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International Journal of Biosciences IJB ISSN: 2220-6655 (Print) 2222-5234 (Online) http://www.innspub.net Vol. 3, No. 1, p. 121-127, 2013 RESEARCH PAPER OPEN ACCESS Assessment of seed to seed allelopathic potential of Corn (Zea mays L.) on seed and seedling growth of some volunteer species Sirous Hassannejad *, Soheila Porheidar Ghafarbi, Ramin Lotfi Department of Plant Eco-physiology, University of Tabriz, Iran Key words: Allelopathy, corn, germination, seedling growth, volunteer species. Article published on January 20, 2013 Abstract The experiment was carried out to investigate the effects of interaction seed to seed between corn (Zea mays L.) with some of volunteer species like sainfoin (Onobrychis sativa var. subinermis.), black cumin (Nigella sativa L.), bitter vetch (Vicia ervilia (L.) Willd.), proso (Panicum miliaceum L.) and jimpson-weed datura (Datura stramonium L.) on seed and seedling growth factors. Results showed that O. sativa had inhibitory effect on Z. mays seedling growth, as shoot and root length and fresh weight of Z. mays by beside of this specie was significantly inhibited. In contrast, V. ervilia and P. miliaceum had stimulatory effect on seedling growth factors of Z. mays, as shoot and root length and fresh weight of Z. mays in the presence of these species was significantly increased. Seedling of P. miliaceum by beside of Z. mays had more germination percentage, shoot fresh weight and root dry weight in comparison with control. Less germination rate of Z. mays by beside of N. sativa was caused the reducing of shoot and root fresh and dry weight of this specie. Our results suggested that the seedling growth is suitable for potential testing of the allelopathic effect of the studied seed to seed interaction under laboratory conditions. In addition, the choice of vigorous seeds can be useful to alleviation of possible allelopathic risks. * Corresponding Author: Sirous Hassannejada Sirous sirous_hasannejad@yahoo.com 121 Hassannejada et al.

Introduction Scientists have focused on the increase of food production needed for the fast expansion of the world s population. Weeds are the major problem in world agriculture because they cause losses in crop yields. Whenever two or more plants occupy the same niche in nature, they compete with each other for various life support requirements (Caton et al., 1999). Seed dispersal and seed germination are critical phases in the life-cycle of the plant during these phases the forces (biotic and abiotic stresses) have a maximum opportunity to exert their influence (Aliloo et al., 2012). A successful establishment of a weed in any ecosystem is attributed to several reasons, such as high growth rate, high reproductive potential, adaptive nature and above all interference by resource depletion and allelopathy (Kohil and Rani, 1994). Allelopathy concerns the effects of one plant on another due to chemicals released by them, or the breakdown products of their metabolites (Willis, 1994). The chemical interactions that occur among plants are known as allelopathy, and organic compounds that play a role in allelopathy are known as allelochemicals, and they become stressful when they are toxic and inhibit growth (Kocaçaliskan, 2006). Allelopathy is an ecological and chemical interaction characterized by stimulatory and inhibitory effects among different plant families. Worldwide, the inhibitory properties of the extracts and residues of many herbal species cohabiting with desired crops on the same field have been a major source of concern (Hussain et al., 2007; Nazir et al., 2007). The allelopathic properties of plants and their metabolites may be effectively used for biological weed management in crop production (Rice, 1984). Allelopathic effects of shoot and root extracts on the seed germination and early seedling growth have been frequently reported (Tanveer et al., 2008), however, few attentions have been given to seed-toseed interaction. In the natural environment, many species shed their seeds to soil surface at same season. Hence, the seed-to-seed allelopathic effect may be one of the mechanisms involved in population establishment and community pattern allelopathic effects on each other. For this reason, the objective of the study was to distinguish allelopathic impacts of Z. mays on seed germination and seedling growth of some weed and crop species. Material and methods In order to determine the effects of interaction seed to seed between (Zea mays L.) with some of weed and crop species like sainfoin (Onobrychis sativa var. subinermis.), black cumin (Nigella sativa L.), bitter vetch (Vicia ervilia (L.) Willd.), proso (Panicum miliaceum L.) and jimpson-weed datura (Datura stramonium L.) on seed and seedling growth factors, an experiment was conducted under laboratory conditions in the University of Tabriz, Tabriz, Iran in 2012. Seeds of Z. mays or weed/crop without another seeds as control were counted and 16 seeds of each species as separate placed within petri dishes which consisted of two layers of filter paper moistened with 4 ml of distilled water. For survey the effects of interaction seed to seed between Z. mays with some of weed and crop species, numbers of 16 seeds of each weed or crop species were regularly placed between the 16 wheat seeds. The distance between adjacent seeds was 1 cm, forming a grid of 4 rows by 4 columns of each species (Z. mays and each weed or crop species). Then, petri dishes were randomly incubated at 20 ºC and germinated seeds (protrusion of radicle by 2 mm) were counted every day up to 9 days. Rate of seed germination was calculated according to Ellis and Roberts (1981): GR= Where n is the number of seeds germinated on day D, D is the number of days from the beginning of the test and R is the mean germination rate. Then percentage of germination was also determined. At the end of test, length and fresh weight of root and shoot were measured. Root and shoot of each sample

