аЯрЁБс>ўџ ‡‰ўџџџ…†џџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџџьЅС5@ јПт9bjbjЯ2Я2 ,n­X­XИ0)џџџџџџˆ€€€€€€€”ќ$ќ$ќ$84%4h%œ”ЋA2&&&(N&N&N&N&N&N&A A A A A A A$нBR/EPDA!€-(N&N&-(-(DA€€N&N&eAх7х7х7-(Œ €N&€N&Aх7-(Aх7х7ћ7к:€€&;N&& UнE)АШќ$Й3ђђ:к;D{A0ЋAњ:,EЋ6тE&;””€€€€E€&;ДN&>Œ&,х7И&$м&QN&N&N&DADA””фx„7X””xTHE INFLUENCE OF MYCORRHIZAE ON PEA YIELD AND QUALITY IN DROUGHT CAUSED STRESS CONDITIONS Suzana KRISTEK1 - Andrija KRISTEK1  Vlado GUBERAC1  Aleksandar STANISAVLJEVI1  Sanda RA`I1 1Faculty of Agriculture, Trg Sv. Trojstva 3, Osijek, Croatia, e-mail: HYPERLINK "mailto:akristek@pfos.hr" skristek@pfos.hr Introduction Mycorrhizas are associates of fungi, representatives of Zygomyceta, Basidiomyceta and Ascomyceta living on the higher plant roots. Albert Bernard Frank in 1885 ( Brundrett et al.,1996) first discovered them. Mycorrhizal fungi fossil residues were found on the roots of the first vascular plants derived from the Devon period approx. 400 mil. years B. C. Botanists think that this symbiotic association played an essential role in developing and spreading vascular plants on the earth. However, regardless of their effect on vascular plants development of mycorrhizal fungi affected soil development. They produce humous components and an organic sticky substance (extra-cellular polysaharides) that hold soil into aggregates resulting in a well structured soil of good porosity and a large mycorrhizas intake in the total microbial bio-mass. These fungi colonize roots in about 90% of the vascular plants. The symbiotic association between higher plants and mycorrhizal fungi is highly-interrelated whereby plants benefit from their symbiont whereas fungi gets photosynthesis-formed carbon compounds. Owing to a dense hyphas network that interweaves large soil volume, an effective root absorption zone increases via hyphas (Barker et al., 1998). However, the mycorrhizal fungi also extracts chemical agents, thereby, making available elements such as phosphorus, iron, zinc, copper, boron, otherwise presented in low concentrations in a soluble form (Gryndler et al., 2003). Mycorrhizas produce antibiotics protecting plants from parasitic fungi and nematodes whereas owing to hypha network around a root, a plant is protected against parasites. Mycorrhized plants are more resistant to a drought-caused stress, have better and more adequate root system and vascular tissue development. Due to the aforesaid, dry conditions are characterized by plants having higher survival percent and higher yield elements compared to non-mycorrhized ones (Augщ et al., 2001; Vosatka et al., 2003). Thanks to the traits of this symbiosis and all benefits possessed by mycorrhized plants compared to non-mycorrhized, an agricultural production does not rely only on the indigenous strains of these fungi but inoculation of seedling seed and root is done by a compatible strain of mycorrhizal fungi. Inoculation is done by a dry treatment or wet one in a way that the inoculum suspended is water prior to inoculation (similar to legume seed inoculation with bio-preparations of the nodule bacteria). This procedure is capable for ensuring effective symbiosis and reducing mineral fertilizer and pesticide requirements being very important from economical and ecological aspect. Material and methods The trial was conducted in the greenhouse within two investigation years (2005 and 2006) on Mollic Gleysols soil type by the completely randomised design in four replicates. Four variants of the seed inoculation and various amounts of water added by irrigation were comprised by the investigation. French variety of the field pea “Bacara” was used. Basic plot size was 9 m2. However, since marginal plot parts of 20 cm wide were not analysed basic plot was 6.76 m2. Sowing was carried out in the row space 20 cm apart, within the row 5 cm whereas sowing depth was about 5 cm. The investigation elements were as follows: A) SEED INOCULATION (A1- non-inoculated seed; A2 – seed inoculated with endomycorrhizal fungus Glomus mossae spora solution); B) AMOUNT OF WATER ADDED BY IRRIGATION (B1 – total of 240 mm m-2 during the growing