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Research Articles

Vol. 13 No. sp5 (2026): Recent Advances in Agriculture

Morphological and biochemical characterisation of seed coat traits in response to mitigate pre-harvest sprouting in soybean (Glycine max (L.) Merr.) genotypes

DOI
https://doi.org/10.14719/pst.15234
Submitted
25 April 2026
Published
28-07-2026

Abstract

Seed coat integrity (porosity, colour and thickness) and structure (hilum, strophiole and micropyle) explain, predict and even allow genotypes resistant to pre-harvest sprouting (PHS) and field weathering under certain environmental conditions. This study examined how morphological and biochemical seed coat characteristics and their role in mitigating PHS in 96 soybean genotypes. All traits showed significant genetic variation, with broad-sense heritability ranging from 0.336 to 0.611. The principal component analysis (PCA) revealed that the first two components explained 62.50 % of total variation. Seed coat proportion correlated strongly with peroxidase activity (POD) (r = 0.511) and negatively with and water absorption capacity (WAC) (r = -0.533). Seeds had black (4 genotypes) and yellow (92 genotypes) seed coats and grey (5 genotypes) and brown (91 genotypes) seed hilums. Physical and biochemical characterisation varied in 96 genotypes, including Birsa Soya 1, Kalitur, Pune 14, Pune 30 (lack colour) and MAUS 61-2 (yellow colour) which were found resistant to field weathering due to hard seed coat (HSC) and seed colour. Black and brown testa genotypes are resistant to field weathering because they have the highest percentage proportion of seed coat, seed thickness (ST) and minimum seed coat permeability than yellow genotypes, which have a delicate seed coat, minimum mechanical strength and maximum seed coat permeability. These findings provide useful selection criteria for breeding soybean cultivars with improved resistance to PHS under humid tropical environments.