were then dried in an oven at 80 ºC for 24 hours (Perry, 1977) and mean dry weight of root and shoot for each treatment at each replicate was determined. Percentage inhibition or stimulation of germination was calculated according to the following equation: Where GST is germination seeds in treatments (neighboring seed to seed) and GSC is germination seeds in control. All the data were analyzed on the basis of experimental design, using MSTATC and SPSS-16 softwares. The means of each trait were compared according to Duncan multiple range test at P 0.05 and standard error values. Excel software was used to draw figures. Fig. 1. Changing in germination percentage of Z. mays affected by some weed or crop species (a) and weed or crop species affected by Z. mays (b). Results The results revealed (Fig. 1a) that germination percentage (G %) of Z. mays was reduced by beside of O. sativa, P. miliaceum and D. stramonium. However, reduction in G% of Z. mays in the presence of the O. sativa, was not significant. In contrast, the highest G % of Z. mays was recorded in seeds by neighbored with N. sativa. Thus, the neighboring of D. stramonium and N. sativa caused maximum inhibition and stimulation in seed germination of Z. mays, respectively. The proximity of Z. mays had no effect on G% of all species. Nonetheless, G % of P. miliaceum by beside of Z. mays was increased (Fig. 1b). As a result, Z. mays had stimulatory effect on P. miliacum, but this weed specie had inhibitory effect on G% of Z. mays (Fig. 1a,b). Fig. 2. Changing in germination rate of Z. mays affected by some weed or crop seed species (a) and weed or crop species affected by Z. mays (b). Fig. 3. Changing in shoot length of Z. mays affected by some weed or crop seed species (a) and weed or crop seed species affected by Z. mays (b). Fig. 2 clearly shows that germination rate (GR) of Z. mays was decreased in the presence of all species. However, alone this reduction was significant for N. sativa. As, GR of Z. mays in the presence of this specie was decreased from 0.30 up to 0.27 than control treatment. GR of all species was inhibited by beside of Z. mays in comparison with control. However, alone N. sativa was significantly affected by Z. mays. As, GR of this specie was 32% less than that of control (Fig. 2b). Hence, the neighboring of Z. mays with N. sativa had reciprocal inhibition for both species (Fig. 2a,b). Fig. 4. Changing in root length of Z. mays affected by some weed or crop species (a) and weed and crop species affected by Z. mays (b). Shoot length (SL) of Z. mays by beside of O. sativa and D. stramonium was inhibited. In contrast, SL of Z.mays in the presence of V. ervilia and P. miliaceum was significantly stimulated. As, SL of Z. mays was 42% and 34% more in compared to the control by beside of these species (Fig. 3a). Figure 3b shows that SL of all species was declined by beside of 123 Hassannejada et al.

Z. mays. However, this reduction was significant only for N. sativa and V. ervilia. As, the inhibitory effect of Z. mays on decrease shoot elongation of these species was 42% and 47% less than that of control. As a result, V. ervilia had the highest stimulatory effect on SL of Z. mays, while this crop had the highest inhibitory effect on SL of this specie (Fig. 3a,b). Fig. 9. Inhibitory or stimulatory effects of some weed or crop species on germination percentage of Z. mays. Fig. 5. Changing in shoot fresh weight of Z. mays or crop species affected by Z. mays (b). Seedling of Z. mays by beside of O. sativa and D. stramonium was produced the lowest root length (RL). But, this specie in the presence of N. sativa, V. ervilia and P. miliaceum had the highest RL. As, RL of Z. mays by beside of P. miliaceum and O. sativa were 20.94% and 12.98% respectively, more and less than that of control treatment (Fig. 4a). Fig. 6. Changing in root fresh weight of Z. mays or crop species affected by Z. mays (b). Fig. 4b indicates that RL of N. sativa, V. ervilia and P. miliaceum was significantly inhibited by beside of Z. mays. As, RL of these species were 54.69%, 48.03% and 50.58% less in comparison with control. According to this result, N. sativa, V. ervilia and P. miliacum had stimulatory effect on RL of Z. mays. However, Z. mays had inhibitory effect on RL of these species (Fig. 4a,b). Fig. 7. Changing in shoot dry weight of Z. mays or crop species affected by Z. mays (b). Fig.8. Changing in root dry weight of Z. mays or crop species affected by Z. mays (b) Shoot fresh weight (SFW) and root fresh weight (RFW) of Z. mays were significantly reduced by beside of O. sativa, N. sativa and D. stramonium. In contrast, SFW and RFW of this specie in the presence of V. ervilia and P. miliacum were significantly increased. As, these quality of Z. mays by beside of V. ervilia were 31.53% and 65.32% and in the presence of N. sativa were 16.66% and 20.88%, respectively more and less than that of control treatments (Fig. 5a & 6a). Seedling of O. sativa and N. sativa by beside of Z. mays had the least SFW. However, this quality of P. miliaceum in the presence of Z. mays was increased (Fig. 5b). Thus, the interaction of Z. mays with P. miliaceum had stimulatory effects on SFW for both species. 124 Hassannejada et al.

