ࡱ>  PRGHIJKLMNO5@ hbjbj22 rlXXϲ&$PNLlej"""""""$R!dE""""uuuX""uu6uu:<{,{" Pk70:h{ }E50er{Rq{0{"Vxul0"""d[zbkuzbINFLUENCE OF BACTERIUM P. FLUORESCENS ON ROOT DECAY AGENT R. SOLANI, STORING PERIOD AND ELEMENTS OF SUGAR BEET YIELD AND QUALITY Suzana Kristek, Andrija Kristek, Milan Pospiail, Renata Glavaa Toki, Jasenka osi ____________________________________________________________________ Dr. Suzana Kristek, assistant professor, Department for Microbiology, Faculty of Agriculture, Trg Sv. Trojstva 3, 31 000 Osijek, Croatia. E-mail:  HYPERLINK "mailto:skristek@pfos.hr" skristek@pfos.hr Dr. Andrija Kristek, full professor, Department for sugar beet breeding, Faculty of Agriculture, Trg Sv. Trojstva 3, 31 000 Osijek, Croatia. Dr. Milan Pospiil, associate professor, Department for Plant Production, Faculty of Agronomy, Svetoaimunska 25, 10 000 Zagreb, Croatia. Renata Glavaa Toki, graduate engineer, Sugar refinery  Kandit Premijer d.o.o.  , Frankopanska 99, 31 000 Osijek, Croatia. Dr. Jasenka osi, assistant professor, Department for Plant Protection, Faculty of Agriculture, Trg Sv. Trojstva 3, 31 000 Osijek, Croatia. ABSTRACT The influence of sugar beet seed inoculation with the bacterium Pseudomonas fluorescens and treatment with the fungicide Thiram42S with and without seed inoculation aiming to control the root decay agents Rhizoctonia solani was studied during a 2 year trial on two soil types (Mollic Gleysols and Eutric Cambisols). The influence of the treatments on parameters of sugar beet yield and quality such as root yield, sugar content, sugar in molasses, sugar yield as well as percentage of the infected and decayed plants as a consequence of parasitic fungi infestation will be described. Values of investigation parameters depended on soil types, hybrids (tolerant and sensitive) and investigation years. The highest average of investigated parameters in both investigation years were obtained on Mollic Gleysols in the variant of the seed inoculated with Pseudomonas fluorescens bacterium (variant 2). On Eutric Cambisols the highest average of these parameters were obtained in the variant of the seed treated with fungicide Thiram 42 S (200 ml/100 kg seed) + inoculated with Pseudomonas fluorescens bacterium (variant 3). Key words: sugar beet, root decay, R. solani, P. fluorescens, storing period, yield, quality. INTRODUCTION During the process of sugar beet cultivation, depending on weather conditions, degree of soil infection with pathogenic microorganisms, and application of soil management, the presence of sugar beet root decay is frequently observed. Pathogenic agents are numerous, inflicting serious damage. According to the occurrence time of root decay symptoms, the agents can be divided into three major groups: agents occurring during the emergence phase, agents occurring in the phase of vegetation, and ones that cause sugar beet root decay during the period of storing. Most frequent decay agents on sugar beet seedlings are fungi Aphanomyces cochlioides, Pythium ultimum and Pythium debarianum, as well as Rhizoctonia solani. In sugar beet vegetation phase plant decay is most frequently caused by fungi Rhizoctonia solani, Fusarium ssp., and bacteria Erwinia ssp. From the root decay agents most damage is inflicted by fungi Rhizoctonia solani. Besides weather conditions that influence this type of root decay, the inadequate soil management irregular soil cultivation resulting in damaged soil structure and rise in soil acidity, is a contributory factor. Apart from the decay agents occurring during cultivation on the field, certain pathogens cause sugar beet decay in the period of storing, and before processing. Storing conditions, as well as plant damages caused when digging are contributory factors in the occurrence of fungal diseases. Most frequent pathogenic microorganisms observed are fungi, such as Rhizoctonia solani, Botrytis cinerea, Penicillium spp., Aspergillus spp. ... Pathogenic fungus Rhizoctonia solani, if compared to the remains sugar beet root decay agents, is able to infect the plant from the emergence phase, during vegetation period, until