Diversity across Ploidy Levels in Mulberry (Morus spp.): A Review of Genetic Resources and Their Utilization

Authors: Y Surjalata Devi; Anowar Hossain
Diversity across Ploidy Levels in Mulberry (Morus spp.): A Review of Genetic Resources and Their Utilization
DIN
IJOEAR-MAR-2026-38
Abstract

Mulberry (Morus spp.) is an economically important perennial crop that serves as the primary food source for the domesticated silkworm (Bombyx mori) and also has potential for fruit production and the utilization of bioactive compounds. This genus exhibits considerable diversity in species characteristics, geographical distribution, chromosome number, and ploidy levels, providing valuable genetic resources for crop improvement. This review surveys mulberry species diversity, chromosome number and genome size variation, and the utilization of genetic resources in the development of improved cultivars. It further summarizes varieties developed through selection, hybridization, mutation breeding, and the use of polyploid germplasm, with emphasis on leaf productivity, environmental adaptation, and suitability for different silkworm-rearing requirements. In addition, reported genes associated with responses to drought, salinity, cold, heat, and powdery mildew are discussed in relation to their potential applications in stress-tolerance improvement. Despite this diversity, mulberry breeding faces challenges associated with variable ploidy levels, high heterozygosity, and long breeding cycles. Integrating conventional breeding with genomics, tissue culture, and emerging genome-editing technologies may help overcome these constraints and facilitate the development of productive, stress-tolerant, and multipurpose cultivars. Overall, this review highlights the importance of conserving and effectively utilizing mulberry genetic resources to support sustainable sericulture and broader agricultural and industrial applications.

Keywords
Morus genetic diversity ploidy mulberry stress tolerance sericulture.
Introduction

Mulberry (Morus spp.) is an ancient, perennial, deciduous plant belonging to the family Moraceae. The genus is widely distributed across Asia and other regions, with considerable diversity among its wild and cultivated forms. East Asia and the Himalayan region are important areas of mulberry diversity. Several species, including Morus indica, M. alba (white mulberry), M. serrata (Himalayan mulberry), and M. laevigata, occur in different geographical regions of India [17], [24]. Mulberry is cultivated primarily for its leaves, which provide the principal food source for the domesticated silkworm (Bombyx mori). During its larval development, the silkworm feeds on mulberry leaves and subsequently spins a cocoon from which raw silk is extracted. Silk production and quality are influenced by leaf quality and the silkworm strain used, including multivoltine, bivoltine, and their hybrids [8], [9]. Beyond sericulture, mulberry fruits are valued for their nutritional and phytochemical constituents, including anthocyanins, antioxidants, fatty acids, and vitamins [11]. Mulberry leaves are also used in health beverages and animal feed because they contain bioactive compounds such as flavonoids, phenolic compounds, and coumarins, which have been investigated for potential health benefits [6], [22].

The genus Morus exhibits considerable variation in leaf morphology, growth characteristics, reproductive biology, fruit traits, and environmental adaptability [30]. This diversity provides valuable opportunities to identify desirable traits and broaden the genetic base of cultivated mulberry. However, the breeding potential of individual species depends on their agronomic characteristics, environmental adaptation, and reproductive compatibility with existing breeding materials. Systematic characterization and evaluation of mulberry genetic resources are therefore essential for identifying suitable material for specific improvement objectives.

Advances in molecular biology and genomics have created further opportunities for mulberry improvement. Research has identified genes associated with responses to abiotic stresses, including drought, salinity, cold, and heat, as well as biotic stresses such as powdery mildew. Understanding the functions of these genes may facilitate the identification and validation of genetic targets for developing stress-tolerant cultivars [19]. Integrating this knowledge with conventional breeding and emerging biotechnological approaches may help address challenges associated with mulberry productivity, environmental adaptation, and disease resistance.

In this context, the present review examines the genetic diversity of Morus species, focusing on geographical distribution, chromosome number, ploidy variation, and genome size. It also summarizes improved mulberry varieties and their utilization, discusses reported stress-responsive genes, and considers the major challenges and future opportunities for integrating conventional breeding with modern biotechnological approaches. The overall aim is to highlight how the conservation and effective utilization of mulberry genetic resources can contribute to the development of productive, stress-tolerant, and multipurpose cultivars for sustainable sericulture and broader agricultural and industrial applications.

