The effect of biostimulants and light wavelengths on the physiology of Cleome gynandra seeds

  • Nkhanedzeni K. Nemahunguni
  • Shubhpriya Gupta
  • Manoj G. Kulkarni
  • Jeffrey F. Finnie
  • Johannes Van StadenEmail author
Original paper


Cleome gynandra L. is used as a vegetable that forms a significant part of the local diet in South Africa and other tropical and subtropical parts of the world. Cleome gynandra seeds are negatively photoblastic (light-induced dormancy) and fail to germinate when planted immediately after harvest. To enhance better cultivation practices, seed germination of C. gynandra using different light wavelengths (red, far-red, green and blue light) with and without organic biostimulants [smoke–water (SW), karrikinolide (KAR1), Kelpak® (KEL) and eckol (ECK)] was tested. Among all the tested biostimulants, the best germination percentage (40%) was observed in seeds treated with SW in the dark. However, the biostimulants did not show any significant effect with the light treatments. In this study, blue light generally promoted (≤ 35%) and red light inhibited (≤ 8%) germination. Furthermore, the effect of different biostimulants (under blue and red light) on biochemical content and enzyme activities was tested in seeds of C. gynandra. Seeds treated with biostimulants in blue light showed an overall increase in protein and total carbohydrate content in comparison to seeds subjected to biostimulants in red light. The α-amylase activity in the seeds was highest in KEL-treated seeds in blue light. Superoxide dismutase and catalase activity was generally higher in blue-light treatment, while peroxidase activity was highest in red light. Enhanced germination under blue light and inhibitory effects with red light is an intriguing phenomenon for C. gynandra seeds, which needs more detailed investigation. The study also indicates the potential application of organic biostimulants (particularly SW) for better seed germination and growth of C. gynandra which can be explored by farmers in the field.


Biostimulants Germination Indigenous crop 



Analysis of variance






Gibberellic acid








Superoxidase dismutase





The authors thank the University of KwaZulu-Natal and the National Research Foundation (UID: 98896), Pretoria for financial support.


The authors thank the University of KwaZulu-Natal and the National Research Foundation (UID: 98896), Pretoria for financial support.


