Allelopathy of Ceratocephalus falcata on enzymatic activity of some crops seeds in germination stage

Document Type : Research Paper


1 Associate Professor Department of Plant Production and Genetics, Maragheh University, Maragheh, Iran

2 Department of chemistry, Ardabil Branch, Islamic Azad University, Ardabil, Iran


Background and objective: Allelopathic weeds by releasing inhibitors to their environment mainly affect the growth and development of adjacent plants. Although the mode of action of these materials is very complex and their targets are varied, however; seed germination inhibition is one of their important targets in most cases. In the present study, we investigated the allelopathic potential of Ceratocephalus falcata on seed germination of maize, wheat and sunflower plants on their hydrolyzing and antioxidant enzymatic systems.
Materials and Methods: The experiment was conducted as factorial in completely randomized design with three replications. Factors included three seed crop varieties of wheat, maize and sunflower (first factor) and the concentration of aqueous extract of C. falcata (second factor) with five levels including control (distilled water), 5, 10, 15 and 20% concentrations.
Results: Germination percentage decreased significantly with increasing concentration of aqueous extract of C. falcata. Germination was suppressed in all species at 10% and higher concentrations. The highest germination percentage was obtained from control treatment of wheat (99.88%) and maize (99.36%) treatments, respectively. At 5% concentration of the extract, germination percentage decreased to below 50% in all studied species. The activity of antioxidant and alpha-amylase enzymes was also affected proportionally by the concentration of extract and plant species. In most species the activity of antioxidant enzymes increased up to 10% concentration of the extract, however, at 20% concentration of the extract their activity decreased significantly compared to the control treatment. The highest activity of superoxide dismutase, catalase, peroxidase and polyphenol oxidase was observed in maize seed at medium levels of the extract, while the lowest activity of superoxide dismutase and catalase was observed in sunflower seed at 20% treatment. This treatments also resulted the lowest peroxidase activity in wheat seeds. The highest and lowest activity of polyphenol oxidase enzyme was related to 10% treatment in wheat seed and 20% concentration in sunflower seed, respectively. The activity of alpha-amylase enzyme decreased gradually with increasing concentration of the extract in all tested species.
Conclusion: The results showed that C. falcata had a very strong allelopathic effect on the germination of the studied plants. According to the results of germination, wheat had the higher sensitivity among the tested plants. The results also showed that allelochemical compounds in the aqueous extract at moderate concentrations increased levels of antioxidant activity implying that oxidative stress during seed germination of the crops has been induced. In addition, the decrease in alpha-amylase activity indicates the interference of inhibitor components with remobilization of seed storages to support the growing embryo. In general, accumulation of oxidative stress damages to seed structures and blocking of seed reserve remobilization were important reasons for drastic reduction of germination percentage in studied seeds that implies bio-herbicide potential of C. falcata.


  1. Asada, K. 1999. The water-water cycle in chloroplasts: scavenging of active oxygen and dissipation of excess photons. Annu. Rev. Plant Physiol. Plant Mol. Biol. 50: 1. 601-639.

    1. Ayyaz Khan, M., Hussain, I., and Ahmad Khan, E. 2008. Allelopathic effect of Eucalyptus (Eucalyptus camaldulensis L.) on germination and seedling growth of wheat (Triticum aestivum L.). Pakistan. J. Weed. Sci. Res. 14: 1-2. 9-18.

    3.Bais, H.P., Vepechedu, R., Gilroy, S., Callaway, R.M., and Vivanco, J.M. 2003. Allelopathy and exatrac plant invasion: from molecules and genes to species interactions. Science. 301: 5638. 1377-1380.

    4.Bogatek, R., Gniazdowska, A., Zakrzewska, W., Oracz, K., and Gawroski, S.W. 2006. Allelopathic effects of sunflower extracts on mustard seed germination and seedling growth. Biol. Plant. 50: 1. 156-158.

    5.Castillo, F.J., and Greppin, H. 1986. Balance between anionic and cationic extracellular peroxidase activities in sedum album leaves after ozone exposure. Physiol Plant. 68: 2. 201-208.

    6.Cruz-Ortega, R., Ayala-Cordero, G., and Anaya, A.L. 2002. Allelochemical stress produced by the aqueous leachate of Callicarpa acuminata: effects on roots of bean, maize and tomato. Physiol Plant. 116: 1. 20-27.