the process of sugar beet storing. Fungicide treatment of seeds, application of tolerant hybrids, or proper soil management does not ensure full plant protection against this pathogen. Moreover, heavily infected soils and weather conditions favourable for development of the pathogen, contribute to the infection of higher or lower degree. The development of the fungi is mainly prevented by treating seeds with fungicides; however, they seriously affect human health and environment whilst targeted pathogenic fungi rapidly develop resistance against them. When growing tolerant hybrids, particularly on slightly infected soils, lower yield values and digestion are observed. Apparently, in newer selection of sugar beet hybrids, the difference is slightly lower if comparing to the older ones. Therefore, bearing in mind environmental aspect of the problem (application of high fungicide doses), seed inoculation with bacteria showing antagonism against pathogenic fungi is an acceptable alternative to chemical pesticide application ( HYPERLINK "javascript:popRef('b1')" Andersen etal., 2003). The best results reported in the studies dealing with this issue, were obtained by inoculating seeds with Pseudomonas fluorescens. Pseudomonas fluorescens belongs to rod-shaped, asporogenous, gram-negative bacteria that are as saprophytes prevalent in soil and water belong to a group of soil microorganisms crucial to the process of soil denitrification. Due to the production of antimicrobial agents, these bacteria also show a distinguished antibiosis against pathogens causing diseases on arable crops, by inactivating their growth and reproduction ( HYPERLINK "javascript:popRef('b20')" Whipps, 2001) For the capacity to synthesize toxic cyclic lipopeptides ( HYPERLINK "javascript:popRef('b19')" Thrane etal., 2000;  HYPERLINK "javascript:popRef('b8')" Koch etal., 2002;  HYPERLINK "javascript:popRef('b1')" Andersen etal., 2003) they are used as effective biological control agents ( HYPERLINK "javascript:popRef('b14')" Nielsen etal., 1999;  HYPERLINK "javascript:popRef('b18')" Sorensen etal., 2001). Many authors have reported that these purified lipopeptides show an antagonistic activity against certain fungi pathogenic on sugar beet roots such as Rhizoctonia solani (Nielsen et al., 2000, 2002; Andersen et al., 2003), Aphanomyces cochlioides, (Nielsen et al., 1999; Sorensen et al., 2001), Pythium ultimum and Pythium debarianum ( HYPERLINK "javascript:popRef('b19')" Lee etal., 2000; Nielsen etal., 2000; Thrane et al., 2000;  HYPERLINK "javascript:popRef('b1')" Andersen etal., 2003). This suggests that bacteria producing lipopeptides could have a potential role in the biocontrol of fungal diseases, which was approved in both laboratory and field trials (Thrane et al., 2000, 2001). ). Lipopeptides may also function as biosurfactants (Desai et al., 1997), which can facilitate bacterial growth on water insoluble carbon sources (Koch et al., 1991; Ron and Rosenberg, 2001) or their interaction with hydrophobic surfaces (Neu, 1996), e. g., surface motility (Lindum et al., 1998). Therefore, seed inoculation with bacteria showing antagonism against pathogenic fungi is an acceptable alternative to chemical pesticide application ( HYPERLINK "javascript:popRef('b1')" Andersen etal., 2003). Moreover, considering the heavily infected soils with the pathogen Rhizoctonia solani and the fact that beneficial bacterium Pseudomonas sp. is not sensitive to a low dose fungicide ( HYPERLINK "javascript:popRef('b16')" Pedersen etal. 2002), it is possible to treat the seeds combining low doses of the chemical agents aiming to control growth and reproduction of the pathogenic fungi, which in consequence, produce positive effect on all the parameters of sugar beet yield and quality. MATERIALS AND METHODS The experiment was set up on two soil types - Mollic Gleysols (FAO, 1998) and Eutric Cambisols (Table 1), with the presence of pathogenic fungi Rhizoctonia solani the sugar beet root decay agent detected from 2001 2004. The experiment was set up in 2004 and 2005 in completely randomized block design, with 4 