While, in neighboring of Z. mays with V. ervilia, this interaction was beneficial for Z. mays but was adversely for V. ervilia (Fig. 5a,b). As a result, the stimulatory effect of Z. mays on SFW of P. miliaceum was more than that of RFW of this specie. Shoot dry weight (SDW) of Z. mays was inhibited by beside of O. sativa, N. sativa and P. miliaceum. In contrast, SDW of this specie was enhanced in the presence of V. ervilia. As, Z. mays in the neighboring of this specie had 25.92% more SDW, compared to the control (Fig. 7a). Seedling of O. sativa, V. ervilia, P. miliaceum and especially D. stramonium was showed less SDW than that of control by beside of Z. mays. However, N. sativa in the presence of this specie was indicated more SDW than that of control (Fig. 7b). presence of allelochemicals that inhibited the processes of seed germination. According to El- Khatib (1977), an indirect association between lower seed germination and allelophatic inhibition may be the consequence of the inhibition of water uptake. Lower seed germination was recorded partly due to the fact that the presence of the some allelochemicals in the seed prevented the growth of seed embryo or caused its death. Allelochemicals are found to be released to environment in appreciable quantities via root exudates, leaf leachates, roots and other degrading plant (seed) residues, which include a wide range of phenolic acids such as benzoic and cinnamic acids, alkaloids, terpenoids and others (Rice, 1995). These compounds are known to modify growth, development of plants, including germination and early seedling growth. Fig. 8a shows that root dry weight (RDW) of Z. mays in the presence of P. miliaceum and D. stramonium was significantly increased. This specie by beside of P. miliaceum had 25.92% more RDW in comparison with control (Fig. 8a). In contrast, RDW of D. stramonium in this interaction was 93.10% less than that of control (Fig. 8b). Also, this quality in O. sativa and V. ervilia was declined by beside of Z. mays. As a result, Z. mays had the highest inhibitory effect on SDW and RWD of this weed specie. In general, seeds of O. sativa, V. ervilia, P. miliaceum and D. stramonium had inhibitory effect on germination percentage of Z. mays. In contrast, N. sativa had stimulatory effect on germination of this crop. Thus, in weed control or crop rotation management, it needs to increase precision in selective of crop. Our results indicated that existing seeds of N. sativa by beside of Z. mays may be beneficial for Z. mays in germination phase (Fig. 9). Discussion The interaction among Z. mays with some of weed or crop species shows that germination percentage and especially germination rate of Z. mays and some of these weed species significantly decreased (Fig. 1 & 2). Reduction in seed germination might be due to Fig. 3-8 indicates that moreover seed germination percentage and rate, seedling growth was also affected by allelochemicals that produced in interaction between Z. mays and some of weed or crop species. This effect was as inhibitory or stimulatory mood. The mechanism of inhibition on the seedling growth caused by allelochemicals can be the result of reduced cell division and/or cell elongation (Iman et al., 2006). Few studies have been reported that legumes crop residues have been stimulatory effects on studied crops. For example, Gill et al., (2000) have reported stimulatory effect of mungbean, sesame and soybean residues on wheat, chickpea and lentil at lower concentration. Improvement in corn yields following legume crops has also been reported (Phetchawee et al., 1985) Phenolic allelochemicals from O. sativa can inhibit amino acid transport and protein synthesis, and the subsequent growth of treated plants. All phenolics could reduce integrity of DNA and RNA (Zeng et al., 2001). In addition, aqueous extracts of O. sativa can increase indoleacetic acid oxidase activity of receptor plants, and then reduce the level of indoleacetic acid (Zeng et al., 2001). 125 Hassannejada et al.