    7.Ding, C., Chachin, K., Ueda, Y., and Wang, C.Y. 2002. Inhibition of loquat enzymatic browning by sulfhydryl compounds. Food Chem. 76: 2. 213- 218.

    8.Du, Y.Y., Wang, P.C., Chen, J., and Song, C.P. 2007 .The comprehensive functional analysis of catalase gene family in Arabidopsis thaliana. N.S.F.C. 121: 1. 16-23.

    9.Du, Y.Y., Wang, P.C., Chen, J., and Song, C.P. 2008. Comprehensive functional analysis of the catalase gene family in Arabidopsis thaliana. J. Integr. Plant Biol. 50: 10. 1318–1326.

    10.El-Shora, H.M., and Abd El-Gawad, A.M. 2015. Physiological and biochemical responses of Cucurbita pepo L. mediated by Portulaca oleracea L. allelopathy. Fresenius Environ. Bull. 24: 1. 386-393.

    1. Escudero, A., Maria, J., Albert, J., Pita, M., and Felix, P.J. 2000. Inhibitory effects of Artemisia herbaalba on the germination of the gypsophyte Helianthemum squamatum. Plant Ecol. 148: 1. 71-80.
    2. Farhoudi, R., and Lee, D. 2013. Allelopatic Effects of Barley Extract (Hordeum vulgare) on Sucrose Synthase Activity, Lipid Peroxidation and Antioxidant Enzymatic Activities of Hordeum spontoneum and Avena ludoviciana. Proc. Natl. Acad. Sci. U.S.A. 83: 3. 447-452.

    13.Foyer, C.H., and Noctor, G. 2005. Oxidant and antioxidant signaling in plants: A reevaluation of the concept of oxidative stress in a physiological context. Plant Cell Environ. 28: 8. 1056-1071.

    14.Ghahreman, A. 1993. Iranian Cormophytes (Systematic Plant). TehranUniv. Press. 2: 31-34. (In Persian)

    15.Giannopolitis, C.N., and Ries, S.K. 1977. Superoxide dismutases: I. Occurrence in higher plants. Plant Physiol. 59: 2. 309-314.

    16.Gill, SS., and Tuteja, N. 2010. Reactive oxygen species and antioxidant machinery in abiotic stress tolerance in crop plants. Plant Physiol. Biochem. 48: 12. 909-30.

    17.Glenn, A. 2008. Allelopathic Interference of Invasive Acacia dealbata on the rice seedling growth, 5th World Congress on Allelopathy, New York, USA.

    1. Gniazdowska, A., and Bogatek, R. 2005. Allelopathic interactions between plants. Multi-site action of allelochemicals. Acta. Physiol. planta. 27: 3. 395-407.

    19.Hatami Hampa, A., Javanmard, A., Alebrahim, M.T., and Sofalian, O. 2018. Allelopathic effects of aqueous extract of sorghum and Acroptilon repens on seedling growth and antioxidant enzymes activity in wheat, Sugar beet, Chenopodium album and Amaranthus retroflexus. J. Plant. Protec. 32: 1. 101-119. (In Persian)

    20.Hoseinzadeh, M., Kiarostami, M., Ilkhanizadeh, M., and Saboora, A. 2009. A study on allelopathic compounds derived from Hordeum spontaneum on carbohydrates, proteins andsome Enzymes of wheat (Triticum aestivum L.). Iran. J. Biol. 22: 3. 392-406. (In Persian)

    21.Inderjit, S.O. 2003. Ecophysiological aspects of allelopathy.Planta. 217: 4. 529-539.

    1. ISTA: Amendments to ISTA Handbook on Seedling Evaluation. 2009. 3rd Edition.

    23.Jabran, K. 2017. Manipulation of Allelopathic Crops for Weed Control. First Ed. New York: Springer International Publishing. Pp: 77-85.

    24.Kar, R.K. 2011. Plant responses to water stress: role of reactive oxygen species. Plant Signal Behav. 6: 11. 1741-5.

    25.Karo, M., Mishra, D. 1976. Catalase, peroxidase and polyphenol oxidase activity during rice leaf senescence. Plant Physiol. 57: 2. 315-319.