repetitions and 24 different variants, including 4 tolerant and 4 sensitive hybrids to the pathogenic fungi, and 3 various seed treating variants: 1 hybrids A hybrids tolerant to the fungi Rhizoctonia solani A1 Laetitia (KWS) A2 Flores (Danisco) A3 Gazeta (Hilleshg) A4 Solea (Strube) B hybrids sensitive to the fungi Rhizoctonia solani B1 Belinda (KWS) B2 Georgina (KWS) B3 Sofarizo (Hilleshg) B4 Buda (Strube) 2 seed treatment C1 control seed treated with fungicide Thiram 42-S (600 ml/100 kg) C2 seed inoculated with Pseudomonas fluorescens bacterium C3 - seed treated with fungicide Thiram 42-S (200 ml/100 kg) + inoculated with Pseudomonas fluorescens bacterium Pseudomonas fluorescens was isolated from the sugar beet seedling rhizosphere ( HYPERLINK "javascript:popRef('b19')" Thrane etal., 2000) and cultivated on King's B medium (Pseudomonas F; Difco catalog No. 0448-17-1). Fluorescent Pseudomonas spp. could be detected by illuminating the agar plates with UV light (254 nm) and randomly picking the fluorescent colinies. The inoculum was applied directly to the seed before sowing at a concentration of about 8נ104 bacteria/seed, i.e. 0.81.4נ1010 bacteria/ha. Row seeding was completed at the end of March, with spacing of 50 cm between rows and 20cm within rows. After sugar beet digging conducted in the mid-October, determination of root yield (t/ha) was done. Simultaneously, two samples of 30 kg for each variant were taken. One sample at time was analyzed after digging, followed by determination of sugar content (%), sugar in molasses (%) and sugar yield (t/ha). Second sample was analyzed after 30 days storing in net bag, in the middle of the sugar beet production prism, and the same parameters were determined. Weather conditions appeared to affect sugar beet growth. Data obtained from Agrometeorological bureau in Osijek (Table 2) were used in weather analysis. Both trial years in comparison to the multiyear average values showed higher humidity and air temperature values, particularly in 2005. During vegetation period (April October) 2005, the rainfall was 622 mm, in 2004 it was slightly lower (551 mm), with multiyear average rainfall of 436 mm. Increased precipitation values favourably dispersed through the year enabled even sugar beet emergence, high field germination values and free thicker root development. However, rise in rainfall values in the period of the highest increase in sugar content (since August 15) when sugar beet needs were poor affected sugar content increase. August 2005 had very high rainfall value (218 mm), followed by September having 72 mm, with air temperature of 17.4 C. Air temperature and precipitation recorded in the period of the highest sugar content increase were above optimum values for sugar beet plant, as well as above multiyear average for the area. Results were processed by modern statistical methods (ANOVA) using computer program by StatSoft Inc. (2001) STATISTICA (data analysis software system), version 6. RESULTS AND DISCUSSION 1. Percentage of infected and decayed plants During 2-year investigations conducted significant differences were shown in the percentage of infected and decayed plants, soil types investigated, tolerant and sensitive hybrids, and hybrids within the two groups (Table 3, 4). On Eutric Cambisols during a 2-year investigation, averagely 10.13 % more infected plants, and 9.69 % more decayed plants were observed if comparing to the results on Mollic Gleysols. This could be due to the heavier infection of the Eutric Cambisols. Furthermore, second trial-year showed higher percentage of infected and decayed plants (precipitation) in both tolerant and sensitive hybrids on both soil types. Both tolerant and sensitive hybrids on Mollic Gleysols had the lowest percentage of infected and decayed plants in the variant of the seeds treated with the bacteria Pseudomonas fluorescens (variant 2). On Eutric Cambisols for heavier infection with the fungi Rhizoctonia solani the lowest percentage of infected and decayed plants was detected in the variant of the seeds inoculated with bacteria Pseudomonas fluorescens