season; B2 – 40 mm m-2 after seeding + 40 mm m-2 in the 2-3 real leaves phase). Soil samples were taken in early spring before the experiment was established. Soil analyses were carried out by standard methods (Table 1): organic matter content was determined by bichromate method, pH in H2O and KCl, phosphorus and potassium content by ammonium-lactate method according to Egner-Riehm-Domingo (Page 1982). Table 1. Chemical properties of soil in experimental localities Investigated propertiesMollic GleysolsLayer (0 – 0.3 m)20052006pH (H2O)7.277.09pH (KCl)6.596.34Humus (%)3.073.31P (mg per 100 g soil)17.4019.20K (mg per 100 g soil)24.1022.90 Soil moisture was measured by tensiometer during field pea vegetation. Two weeks before seeding, soil moisture was approximately 65% of max. soil water capacity. At irrigation of 240 mm m-2 (during the growing season), soil moisture was maintained within 70-100% of max. soil water capacity. While adding water of 40 mm m-2 after seeding and 40 mm m-2 at the 2-3 real leaves phase, soil moisture ranged within 70-80% after irrigation, 40-45% in the grain filling phase and dropped to approximately 30-35% of max. soil water capacity in the maturation phase. Fertilization with 450 kg ha-1 NPK 15:15:15 was conducted prior to seeding. Due to weeds occurrence, the experimental plot was treated with herbicide (Basagran – 2 l ha-1) after emergence. Green mass yield per plant were determined in the blooming phase (approx. 80% of the blooming plants). Pea harvest was followed by determination of grain yield, mean count of pods per plant, mean count of grains per pod an as well as plant count per square. Nitrogen concentration in grains was determined by Kjeldahl method. Results and discussions Obtained results show that the highest values of the investigated parameters were achieved in the variants of seed inoculated with mycorrhizal fungi. This difference is especially obvious at irrigation with 80 mm water per m2 (Table 2). The highest average yield of the green mass was obtained in the mycorrhized seed variants with irrigation of 240 mm water per m2. Importance of pea irrigation in investigation results obtained Lesznyрk and Hunyadi Borbщli (2005). Mycorrhized plants obtained higher green mass yield by 124.2–159.1% at irrigation of 80 mm water per m2 compared to non-mycorrhized control. Achieved results are in the harmony with the investigation results obtained by many authors (Martin and Jamieson, 1996; Douds and Nagahashi, 2000) who reported that plants inoculated with mycorrhizal fungi in the drought-caused stress conditions had higher survival percent, thus, obtained higher green mass yield as well as higher phosphorus and nitrogen content in plant mass and grain. Table 2. Average values of investigated parameters during both investigation years Investigated parameter YearSeed inoculation LSDNon-inoculated seedInoculated seedWater added by irrigation (mm m-2)24080240800.050.01Green mass yield (g m-2)2005547.10158.90705.60411.806.189.712006543.40174.90571.30392.054.746.70Grain yield (g m-2)2005317,32133.05365.15291.054.015.402006302.80103.50341.98290.404.206.11Number of pods per plant20056.062.407.954.890.681.1620066.032.617.214.650.240.39Number of grains per pod20055,293.416.474.570.310.4320065,213.556.044.350.150.20Grain nitrogen conc. (%)20053,443.204.193.810.050.0820063,403.184.113.750.060.12Average number of plants per m2200591.6049.4097.8078.051.161.75200689.8543.9695.7572.800.390.55 The highest average values of grain yield, average number of pods per plant and average number of grains per pod were obtained in the variants of inoculated seed with mycorrhizal fungus G. mossae, regardless amount of water added by irrigation. Grain yield was in significantly positive correlation with green mass yield (r = 0.975**). Linderman (1994) stated that seed inoculated with arbuscular mycorrhizal fungi improved water and nutrients uptake in drought-caused stress conditions. He also reported that plants had better vigour, were more resistant to nematodes and disease agents resulting in higher yield elements. The highest grain nitrogen ratio