Conclusion

Mulberry (Morus spp.) exhibits considerable diversity in chromosome number, ploidy level, genetic characteristics, and potential utilization, providing valuable resources for sericulture, fruit production, and bioactive-compound development. Improved varieties demonstrate how this diversity can be utilized to enhance leaf productivity, meet the feeding requirements of silkworms at different developmental stages, and adapt cultivation to diverse environments. In addition, studies of stress-responsive genes have identified potential molecular targets for improving tolerance to abiotic stresses and diseases. Nevertheless, challenges associated with ploidy variation, high heterozygosity, reproductive biology, and long breeding cycles remain important constraints. Integrating conventional breeding with genomics, tissue culture, and emerging genome-editing technologies may help improve the identification and utilization of valuable genetic resources. Such an integrated approach could contribute to the development of productive, resilient, and multipurpose mulberry cultivars for sustainable sericulture and broader agricultural and industrial applications.

Agriculture Journal IJOEAR Call for Papers

References
  1. Arora, V., Ghosh, M. K., Pal, S., & Gangopadhyay, G. (2017). Allele specific CAPS marker development and characterization of chalcone synthase gene in Indian mulberry (Morus spp., family Moraceae). PLoS One, *12*(6), e0179189.
  2. Bian, C., Long, D., Liu, X., Wei, C., Gong, J., & Zhao, A. (2015). Cloning and expression to salt stress of Na⁺/H⁺ antiporter gene (MnNHX1) in mulberry tree. Scientia Silvae Sinicae, *51*(8), 16-25.
  3. Central Sericultural Research and Training Institute. (2012). Mulberry varieties suitable for eastern and north eastern India [Brochure]. Central Silk Board, Ministry of Textiles, Government of India. https://www.csrtiber.res.in/Brochure_pamphlets/Mul_ver_E%26NE_India.pdf
  4. Central Sericultural Research and Training Institute. (n.d.). Improved mulberry varieties. Central Silk Board, Ministry of Textiles, Government of India. https://www.csrtiber.res.in/Brochure_pamphlets/Brochure.htm
  5. Central Silk Board. (2020). Note on sericulture: Research and development highlights, 2020–21. Ministry of Textiles, Government of India. https://www.csb.gov.in/sites/default/files/what_s_new/files/Note-on-Seri-CSB-2020-21.pdf
  6. Chae, J.-Y., Lee, J.-Y., Hoang, I.-S., Wang-Bo, D., Choi, P.-W., Lee, W.-C., Kim, J.-W., Kim, S.-Y., Choi, S.-W., & Rhee, S.-J. (2003). Analysis of functional components of leaves of different mulberry cultivars. Journal of the Korean Society of Food Science and Nutrition, *32*(1), 15-21.
  7. Debaraj, Y., Singh, R., & Bajpai, A. K. (2010). Evaluation of some mulberry varieties for yield and quality through bioassay in Manipur, North East India. Uttar Pradesh Journal of Zoology, *30*(2), 165-168. https://mbimph.com/index.php/UPJOZ/article/view/276
  8. Devi, Y. S., Hossain, A., Lalremruatpuia, V., & Raviraj, V. S. (2024). Effect of UV radiation on physiological performance of silkworm Bombyx mori. Biological Forum, *16*(6), 221-229.
  9. Devi, Y. S., Verma, A. K., Nalavadi, C., Hossain, A., & Pramanick, B. (2024). Optimizing preservation duration of mulberry silkworm crossbreed eggs in cold storage. Biological Forum, *16*(8), 249-356.
  10. Erarslan, Z. B., Karagöz, S., & Kültür, Ş. (2021). Comparative morphological and anatomical studies on Morus species (Moraceae) in Turkey. Turkish Journal of Pharmaceutical Sciences, *18*(2), 157.
  11. Kattil, A., Dash, K. K., Shams, R., & Sharma, S. (2024). Nutritional composition, phytochemical extraction, and pharmacological potential of mulberry: A comprehensive review. Future Foods, *9*, 100295.
  12. Li, R., Su, X., Zhou, R., Zhang, Y., & Wang, T. (2022). Molecular mechanism of mulberry response to drought stress revealed by complementary transcriptomic and iTRAQ analyses. BMC Plant Biology, *22*(1), 36.
  13. Liu, C., Hu, J., Fan, W., Zhu, P., Cao, B., Zheng, S., & Zhao, A. (2021). Heterotrimeric G-protein γ subunits regulate ABA signaling in response to drought through interacting with PP2Cs and SnRK2s in mulberry (Morus alba L.). Plant Physiology and Biochemistry, *161*, 210-221.