combined with lower doses of Thiram 42-S fungicide (variant 3). An average percentage of infected sugar beet plants with tolerant hybrids during 2-year trial on both soil types in variant 1 (control seed treated with fungicide Thiram 42 S, 600 ml/100 kg seed) was by 4.10 % lower, while an average percentage of decayed plants was by 3.86 % lower in comparison to the sensitive ones. The lowest percentage of infected and decayed plants on both soil types was obtained with Flores, a tolerant hybrids and Georgine, a sensitive one. The results obtained are in agreement with those of Whipps (2001) who reported that the plants inoculated with bacteria Pseudomonas fluorescens were characterized with rapid initial growth that enabled fast undergoing through the most vulnerable phase when the damage of the pathogen affecting was the most pronounced. Due to the pronounced antagonism of the bacterium to the root decay agent of sugar beet fungi Rhizoctonia solani high survival percentage of inoculated plants was obtained in comparison to the non-inoculated ones and decrease in the damage of infected plants, which consequently, influenced the sugar beet root yield. 2. Root yield Sugar beet root yield depended on soil types, investigation years and hybrid. The highest root yield during 2-year trial was obtained on Mollic Gleysols (Table 5), with a hybrid sensitive to pathogenic fungi Rhizoctonia solani Georgina, in the variant of the seed inoculated with Pseudomonas fluorescens bacterium (variant 2). The remains hybrid (tolerant and sensitive) obtained statistically very significant lower values of the parameter (P<0.01). The highest root yield per ha with tolerant hybrid was obtained in 2004 with Laetitia hybrid, also in the variant of the seed inoculated with Pseudomonas fluorescens bacterium (variant 2). However, no significant differences were recorded between variants 2 and 3 (seed treated with fungicide Thiram 42 S (200 ml/100 kg seed) + inoculated with Pseudomonas fluorescens bacterium). In the second year of investigation the highest value of the parameter was obtained with Flores hybrid in the variant of the seed treated with fungicide Thiram 42 S (200 ml/100 kg seed) + inoculated with Pseudomonas fluorescens bacterium (variant 3), though no significant differences were recorded between variants 3 and 2 (seed inoculated with Pseudomonas fluorescens bacterium). Hybrids tolerant to pathogenic fungi Rhizoctonia solani Flores and Gazeta were applied as new hybrids in the second year of investigation when they were introduced in a production process. On Eutric Cambisols (Table 6) in both investigation years the highest root yield with tolerant hybrids was obtained with Laetitia, in the variant of the seed treated with fungicide Thiram 42 S (200 ml/100 kg seed) + inoculated with Pseudomonas fluorescens bacterium (variant 3). In both investigation years the highest root yield in hybrids sensitive to the pathogenic fungi Rhizoctonia solani was obtained with Georgina, in the variant of the seed treated with fungicide Thiram 42 S (200 ml/100 kg seed) + inoculated with Pseudomonas fluorescens bacterium (variant 3). In both cases the remains variants obtained statistically lower values of the parameter. In the second year of investigation on both soil types lower root yield was obtained due to the less favourable weather conditions during sugar beet vegetation. On Mollic Gleysols in the second year of investigation tolerant hybrids obtained up to 10.09 17.72% and hybrids sensitive to the pathogenic fungi Rhizoctonia solani up to 3.00 10.20% lower root yields in comparison to the first year of investigation. On Eutric Cambisols the difference in tolerant hybrids was 8.50 16.00% and 2.40 9.00% in sensitive ones. 3. Sugar content Sugar content in the samples analyzed immediately after digging and after 30 days storing period depended, respectively, on seed treatment, hybrid (tolerant and sensitive to Rhizoctonia solani ), type of the soil and investigation year. On Mollic Gleysols (Table 7) sugar content in hybrids tolerant to pathogenic fungi Rhizoctonia solani (samples analyzed