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Results and discussion

Evaluated wheat varieties showed significant genetic variation in their allelopathic activities ranging between +10.9% (cv. Naveed) -- -3O.2% (cv. PI 4512) in terms of their effect on increase or decrease in dry weight of Avenue, and +14.0% (cv. Naveed) -- -29.8% (cv. Inia) on its height. Nine of the evaluated varieties exhibited more than 20% inhibition of the dry weight of the test weed. However, seven genotypes/varieties e.g. Baiyat, Ghods, Inia, Kavkaz, Khazar-1, PI 4512 and Surkhu caused 25.8, 27.9, 28.3, 25.7, 28.3, 30.2 and 26. 1% reduction in the dry weight, respectively, indicating the existence of gene(s) for allelochemical(s) production which in turn inhibit the growth of A. ludoviciana. Interestingly, Bezostaya-1, Naveed and Niknejad varieties exerted a positive allelopathy, and promoted the dry weight of Avenue by 6.6, 10.9 and 10.4%, respectively (Table 1).

When the number of wheat seeds were increased in the assay boxes, the extent of inhibition was further enhanced. However, varieties, Bezostaya-1, Naveed and Niknejad that had positive effects on the growth of the test weed at low seed density, showed some inhibitory effect but there was no significant change in the height of wheat plants when grown with increased density (Table 2). This indicates that the adverse or inhibitory effect of wheat varieties on Avenue is mediated through the release of chemical inhibitors (allelopathy) rather than simple competition, otherwise growth of wheat plants would have also been affected. Furthermore, none of the wheat varieties showed any autotoxicity.

It is generally presumed that plant species with high growth vigour are better competitors of weeds and are allelopathically superior. However, we could not get any correlation between the growth (height) and allelopathic activity of different wheat varieties. Bezostaya-1, a variety with maximum height (35.1 cm) caused a 6.6% promotion in Avenue dry weight whereas Inia with a comparatively less height (26.3 cm) inhibited the test weed by 28.3%. Naveed and Niknejad varieties had almost similar plant heights with those of Karaj-1, Karaj-3 and Khazar-1 but the earlier ones had a 10% promotory effect while the latter varieties inhibited the test weed by 13.0% to 28.3% (Fig. 1). These results indicate that in this case the allelopathic activity is independent of the early growth vigour of wheat varieties. In order to see any possible genetic variation in allelopathic activity of wheat genotypes at different growth stages (the age of plants) their effect on dry matter and. plant height of Avenue was studied. All the allelopathic varieties showed a similar trend of inhibition and the allelopathic activity increased with the advancing age of wheat seedlings (Fig. 2 and Fig. 3). Out of the tested wheat genotypes, Inia, Jenab, Karaj-3, Kavkaz, Khazar-1, PI -4512 and Surkhu showed more than 60% inhibition in the growth rate of Avenue during 22nd to 26th day (Table 1, Fig. 2 and Fig. 3). PI-4512 caused the maximum inhibition (74.6%). Inhibition caused by different wheat genotypes can he attributed to the presence of major gene(s) controlling the production of allelochemicals like acetic, propionic, butyric, vanillic, syringic, p-coumaric, p-hydroxybenzoic and hydroxamic (2,4-dihydroxy-7-methoxy-1,4-benzoxazin-3-one) acids which are known to cause reduction in plant growth, total biomass and act as herbicide (Tang and Waiss 1978; Perez 1990; Alsaadawi et al. 1998).

These preliminary results indicating great genetic variability in bread wheat genotypes for allelochemical production can be of paramount importance in view of the success achieved with allelopathic rice (Olofsdotter 1998), in the control of weeds and reduction in the use of herbicides. These results have also revealed that it is not the early growth vigour of some of the wheat genotypes, helpful in the control or reduction of weed population in wheat field, rather it is the ability of wheat genotypes to produce certain types of allelochemicals which suppress, inhibit or kill the competing weed. Possibility of identifying strongly allelopathic wheat varieties possessing major gene(s) to control weeds under field. condition lies in screening a large number of genetically diverse genotypes and their wild relatives. Putnam and Duke (1978) have suggested that wild types and progenitors of existing crops may have possessed high allelopathic activity and this character was reduced or lost as they were hybridised and selected for other useful characteristics. The facts that allelopathic activity is genetically inherited (Panchuk and Prutenskaya 1973), and is caused in synergistic manner involving a complex of chemicals (Rizvi et al. 1992) which is produced under the action of more than one gene (Alborn et al. 1992) make the wheat allelopathy research very attractive. However, unfortunately in wheat which is one of the most widely researched and grown crop throughout the world, the research on its allelopathic trait is negligible. These studies clearly indicate the possibilities of exploiting allelopathic trait through genetic manipulations while breeding new wheat cultivars which will help the farmers in reducing the cost of production and at the same time the approach will be environment friendly.

Allelopathic crop varieties may not only lead to the reduction in the use of herbicides, these could also be used as a 'source' of allelopathic trait for breeding crops with strong 'weed resistance'. However, evaluation of a large number of genetically diverse wheat genotypes, identification of useable gene(s) controlling allelochemicals production, inheritance, their mode of action and non-target toxicity, are prerequisite before allelopathic wheat genotypes can he used commercially. Experiments are in progress on the above mentioned aspects.

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