References

  1. 1. Archana P, Taware SP, Oak MD, Tamhankar SA, Rao VS. Improvement of oil quality in soybean (Glycine max (L.) Merrill) by mutation breeding. J Am Oil Chem Soc. 2007;84:1117–24. https://doi.org/10.1007/s11746-007-1146-1
  2. 2. Ministry of Agriculture & Farmers Welfare. Final estimates of production of major agricultural crops for 2024–25. Government of India. https://agricoop.gov.in/en/StatEstimates
  3. 3. Yang R, Harrison M, Yang YC, Wang S, Sergey M, Zhao H, et al. Preharvest sprouting in cereals: global incidence, impacts and mitigation strategies. Field Crops Res. 2025;333:110111. https://doi.org/10.1016/j.fcr.2025.110111
  4. 4. Bhatia VS, Bhatnagar PS, Joshi OP. Screening of Indian soybean genotypes for seed longevity as affected by field weathering. Soybean Genetics Newsletter. 1996;23:102–4.
  5. 5. Nautiya PC, Bandyopadhyay A, Zala PV. In situ sprouting and regulation of fresh seed dormancy in Spanish type groundnut (Arachis hypogaea L. ssp. fastigiata var. vulgaris). Field Crops Res. 2001;70:233–41. https://doi.org/10.1016/S0378-4290(01)00143-5
  6. 6. Sohn SI, Pandian S, Kumar TS, Zoclanclounon YAB, Muthuramalingam P, Shilpha J, et al. Seed dormancy and preharvest sprouting in rice: an updated overview. Int J Mol Sci. 2021;22:11804. https://doi.org/10.3390/ijms222111804
  7. 7. Kueneman EA, Dassou S. Screening methodology for resistance to field weathering of soybean seed. Crop Sci. 1982.
  8. 8. Ahmad S, Khulbe RK, Roy D. Evaluation of mungbean (Vigna radiata) germplasm for pre-harvest sprouting tolerance. Legume Res. 2014;37:259–63. https://doi.org/10.5958/j.0976-0571.37.3.039
  9. 9. Alvarez PJC, Krzyzanowski FC, Mandarino JMG, Franca NJB. Relationship between soybean seed coat lignin content and resistance to mechanical damage. Seed Sci Technol. 1997;25:209–14.
  10. 10. Carbonell SAM, Krzyzanowski FC. The pendulum test for screening soybean genotypes for seeds resistant to mechanical damage. Seed Sci Technol. 1995;23:331–9.
  11. 11. Petr S, Vanessa V, Matthew WB, Ales S, Richard DT. The role of the testa during development and in establishment of dormancy of the legume seed. Front Plant Sci. 2014;5:351. https://doi.org/10.3389/fpls.2014.00351
  12. 12. Zhou S, Sekizaki H, Yang Z, Sawa S, Pan J. Phenolics in the seed coat of wild soybean (Glycine soja) and their significance for seed hardness and seed germination. J Agric Food Chem. 2010;58:10972–8. https://doi.org/10.1021/jf102694k
  13. 13. Finch-Savage W, Leubner-Metzger G. Seed dormancy and the control of germination. New Phytol. 2006;171:501–23. https://doi.org/10.1111/j.1469-8137.2006.01787.x
  14. 14. Gao X, Hu CH, Li HZ, Yao YJ, Meng M, Dong J, et al. Factors affecting pre-harvest sprouting resistance in wheat (Triticum aestivum L.): a review. J Anim Plant Sci. 2013;23:556–65.
  15. 15. Sunayana R, Ramendra NS, Raj NSD. Variation in seed dormancy and α-amylase activity in Indian rice (Oryza sativa) accessions. Indian J Agric Sci. 2013;83:56–62.
  16. 16. Singh A, Khulbe RK, Panwar RK. Evaluation of blackgram (Vigna mungo) germplasm for pre-harvesting sprouting tolerance. J Food Legumes. 2012;25:183–6.
  17. 17. Dorian Q, Robin A. Seed dispersal and crop domestication: shattering, germination and seasonality in evolution under cultivation. Annu Plant Rev. 2009;38:238–95.
  18. 18. Humphry ME, Lambrides CJ, Chapman SC, Aitken EAB, Imrie BC, Lawn RJ. Relationships between hard-seededness and seed weight in mungbean assessed by QTL analysis. Plant Breed. 2005;124:292–8.
  19. https://doi.org/10.1111/j.1439-0523.2005.01084.x
  20. 19. Kumar G, Praveen M, Pallavi N, Swapna, Shahana F, Reddy GE, Rakesh G. Genetic variability and correlation studies for pre-harvest sprouting tolerance and associated traits in soybean (Glycine max L. Merrill). Curr J Appl Sci Technol. 2021;40(4):1–10. https:// doi.org/10.9734/cjast/2021/v40i431290
  21. 20. Rani R, Raza G, Ashfaq H, Rizwan M, Shimelis H, Tung MH, Arif M. Analysis of genotype × environment interactions for agronomic traits of soybean (Glycine max (L.) Merr.) using association mapping. Front Genet. 2023;13:1090994. https://doi.org/10.3389/fgene.2022.1090994
  22. 21. Fufa WG, Nepir G. Assessment of genetic variability, heritability and genetic advance in soybean (Glycine max (L.) Merrill) genotypes at Assosa, western Ethiopia. Asian J Biol Sci. 2025;18:83–91. https://doi.org/10.3923/ajbs.2025.83.92
  23. 22. Raju JT, Vasudevan SN, Basavae Gowda SR, Doddagoudar TC, Suma B, Kisan Beladhadi RV. SSR markers linked to hard seediness and electrolyte leakage in soybeans. Pharma Innov. 2022;11:617–20.
  24. 23. Changrong Y, Sripichitt P, Sunanta J, Vipa H, Arom S. Modifying controlled deterioration for evaluating field weathering resistance of soybean. Kasetsart J Nat Sci. 2007;41(2).
  25. 24. Alexander HM, Cummings CL, Khan L, Snow A. Seed size variation and predation of seeds produced by wild and crop-wild sunflowers. Ann Bot. 2001;88:623–7. https://doi.org/10.2307/2657061
  26. 25. Egli DB, TeKrony DM, Wiralaga RA. Effect of soybean seed vigor and size on seedling growth. J Seed Tech. 1990;14(1):1–12. http://www.jstor.org/stable/23432656
  27. 26. Ragus LN. Variation in water absorbing capacity and other seed characters of soybean (Glycine max (L.) Merr.). Philipp J Crop Sci. 1986;11:195–201.