was achieved in the mycorrhized variants with irrigation of 240 mm water per m2. Irrigation of 80 mm water per m2 was characterized by grain nitrogen concentration in mycorrhized plants being higher by 10.3–19.1% compared to the same one in non-mycorrhized plants. Although the seed was not inoculated with nodule bacteria, nodules were formed on the pea root in the blooming phase as a result of symbiosis with indigenous strains rhizobium bacteria. The nodules interior was red indicating active performance of atmospheric nitrogen uptake. Based on their investigation Xie et al. (1995) stated that symbiosis of plant with nodule bacteria stimulated symbiosis development in the soybean plant between plants and mycorrhizal fungi. The highest number of plants per m2 was achieved in the first investigation year in the variant of the seed inoculated with mycorrhizal fungus G. mossae. Inoculated variants had by 6.4% - 6.8% higher average number of plants per m2 compared to non-inoculated control at irrigation of 240 mm water per m2. This difference was significantly higher (57.9%-65.6%) at irrigation of 80 mm water per m2. Douds and Nagahashi (2000) reported that based on their investigation results mycorrhizal fungi improve uptake of less mobile soil nutrients in the water shortage conditions. They also improve water absorption due to hyphas mass interweaving a large soil volume. Finally, the aforesaid brings about higher percent of plants survival in the drought conditions as well as better quality of the obtained agricultural product. Conclusion Based on the obtained results of the conducted investigations it can be concluded that seed inoculated by compatible strains of mycorrhizal fungi Glomus mossae lead to better field pea plant survival in the drought stress condition and yield elements and field pea quality increase at the same time. It is from economical aspect of special importance. References Augщ R. M. - Kubikova E. - Moore J. L. : 2001. Foliar dehydration tolerance of mycorrhizal cowpea, soybean and bush bean - New Phytologist, vol. 151 no. 2 535-542pp. Barker S. J. - Tagu D. - Delp G. : 1998. Regulation of root and fungal morphogenesis in mycorrhizal symbioses - Plant Physiol., vol. 116 no. 4 1201-1207pp. Brundrett M. - Bougher N. - Dell B. - Grove T. - Malajczuk N. : 1996. Working with Mycorrhizas in Forestry and Agriculture. Monograph 32. Australian Centre for International Agricultural Research. Canberra. Douds D. D. - Nagahashi G. : 2000. Signaling and Recognition Events Prior to Colonization of Roots by Arbuscular Mycorrhizal Fungi. In: Podila G. K. and Douds D. D. (ed.): Current Advances in Mycorrhizae Research. St. Paul, The American Phytopathological Society 11 –19 pp. Gryndler M. - Hrselova H. - Malcova R. - Cajthaml T. - Halada P. : 2003. Locally accumulated extractable coumpounds in mycorrhizal parts of maize roots suppress the growth of Hyphae of G. intraradices -. Folia Geobotanica, vol. 38 no. 2 125-138pp. Lesznyрk M. – Hunyadi Borbщli Ш. : 2005. Effect of water and nutrient supply on the yield and protein yield of pea – Cereal Research Commun., vol. 33 no. 1 109-112pp. Linderman R. G. : 1994. Role of VAM fungi in biocontrol. In: Pfleger F. L. and Linderman R. G. (ed.): Mycorrhizae and Plant Health. Minnesota, APS 1-25pp. Martin R. J. - Jamieson P. D. : 1996. Effect of timing and intensity of drought on the growth yield of field peas (Pisum sativum L.) - New Zeland Journal of Crop and Horticultural Science, vol. 24 no. 2 167-174pp. Page A.L. : 1982. Methods in Soil Analysis - American Society of America, Madison, Wisconsin. Vosсtka M. - Dodd J. C. - Patten R. - Alten H. - Hutter I. - Blal B. - Dorrego A. - Estaun V. - Kapulnik Y. - Giovannetti G. - Cassels A. - Gianinazzi S. : 2003. A joint initiative for the use of mycorrhizae in plant production (the establishment of the Federation of European Mycorrhizal Fungi Producers) - Folia geobotanica., vol. 38 no. 2 235-237pp. Xie Z. P. - Staehelin C. - Vierheilig H. - Wiemken A. - Jabbouri S. - Broughton W. J. - Vogeli Lange R. - Boller T. : 1995. 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