    26.Kato-Noguchi, H., and Macias, F.A. 2005. Effects of 6-methoxy-2 benzoxazoline on Germination and amylase activity in lettuce seeds. J. Plant physiol. 162: 12. 1304-1307.

    27.Li, Zh., Wang, H., Ruan, Q.X., Pan, C.D., and Jiang, D.A. 2010. Phenolics and plant allelopathy. Molecules. 15: 12. 8933-8952.

    28.Lorenzo, P., Palomera-Pe´Rez A., Reigosa, M.J., and Gonzal, L. 2011. Allelopathic interference of invasive Acacia dealbata link on the physiological parameters of native understory species. Plant Ecol. 212: 3. 403-411.

    29.Macías, F.A., Galindo, J.C.G., Castellano, D., and Velasco, R.F. 2008. Sesquiterpene Lactones with Potential use as Natural Herbicide Models (I): Trans, trans-germacranolides. J. Agr. Food. Chem. 47: 10. 4407-4414.

    1. Machado, S. 2007. Allelopathic Potential of Various Plant Species on Downy Brome: Implications for Weed Control in Wheat Production. Agron J. 99: 1. 127-132.

    31.Mahdavikia, F., Saharkhiz, M.J., and Karami, A. 2017. Defensive response phenolic compounds of radish seedlings to the oxidative stress arising from in the extract of peppermint (Mentha piperita L.). Sci. Hortic. 214: 1. 133-140.

    1. Manikandam, R., Mobarack, H.M., Dhumal, K.N., and Murugesan, K. 2007. Influence of stanozolol and wedelia extracts on physiology of cotton var. Anjali. J. Maharashtra Agric. Univ. 32: 1. 76-79.

    33.Mishra, A. 2015. Allelopathic properties of Lantana camara. Int. J. Basic Clin. Study. 3: 1. 13-28.

    34.Mittler, R. 2004. Oxidative stress, antioxidants and stress tolerance. Trends. Plant. Sci. 7: 9. 405-410.

    35.Mittler, R., and Poulos, T.L. 2005. Ascorbate peroxidase. In antioxidants and reactive oxygen species in plants N. Smirnoff (Ed.). Blackwell Publishing Ltd, Chennai, India. Pp: 87-100.

    36.Moller, I.M., and Sweetlove, L.J. 2010. ROS signaling-specificity is required. Trends Plant Sci. 15: 7. 370-374.

    37.Newman, E.I., and Rovira, A.D. 2005. Allelopathy among some British grassland species. Ecology. 63: 3. 727-737.

    38.Omidpanah, N., Asrar, Z., and Moradshahi, A. 2011. Allelopathic potential of Zhumeria majdae essential oil on Brassica napus (Talaye cultivar). Iran. J. Plant. Biol. 7: 1. 1-9. (In Persian)

    39.Omidpanah, N., Moradshahi, A., and Asrar, Z. 2012. Investigation on allelopathic potential of Zhuceria majdae Rech. Essential oil on two wheat cultivars. Iran. J. Med. Arom. Plant. 28: 3. 198-209. (In Persian)

    40.Oracz, K., Bailly, C., Gniazdowska, A., Côme, D., Corbineau, D., and Bogatek, R. 2007. Induction of oxidative stress by sunflower phytotoxins in germinating mustard seeds. J. Chem. Ecol. 33: 2. 251-264.

    41.Parera, C., and Cantliffe, D. 1994. Dehydration rate after solid matrix alters seed performance of shrunken-2 corn. J. Am. Soc. Hortic. Sci. 119: 3. 629-635.

    42.Rassam, G.H., and Dadkhah, A. 2013. The effects of drought stress on germination and heterotrophic seedling growth characteristics of lentil (Lens culinaris Medik.). J. Agron. Sci. 5: 9. 13-24. (In Persian)

    43.Saberi, M., Shahriar, R., Jafari, M., Tarnian, A., and Safari, H. 2011. Allelopathic effect of Thymus kotschyanus on seed germination and initial growth of Bromus inermis and Agropyron elongatum. J. Watershed. Manag. Res. 93: 1. 18-25. (In Persian)

    1. Saboora, A., Amiri, N., and Kiarostami, Kh. 2012. Study allelopathic effect of some phenolic acids on the protein content and activity of some antioxidant enzymes of Amaranthus retroflexus. J. Appl. Biol. 25: 1. 53-73. (In Persian)
    2. Sasaki, K., Iwai, T., Hiraga, S., Kuroda, K., Seo, S., Mitsuhara, I., Miyasaka, A., Iwano, M., Ito, H., and Matsui, H. 2004. Ten rice peroxidases redundantly respond to multiple stresses including infection with rice blast fungus. Plant .Cell. Physiol. 45: 10. 1442-1452.