Einhelling (2002) reported that individual compounds can have multiple phytotoxic effects on a plant. There was a reduction in radicle growth of the germinating seeds because the seeds allelopathic might have contained allelochemicals that affected the enzymes responsible for plant hormons synthesis (Fujii et al., 1991). There is an inhibitory effect on root elongation when the root comes into contact with the matter with inhibitory chemicals as described in early works with various crops and weeds (Qasem, 1995). Gill S, Anoliefo O. Iduoze U. 2000. Allelopathic effects of aqueous extract from siamm weed on the growth of cowpea. Department Botany, University of Botany, Nigeria. Hussain S, Siddiqui S, Khalid S, Jamal A, Qayyum A, Ahmad Z. 2007. Allelopathic potential of Senna (Cassia angustifolia Vahl.) on germination and seedling characters of some major cereal crops and their associated grassy weeds. Pakistan Journal of Botany 39, 1145-1153. References Aliloo AA, Shahabivand S, Farjam L, Heravi S. 2012. Allelopathic effects of Pine Needle extracts on germination and seedling growth of Ryegrass and Kentucky bluegrass. Advances in Environmental Biology, 6, 2513-2518. Caton BP, Mortimer AM, Hill TC, Gibson JE, Fisher AJ. 1999. Weed morphology effects on competitiveness for light in direct-seeded rice. Proc. 17 th Asian-Pacific, weed science society conference, Bangkok, 1, 116-120. Iman A, Wahab S, Rastan M, Halim M. 2006. Allelopathic effect of sweet corn and vegetable soybean extracts at two growth stages on germination and seedling growth of corn and soybean varieties. Journal of Agronomy 5, 62-68. Kocaçalışkan İ. 2006. Allelopati, Bizim Büro Basım Yayın, Ankara. Kohil KK, Rani D. 1994. Parthenium hysterophorus--a review. Research Bulletin (Sci.) Panjab University. 44, 105-149. Einhellig FA. 2002. The physiology of allelochemical action: clues and views. In Allelopathy: From Molecules to Ecosystems (Eds., Reigosa, M.J. & Pedrol, N.). Science Publishers, Enfield, New Hampshire, p. 1-23.. El-Khatib AA. 1997. Does allelopathy involves in the association pattern of Trifolium resupinatu. Biologia Plantarum 40, 425-431. Ellis RH, Roberts EH. 1981. The quantification of ageing and survival in orthodoxes seeds. Seed Sci. and Technol. 9, 373-409. Fujii Y, Furukawa M, Hayakawa Y, Sugahara K, Shibuya T. 1991. Survey of Japanese medicinal plants for the detection of allelopathic properties. Weed Research (Tokyo) 36, 36-42. Nazir T, Uniyal AK, Todari NP. 2007. Allelopathic behaviour of three medicinal plant species on traditional agriculture crops of Garhwal Himalaya, India. Agroforestry System 69, 183-187. Perry, D.A. 1977. A vigour test for seed of barley (Hordeum vulgare L.), based on measurement of plumule growth. Seed Sci. and Technol. 5, 709-719. Phetchawee S, Vibulsukh N, Theppoolpon M, Masarugson W. 1985. Long term effect of mulching with fertilizer under corn-legumes intercropping on crop yield and improvement of soil chemical and physical properties. In: Thailand National Corn and Sorghum Program 1985 AnnualReport. Department of Agriculture, Kasetsart University and CIMMYT pp: 204-212. 126 Hassannejada et al.

Quasem JR. 1995. The allelopathic effect of three Amaranthus spp. (Pigweeds) on wheat (Triticum durum). Weed Research 35, 41-49. Rice EL. 1984. Allelopathy, 2nd Edn. Academic Press, Orlando, Florida. Science Publishers, Enfield, New Hampshire. Rice EL. 1995. Biological control of weeds and plant diseases. Univ. Oklahoma Press, Norman. seedling growth of crops. Allelopathy Journal 21, 317-327. Willis RJ. 1994. Terminology and trends in allelopathy. Allelopathy Journal 1, 6-28. Zeng RS, Luo SM, Shi YH. 2001. Physiological and biochemical mechanism of allelopathy of secalonic acid on higher plants. Agronomy Journal 93, 72-79. Tanveer A, Tahir M, Nadeem MA, Younis K, Aziz A, Yaseen M. 2008 Allelopathic effects of Xanthium strumarium L. on seed germination and 127 Hassannejada et al.