  28. 27. Presley JT. Relation of protoplast permeability to cotton seed viability and predisposition to seedling disease. Plant Dis Rep. 1958;42:852. https://www.cabidigitallibrary.org/doi/full/10.5555/19581102834
  29. 28. Castillo MDP, Stenstrom J, Ander P. Determination of manganese peroxidase activity with 3-methyl-2-benzothiazolinone and 3-(dimethylamino) benzoic acid. Ann Chem Sci Res. 1994;218:399–404. https://doi.org/10.1006/abio.1994.1198
  30. 29. Nnenna EO, Kalu CM, Nnorom C. Estimation of protein content and amino acid compositions in selected plant samples using UV-Vis spectrophotometric method. Am J Food Sci Health. 2016;3:41–6.
  31. 30. MA F. Cracks in the palisade cuticle of soybean seed coats correlate with their permeability to water. Ann Bot. 2004;94:213–28. https://doi.org/10.1093/aob/mch133
  32. 31. Sun Y, Gong Y. Research advances on the hard seededness trait of soybean and the underlying regulatory mechanisms. Front Plant Sci. 2024;15:1419962. https://doi.org/10.3389/fpls.2024.1419962
  33. 32. Zhang L, Jia R, Liu L, Shen W, Fang Z, Zhou B, Liu B. Seed coat colour and structure are related to the seed dormancy and overwintering ability of crop-to-wild hybrid soybean. Ann Bot Plants. 2023;15:eplad081. https://doi.org/10.1093/aobpla/plad081
  34. 33. Bewley JD, Bradford KJ, Hilhorst HWM, Nonogaki H. Seeds: Physiology of Development, Germination and Dormancy. 3rd ed. Springer; 2013. https://doi.org/10.1007/978-1-4614-4693-4
  35. 34. Salinas R, Roque MC, Celina B, Vilma B, Ana MY. Electrical conductivity of soybean seed cultivars and adjusted models of leakage curves along the time. Rev Caatinga. 2010;23:1–7.
  36. 35. Cavalcante JA, Gadotti GI, Pinheiro RM, Silva RNO, de Oliveira FK, de Moraes DM. Vigor and anaerobic metabolism of soybean seeds evaluated by ethanol test. J Seed Sci. 2023;45:e202345007. https://doi.org/10.1590/2317-1545v45263825
  37. 36. Gillikin JW, Graham JS. Purification and developmental analysis of the major anionic peroxidase from the seed coat of Glycine max. Plant Physiol. 1991;96:214–20. https://doi.org/10.1104/pp.96.1.214
  38. 37. Capeleti I, Bonini EA, Ferrarese MDLL. Lignin content and peroxidase activity in soybean seed coat susceptible and resistant to mechanical damage. Acta Physiol Plant. 2005;27:103–8. https://doi.org/10.1007/s11738-005-0042-2
  39. 38. Ranathunge K, Shao S, Qutob D. Properties of the soybean seed coat cuticle change during development. Planta. 2010;231:1171–88. https://doi.org/10.1007/s00425-010-1118-9
  40. 39. Guo B, Sun L, Jiang S, Ren H, Sun R, Wei Z, et al. Soybean genetic resources contributing to sustainable protein production. Theor Appl Genet. 2022;135:4095–121. https://doi.org/10.1007/s00122-022-04222-9
  41. 40. Kholmurodova G, Gulchekhra T, Murod R, Rano Y. Analysis of protein and oil content in seeds of soybean collection varieties. E3S Web Conf. 2022;377:03016. https://doi.org/10.1051/e3sconf/202337703016
  42. 41. Baroniya SS, Jumrani K, Baroniya M, Guruprasad KN, Landi M, Kataria S. Intraspecific variation in photosynthetic efficiency in soybean (Glycine max L.) varieties towards solar ultraviolet radiations. Photosynthetica. 2022;61:203–14. https://doi.org/10.32615/ps.2022.048
  43. 42. Mishra N, Jiang C, Chen L, Paul A, Chatterjee A, Shen G. Achieving abiotic stress tolerance in plants through antioxidative defense mechanisms. Front Plant Sci. 2023;14:1110622. https://doi.org/10.3389/fpls.2023.1110622
  44. 43. Wilson RF. Seed composition. In: Boerma HR, Specht JE, editors. Soybeans: Improvement, Production and Uses. 3rd ed. ASA, CSSA and SSSA; 2004. p. 621–68. https://doi.org/10.2134/agronmonogr16.3ed.c13
  45. 44. Panthee DR, Pantalone VR, West DR, Saxton AM, Sams CE. Quantitative trait loci for seed protein and oil concentration and seed size in soybean. Crop Sci. 2005;45:2015–22. https://doi.org/10.2135/cropsci2004.0720
  46. 45. Rao PS, Madhulety TY, Ankaiah R, Volet SR. Morpho-physiological trait variation of pre-harvest sprouting tolerance to simulated rain in mungbean (Vigna radiata). Indian J Agric Sci. 2023;93:269–73. https://doi.org/10.56093/ijas.v93i3.128735
  47. 46. Gupta S, Aski M, Mishra GP, Yadav PS, Tripathi K, Lal SK, et al. Genetic variation for tolerance to pre-harvest sprouting in mungbean (Vigna radiata) genotypes. PeerJ. 2024;12:e17609. https://doi.org/10.7717/peerj.17609
  48. 47. Pinheiro DT, dos Santos Dias DCF, Silva JM, Gama GFV, León MJZ, da Silva LJ. Assessment of the tolerance of soybean seeds to weathering deterioration in the pre-harvest phase by multivariate analysis. J Seed Sci. 2022;44:e202244035. https://doi.org/10.1590/2317-1545v44263001
  49. 48. Abati J, Zucareli C, Brzezinski C, Lopes I, Krzyzanowski F, Cardoso M, Henning F. Water absorption and storage tolerance of soybean seeds with contrasting seed coat characteristics. Acta Sci Agron. 2022;44:e53096. https://doi.org/10.4025/actasciagron.v44i1.53096
  50. 49. Yunuskhanov MSh, Jaynakov ZL, Abdurazakova. Soybean seed peroxidase. J Propulsion Technol. 2023;44:4. https://doi.org/10.52783/tjjpt.v44.i4.969
  51. 50. Soppe WJJ, Bentsink L. Seed dormancy back on track; its definition and regulation by DOG1. New Phytol. 2020;228:816–9. https://doi.org/10.1111/nph.16592

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