    46.Scott, S.J., Jones, R.A., and Williams, W.A. 1984. Review of data analysis methods for seed germination. Crop Sci. 24: 6. 1192-1199.

    1. Shalini, V.R., and Dubey, S. 2003. Lead toxicity induced lipid per oxidation and alters the activities of antioxidant enzymes in growing rice plant. Plant Sci. 164: 4. 645-655.

    48.Sharma, P., Jha, A., Dubey, R., and Pessarakli, M. 2012. Reactive oxygen species, oxidative damage, and antioxidative defense mechanism in plants under stressful conditions. J. Bot. 14: 1. 1-26.

    49.Singh, H.P., Batish, D.R., Kaur, S., Arora, K., and Kohli, R.K. 2006. a-pinene inhibits growth and induces oxidative stress in roots. Ann Bot. 98: 6. 1261-1269.

    50.Soltani, A., Gholipoor, M., and Zeinali, E. 2006. Seed reserve utilization and seedling growth of wheat as affected by drought and salinity. Environ. Exp. Bot. 55: 1. 195-200.

    51.Sudhakar, C., Lakshmi, A., and Giridara, Kumar, S. 2001. Changes in the antioxidant enzyme efficacy in two high yielding genotypes of mulberry (Morus Alba L.) under Naci salinity. Plant Sci. 161: 3. 613-619.

    52.Taheri, Gh. 2015. The Study of allelopathic mechanism of aqueous extract of Ferula flabelliloba in germinating seeds of Alyssum szowitsianum. J. Plant. Protec. 29: 1. 134-143. (In Persian)

    53.Thipyapong, P., Melkonian, J., Wolfe, D.W., and Steffens, J.C. 2004. Suppression of polyphenol oxidases increases stress tolerance in tomato. Plant Sci. 167: 4. 693-703.

    54.Tigre, R.C., Silva, N.H., Santos, M.G., Honda, N.K., Falcao, E.P.S., and Pereira, E.C. 2012. Allelopathic and bioherbicidal potential of Cladonia verticillaris on the germination and growth of lactuca sativa. Ecotoxicol. Environ. Saf. 84: 1. 125-132.

    55.Varier, A., and Yaduraju, N. 1996. Field emergence of cabbage seed as affected by hydro and osmo priming treatment. Seed Res. 23: 2. 116-117.

    56.Wen-Bin, W., Yun-Hee, K., Haeng-Soon, L., Ki-Yong, K., and Xi-Ping, D. 2009. Analysis of antioxidant enzyme activity during germination of alfalfa under salt and drought stresses. Plant Physiol. Biochem. 47: 7. 570-577.

    57.Yang, T., and Poovaiah, B.W. 2002. Hydrogen peroxide homeostasis: Activation of plant catalase by calcium/calmodulin. Proc. National Acad. Sci. USA. 99: 6. 4097-4102.

    58.Yarnia, M., Khorshidi Benam, M.B., and Farajzade Memari Tabrizi, E. 2009. Allelopathic effects of sorghum extracts on Amaranranthys retroflexus seed germination and growth. J. Food, Agri. Environ. 7: 3-4. 770-774.

    59.Yilmaz, H., Taskin, T., and Otludil, B. 2003. Polyphenol oxidase activity during rooting in cuttings of grape (Vitis vinifera L.) varieties.Turk. J. Botany. 27: 6. 495-498.

    60.Yu, J.Q., Fye, S., Zhang, M.F., and Hu, W.H. 2003. Effects of root exudates and aqueous root extract of cucumber and allelochemicals on photosynthesis and antioxidant enzymes in cucumber. Biochem. Syst. Ecol. 31: 2. 129-139.

    61.Zanganeh, H., and Farhoudi, R. 2011. Allelopathic Investigation of Barley Extract on Germination, Seedling Growth and Lipoxygenase Enzyme Activity of Cultivar Karun. Second National Seed Technology Conference. Pp. 123-127. (In Persian)