  1. Bandyopadhyay U, Das D, Banerjee RK (1999) Reactive oxygen species: oxidative damage and pathogenesis. Curr Sci 77:658–666Google Scholar
  2. Baskin CC, Baskin JM (1998) Seeds: ecology, biogeography, and evolution of dormancy and germination. Academic Press, San DiegoGoogle Scholar
  3. Baxter B, Van Staden J, Granger J, Brown NAC (1994) Plant-derived smoke and smoke extracts stimulate seed germination of the fire-climax grass Themeda triandra Forssk. Environ Exper Bot 34:217–223CrossRefGoogle Scholar
  4. Beers RF, Sizer IW (1952) A spectrophotometric method for measuring the breakdown of hydrogen peroxide by catalase. J Biol Chem 195:133–140PubMedPubMedCentralGoogle Scholar
  5. Bewley ID, Black M (1994) Seeds: physiology of development and germination, 2nd edn. Plenum Press, New YorkCrossRefGoogle Scholar
  6. Bian ZH, Yang QC, Liu WK (2015) Effects of light quality on the accumulation of phytochemicals in vegetables produced in controlled environments: a review. J Sci Food Agric 95:869–877PubMedCrossRefPubMedCentralGoogle Scholar
  7. Borthwick HA, Hendricks SB, Toole EH, Toole VK (1954) Action of light on lettuce seed germination. Bot Gaz 115:205–225CrossRefGoogle Scholar
  8. Bradford MM (1976) A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding. Anal Biochem 72:248–254PubMedPubMedCentralCrossRefGoogle Scholar
  9. Burki HM, Schroeder D, Lawrie J, Cagan L, Vrablova M, El Aydam M, Szentkiralyi F, Ghorbani R, Juttersonke B, Ammon HU (1997) Biological control of pigweeds (Amaranthus retroflexus, L. A. Powellii, S. Watson and A. bouchonii Thell.) with phytophagous insects, fungal pathogens and crop management. Integrated Pest Manag Rev 2:51–59CrossRefGoogle Scholar
  10. Chander R, Kapoor NK (1990) High density lipoprotein is a scavenger of superoxide anions. Biochem Pharmacol 40:1663–1665PubMedCrossRefPubMedCentralGoogle Scholar
  11. Chweya J, Eyzaguirre P (1999) The biodiversity of traditional leafy vegetables. International Plant Genetic Resources Institute, Italy, p 182Google Scholar
  12. Chweya J, Mnzava NA (1997) Cat’s whiskers (Cleome gynandra L): promoting the conservation and use of underutilized and neglected crops. 11. International Plant Genetic Resources Institute, Rome, p 54Google Scholar
  13. Cone JW, Jaspers PAPM, Kendrick RE (1985) Biphasic fluence-response curves for light induced germination of Arabidopsis thaliana seeds. Plant, Cell Environ 8:60–612CrossRefGoogle Scholar
  14. Dong C, Fu Y, Liu G, Liu H (2014) Growth photosynthetic characteristics, antioxidant capacity and biomass yield and quality of wheat (Triticum aestivum L.) exposed to LED light sources with different spectra combinations. J Agron Crop Sci 200:219–230CrossRefGoogle Scholar
  15. Flematti GR, Ghisalberti EL, Dixon KW, Trengove RD (2005) Synthesis of the seed germination stimulant 3-methyl-2H-furo[2,3-c]pyran-2-one. Tetrahedron Lett 46:5719–5721CrossRefGoogle Scholar
  16. Giannopolitis CN, Ries SK (1977) Superoxide dismutases I. Occurrence in higher plants. Plant Physiol 59:309–314PubMedPubMedCentralCrossRefGoogle Scholar
  17. Ginzo HD (1978) Red and far red inhibition of germination in Aristida murina Cav. Z Pflanzenphysiol 90:303–307CrossRefGoogle Scholar
  18. Gupta S, Hrdlička J, Ngoroyemoto N, Nemahunguni NK, Gucký T, Novák O, Kulkarni MG, Doležal K, Van Staden J (2019) Preparation and standardisation of smoke–water for seed germination and plant growth stimulation. J Plant Growth Regul. CrossRefGoogle Scholar
  19. Hilton JR (1982) An unusual effect of the far-red absorbing form of phytochrome: photoinhibition of seed germination in Bromus sterilis L. Planta 155:524–528PubMedCrossRefPubMedCentralGoogle Scholar
  20. Hilton JR (1984) The influence of temperature and moisture status on the photoinhibition of seed germination in Bromus sterilis L. by the far-red absorbing form of phytochrome. New Phytol 97:369–374CrossRefGoogle Scholar
  21. Hrdlička J, Gucký T, Novák O, Kulkarni MG, Gupta S, Van Staden J, Doležal K (2019) Quantification of karrikins in smoke water using ultra-high performance liquid chromatography–tandem mass spectrometry. Plant Methods 15:81PubMedPubMedCentralCrossRefGoogle Scholar
  22. Kaul HP, Aufhammer W, Laible B, Nalborczyk E, Pirog S, Wasiak K (1996) The suitability of amaranth genotypes for grain and fodder use in Central Europe. Die Bodenkultur 47:173–181Google Scholar
  23. Kendrick RE, Kronenberg GHM (1994) Photomorphogenesis in plants, 2nd edn. Kluwer Academic Publishers, DordrechtCrossRefGoogle Scholar
  24. Kokwaro JO (1976) Medicinal plants of East Africa. East African Literature Bureau, Kampala, Nairobi, Dar-Es-Salaam. McGraw-Hill Book Co, New York, pp 106–107Google Scholar
