka, I. І., Nakloka, О. P., & Karychkovska, S. P. (2022). Breeding and genetic improvement of soft winter wheat with the use of spelt wheat. Agronomy Research, 20(1), 91–102.
Frantová, N., Rábek, M., Elzner, P., Středa, T., Jovanović, I., Holková, L., Martinek, P., Smutná, P., & Prášil, I. T. (2022). Different drought tolerance strategy of wheat varieties in spike architecture. Agronomy, 12(10), 2328.
Guo, Z., Zhao, Y., Röder, M. S., Reif, J. C., Ganal, M. W., Chen, D., & Schnurbusch, T. (2018). Manipulation and prediction of spike morphology traits for the improvement of grain yield in wheat. Scientific Reports, 8(1), 14435.
Halder, J., Gill, H. S., Zhang, J., Altameemi, R., Olson, E., Turnipseed, B., & Sehgal, S. K. (2023). Genome wide association analysis of spike and kernel traits in the U.S. hard winter wheat. The Plant Genome, 16(1), e20300.
Hospodarenko, H. M., Kostogriz, P. V., Liubich, V. V., Parіi, M. F., Poltoretskii, S. P., Polianetska, I. O., Riabovol, I. S., Rjabovol, L. O., & Suhomud, O. G. (2016). Pshenytsia spelta [Spelt wheat]. Sіk Group Ukraine, Kyiv (in Ukrainian).
Ji, Z., Liu, X., Yan, F., Wu, S., & Du, Y. (2025). The genetic basis of wheat spike architecture. Agriculture, 15(15), 1575.
Johnson, E. R., Nalam, V. J., Zemetra, R. S., & Riera-Lizarazu, O. (2008). Mapping the compactum locus in wheat (Triticum aestivum L.) and its relationship to other spike morphology genes of the Triticeae. Euphytica, 163, 193–201.
Konopatskaia, I., Vavilova, V., Blinov, A., Goncharov, N. P. (2017). Spike morphology genes in wheat species (Triticum L.). Proceedings of the Latvian Academy of Sciences, Section B, Natural, Exact, and Applied Sciences, 70(6), 345–355.
Liu, H., Shi, Z., Ma, F., Xu, Y., Han, G., Zhang, J., Liu, D., An, D. (2022). Identification and validation of plant height, spike length and spike compactness loci in common wheat (Triticum aestivum L.). BMC Plant Biology, 22, 568.
Liu, M., Zhao, Q., Qi, F., Stiller, J., Tang, S., Miao, J., Vrána, J., Holušová, K., Liu, D., Doležel, J., Manners, J. M., Han, B., & Liu, C. (2018). Sequence divergence between spelt and common wheat. Theoretical and Applied Genetics, 131(5), 1125–1132.
Liu, Y., Yu, R., Shen, L., Sun, M., & Peng, Y. (2024). Genomic insights into the modifications of spike morphology traits during wheat breeding. Plant, Cell and Environment, 47(12), 5470–5482.
Luo, X., Yang, Y., Lin, X., & Xiao, J. (2023). Deciphering spike architecture formation towards yield improvement in wheat. Journal of Genetics and Genomics, 50(11), 835–845.
Shen, K., Ye, B., Yu, X., Shen, P., Yu, R., Yin, C., & He, Z. (2025). Dissection of genomic drivers of spike morphology changes in wheat by high throughput phenotyping. Cell Reports, 44(8), 116120.
Simons, K. J., Fellers, J. P., Trick, H. N., Zhang, Z., Tai, Y. S., Gill, B. S., & Faris, J. D. (2006). Molecular characterization of the major wheat domestication gene Q. Genetics, 172(1), 547–555.
Sormacheva, I., Golovnina, K., Vavilova, V., Kosuge, K., Watanabe, N., Blinov, A., & Goncharov, N. P. (2015). Q gene variability in wheat species with different spike morphology. Genetic Resources and Crop Evolution, 62, 837–852.
Sun, F., Zheng, S., Li, Z., Gao, Q., & Jiang, N. (2025). Analysis of wheat spike morphological traits by 2D imaging. Plant Phenomics, 7(3), 100096.
Sіchkar, S. M., Morgun, V. V., & Dubrovna, O. V. (2016.) Uspadkuvannia morfolohichnykh oznak hibrydiv F1–F2 T. spelta × T. aestivum [Inheritance of morphological traits of F1–F2 T. spelta × T. aestivum hybrids]. Physiology of Plants and Genetics, 48(4), 344–355 (in Ukrainian).
Takač, V., Tóth, V., Rakszegi, M., Mikić, S., Mirosavljević, M., & Kondić Špika, A. (2021). Differences in processing quality traits, protein content and composition between spelt and bread wheat genotypes grown under conventional and organic production. Foods, 10(1), 156.
Wang, M., Lu, J., Liu, R., Li, Y., Ao, D., Wu, Y., & Zhang, L. (2023) Identification and validation of a major quantitative trait locus for spike length and compactness in the wheat (Triticum aestivum L.) line Chuanyu12D7. Frontiers in Plant Science, 14, 1186183.
Wiwart, M., Szafrańska, A., & Suchowilska, E. (2023). Grain of hybrids between spelt (Triticum spelta L.) and bread wheat (Triticum aestivum L.) as a new raw material for breadmaking. Polish Journal of Food and Nutrition Sciences, 73(3), 265–277.
Wolde, G. M., & Schnurbusch, T. (2019). Inferring vascular architecture of the wheat spikelet based on resource allocation in the branched headt (bht-A1) near isogenic lines. Functional Plant Biology, 46(11), 1023–1035.
Xie, Q., Li, N., Yang, Y., Lv,Y., Yao, H., Wei, R., Sparkes, D. L., & Ma, Z. (2018). Pleiotropic effects of the wheat domestication gene Q on yield and grain morphology. Planta, 247(5), 1089–1098.
Zhai, H., Feng, Z., Li, J., Liu, X., Xiao, S., Ni, Z., & Sun, Q. (2016). QTL analysis of spike morphological traits and plant height in winter wheat (Triticum aestivum L.) using a high-density SNP and SSR-based linkage map. Frontiers in Plant Science, 7, 1617.
Zhou, H., Riche, A. B., Hawkesford, M. J., Whalley, W. R., Atkinson, B. S., Sturrock, C. J., & Mooney, S. J. (2021). Determination of wheat spike and spikelet architecture and grain traits using X ray Computed Tomography imaging. Plant Methods, 17, 26.
Żuk-Gołaszewska, K., Majewska, K., Gołaszewski, J., & Tyburski, J. (2022). Nutritional properties of organic spelt wheats in different growth stages and the resulting flours. Journal of Elementology, 27(3), 645–662.
Authors retain copyright and grant the journal right of first publication with the work simultaneously licensed under a Creative Commons «Attribution» 4.0 License that allows others to share the work with an acknowledgement of the work's authorship and initial publication in this journal.