  14. Ma, G., Chai, X., Hou, G., Zhao, F., & Meng, Q. (2022). Phytochemistry, bioactivities and future prospects of mulberry leaves: A review. Food Chemistry, *372*, 131335.
  15. Rajan, M. V., Dandin, S. B., Magadum, S. B., & Datta, R. K. (2000). Nutritional evaluation of mulberry (Morus spp.) genotypes through silkworm growth studies.
  16. Ramesha, A., Dubey, H., Vijayan, K., Ponnuvel, K. M., Mishra, R. K., & Suresh, K. (2020). Genome wide characterization revealed MnMLO2 and MnMLO6A as candidate genes involved in powdery mildew susceptibility in mulberry. Molecular Biology Reports, *47*(4), 2889-2900.
  17. Rao, A. A., Thangavelu, K., & Sharma, K. R. (2005). Distribution and variation of mulberry genetic resources (Morus spp.) in the high altitude and cold deserts of Ladakh Himalayan region. Indian Journal of Plant Genetic Resources, *18*(2), 255-259.
  18. Sarkar, K. (2020). Studies on the effect of different types of feeding on the commercial characters of mulberry silkworm (Bombyx mori L.) in West Bengal: A review. International Journal of Agriculture, Environment and Biotechnology, *13*(3), 305-321.
  19. Sarkar, T., Mogili, T., & Sivaprasad, V. (2017). Improvement of abiotic stress adaptive traits in mulberry (Morus spp.): An update on biotechnological interventions. 3 Biotech, *7*(3), 214.
  20. Sliwinska, E., Loureiro, J., Leitch, I. J., Šmarda, P., Bainard, J., Bureš, P., & Galbraith, D. W. (2022). Application-based guidelines for best practices in plant flow cytometry. Cytometry Part A, *101*(9), 749-781.
  21. Song, F., Li, Y., Huang, J., Lu, W., Guo, Z., & Deng, W. (2024). Integrated transcriptomic and proteomic analyses reveal the impact of drought and heat stress combination on Morus alba. Environmental and Experimental Botany, *228*, 105988.
  22. Song, W., Wang, H. J., Bucheli, P., Zhang, P. F., Wei, D. Z., & Lu, Y. H. (2009). Phytochemical profiles of different mulberry (Morus sp.) species from China. Journal of Agricultural and Food Chemistry, *57*(19), 9133-9140.
  23. Tikader, A., & Dandin, S. B. (2008). Foliar morphology and venation pattern in different mulberry (Morus spp.).
  24. Tikader, A., & Roy, B. N. (2002). Genetic divergence in mulberry (Morus spp.). The Indian Journal of Genetics and Plant Breeding, *62*(1), 52-54.
  25. Wang, H., Liu, J., Xu, X., Li, Y., Yuan, J., Zeng, Y., & Ma, B. (2024). MGCPdb, a collective resource for mulberry genome size, chromosome number, and ploidy. Forestry Research, *4*, e027.
  26. Wang, M., Zhu, M., Qian, J., Yang, Z., Shang, F., Egan, A. N., & Liu, L. (2024). Phylogenomics of mulberries (Morus, Moraceae) inferred from plastomes and single copy nuclear genes. Molecular Phylogenetics and Evolution, *197*, 108093.
  27. Xia, Z., Dai, X., Fan, W., Liu, C., Zhang, M., Bian, P., & Zhao, A. (2022). Chromosome-level genomes reveal the genetic basis of descending dysploidy and sex determination in Morus plants. Genomics, Proteomics & Bioinformatics, *20*(6), 1119-1137.
  28. Xuan, Y., Ma, B., Li, D., Tian, Y., Zeng, Q., & He, N. (2022). Chromosome restructuring and number change during the evolution of Morus notabilis and Morus alba. Horticulture Research, *9*, uhab030.
  29. Xuan, Y., Wang, S., Li, S., Yuan, J., Zhou, Q., & He, N. (2023). Chromosome constitution and genetic relationships of Morus spp. revealed by genomic in situ hybridization. BMC Plant Biology, *23*(1), 428.
  30. Yang, C. X., Liu, S. Y., Zerega, N. J., Stull, G. W., Gardner, E. M., Tian, Q., & Yi, T. S. (2023). Phylogeny and biogeography of Morus (Moraceae). Agronomy, *13*(8), 2021.
  31. Zeng, Q., Chen, M., Wang, S., Xu, X., Li, T., Xiang, Z., & He, N. (2022). Comparative and phylogenetic analyses of the chloroplast genome reveal the taxonomy of the Morus genus. Frontiers in Plant Science, *13*, 1047592.
  32. Zhao, M., Zhou, G., Liu, P., Wang, Z., Yang, L., Li, T., & Lin, T. (2024). The role of MaFAD2 gene in bud dormancy and cold resistance in mulberry trees (Morus alba L.). International Journal of Molecular Sciences, *25*(24), 13341.
Article Preview