  25. Kulkarni MG, Sparg SG, Van Staden J (2006) Dark conditioning, cold stratification and a smoke-derived compound enhance the germination of Eucomis autumnalis subsp. autumnalis seeds. S Afr J Bot 72:157–162CrossRefGoogle Scholar
  26. Liu JG, Zhang XL, Sun YH, Lin W (2010) Antioxidative capacity and enzyme activity in Haematococcus pluvialis cells exposed to superoxide free radicals. Chin J. Oceanol. Limnol 28:1–9CrossRefGoogle Scholar
  27. Makokha AO, Ombwara FK (2002) Challenges and opportunities in commercial production of indigenous vegetables in Kenya. In: Wesonga JM, et al (eds) Proceedings of the second horticultural seminar—sustainable horticultural production in the tropics. Juja KenyaGoogle Scholar
  28. Malcoste R, Tzanni H, Jacques R, Rollin P (1972) The influence of blue light on dark-germinating seeds of Nemophila insignis. Planta 103:24–34PubMedCrossRefPubMedCentralGoogle Scholar
  29. Masondo NA, Kulkarni MG, Finnie JF, Van Staden J (2019) Influence of biostimulants-seed-priming on Ceratotheca triloba germination and seedling growth under low temperatures, low osmotic potential and salinity stress. Ecotoxicol Environ Saf 147:43–48CrossRefGoogle Scholar
  30. Mishra SS, Moharana SK, Dash MR (2011) Review on Cleome gynandra. Int J Res Pharm Chem 1:681–688Google Scholar
  31. Muasya RM, Simiyu JN, Muui CW, Rao NK, Dulloo ME, Gohole LS (2009) Overcoming seed dormancy in Cleome gynandra L. to improve germination. Seed Technol 31:134–143Google Scholar
  32. Müller K, Tintelnot S, Leubner-Metzger G (2006) Endosperm-limited Brassicaceae seed germination: abscisic acid inhibits embryo-induced endosperm weakening of Lepidium sativum (cress) and endosperm rupture of cress and Arabidopsis thaliana. Plant Cell Physiol 47:864–877CrossRefGoogle Scholar
  33. Mustafa M, Lee S (1977) Biological effects of environmental pollutants: Methods for assessing biochemical changes. Assessing toxic effects of environmental pollutants. Ann Arbor Science Publishers, Ann Arbor, pp 105–120Google Scholar
  34. Nawaz T, Ahmad N, Ali S, Khan M, Fazal H, Khalil SA (2018) Developmental variation during seed germination and biochemical responses of Brassica rapa exposed to various colored lights. J Photochem Photobiol, B 179:113–118CrossRefGoogle Scholar
  35. Ochuodho JO, Modi AT (2005) Temperature and light requirements for seed germination of Cleome gynandra L. S Afr J Plant Soil 22:49–54CrossRefGoogle Scholar
  36. Rengasamy KRR, Kulkarni MG, Stirk WA, Van Staden J (2015) Eckol-a new plant growth stimulant from the brown seaweed Ecklonia maxima. J Appl Phycol 27:581–587CrossRefGoogle Scholar
  37. Sadasivam S, Manickam A (1996) Biochemical methods, 3rd edn. New Age International Publishers, New Delhi, p 270Google Scholar
  38. Sharma HS, Fleming C, Selby C, Rao JR, Martin T (2014) Plant biostimulants: a review on the processing of macroalgae and use of extracts for crop management to reduce abiotic and biotic stresses. J Appl Phycol 26:465–490CrossRefGoogle Scholar
  39. Simlat M, Ślęzak P, Moś M, Warchoł M, Ptak A (2016) The effect of light quality on seed germination, seedling growth and selected biochemical properties of Stevia rebaudiana Bertoni. Sci Hort 211:295–304CrossRefGoogle Scholar
  40. Somboon S, Pimsamarn S (2006) Biological activity of Cleome spp. extracts against the rice weevil, Sitophilus oryzae L. J Agric Sci 37:232–235Google Scholar
  41. Sunmonu TO, Kulkarni MG, Van Staden J (2016) Smoke-water, karrikinolide and gibberellic acid stimulate growth in bean and maize seedlings by efficient starch mobilization and suppression of oxidative stress. S Afr J Bot 102:4–11CrossRefGoogle Scholar
  42. Thanos CA, Georghlou K, Douma OJ, Cl Marangaki (1991) Photoinhibition of seed germination in Mediterranean maritime plants. Ann Bot 68:469–475CrossRefGoogle Scholar
  43. Van Staden J, Brown NAC, Jäger AK, Johnson TA (2000) Smoke as a germination cue. Plant Species Biol 15:167–178CrossRefGoogle Scholar
  44. Van Staden J, Jäger AK, Light ME, Burger BV (2004) Isolation of the major germination cue from plant-derived smoke. S Afr J Bot 70:654–659CrossRefGoogle Scholar
  45. VanDer Woude WJ, Toole VK (1980) Studies of the mechanism of enhancement of phytochrome-dependent lettuce seed germination by prechilling. Plant Physiol 66:220–224PubMedPubMedCentralCrossRefGoogle Scholar

Copyright information

© Springer Nature B.V. 2019

Authors and Affiliations

  • Nkhanedzeni K. Nemahunguni
    • 1
  • Shubhpriya Gupta
    • 1
  • Manoj G. Kulkarni
    • 1
  • Jeffrey F. Finnie
    • 1
  • Johannes Van Staden
    • 1
    Email author
  1. 1.Research Centre for Plant Growth and Development, School of Life SciencesUniversity of KwaZulu-Natal PietermaritzburgScottsvilleSouth Africa

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