immediately after digging) ranged from 11.90-17.01%, while sensitive hybrids had 13.10-17.28%. By sample analysis after 30 days storing sugar content in tolerant hybrids ranged from 11.81-16.91%, while sensitive hybrids had 12.22-17.20%. The highest differences between tolerant and sensitive hybrids were recorded in the control (seed treated with fungicide Thiram 42 S, 600 ml/100 kg seed). Differences in sugar content between samples analyzed immediately after digging and after 30 days storing with hybrids tolerant to Rhizoctonia solani ranged from 0.4-1.4%. Sensitive hybrids obtained significantly higher differences (5.2-8.4%). Differences in sugar content between the samples analyzed immediately after digging and after 30 days storing in tolerant and sensitive hybrids were comparable in variant 2 (seed inoculated with Pseudomonas fluorescens bacterium) and in variant 3 (seed treated with fungicide Thiram 42 S (200 ml/100 kg seed) + inoculated with Pseudomonas fluorescens bacterium). In some cases within the hybrids, significant differences were recorded between the seed treated variants; however, no rules can be followed regarding hybrids, type of soil and investigation year. Apparently, in the second year of investigation these values were, as well, slightly lower. On Eutric Cambisols (Table 8) sugar content in hybrids tolerant to pathogenic fungi Rhizoctonia solani (samples analyzed immediately after digging) ranged from 11.46-16.89%, while sensitive hybrids had 12.07-17.20%. By sample analysis after 30 days storing sugar content in tolerant hybrids ranged from 11.31-16.81%, while sensitive hybrids had 11.46-17.09%. With tolerant hybrids differences in sugar content in variant 1 sugar samples (control seed treated with fungicideThiram 42 S (600 ml/100 kg seed) analyzed immediately after digging and after 30 days storing were slightly higher than recorded on Mollic Gleysols and ranged from 1.3-3.6%. On the other hand, in sensitive hybrids the aforesaid differences were lower than recorded on Mollic Gleysols measuring 3.6-5.9%. Differences in sugar contents between tolerant and sensitive hybrids, in variants 2 (seed inoculated with Pseudomonas fluorescens bacterium) and 3 ((seed treated with fungicide Thiram 42 S (200 ml/100 kg seed) + inoculated with Pseudomonas fluorescens bacterium) between samples analyzed immediately after digging and after 30 days storing were significantly lower in comparison to the control. However, the highest values of the parameter investigated on Mollic Gleysols, and within the hybrids, were obtained in variant 2, while on Eutric Cambisols variant 3 had the highest parameter values. Namely, on Eutric Cambisols higher infection intensity was recorded with the pathogenic fungi Rhizoctonia solani. This is in agreement with the trial of Pedersen et al (2000) who have reported that soils heavily infected with pathogenic fungi Rhizoctonia solani, and regarding the fact that beneficial bacterium Pseudomonas sp. is usually not especially fungicides sensitive, seed is possible to be treated in combination with low doses of the fungicide aiming at effective control of pathogenic fungi growth and reproduction resulting in a positive effect on all sugar beet yield and quality indicators. 4. Sugar in molasses Sugar content in molasses depended, respectively, on seed treated variant, hybrid properties (tolerant, sensitive), soil type investigated, and year of investigation (weather conditions during vegetation phase of sugar beet plant). On Mollic Gleysols (Table 9) the lowest values of the parameter were obtained in variant 2 (seed inoculated with Pseudomonas fluorescens bacterium). This is in agreement with the results of Lifshitz et al. (1987) and Whipps (2001) who stated that Pseudomonas fluorescens as plant growth promoting bacteria increased solubility of phosphorus in inorganic forms round the active root zone. By mobilizing soil phosphorus plants vigour improves and yield increases by 1520% (great root availability reduce negative nitrogen effect on sugar beet plant achieving balanced plant nutrition and reduced production of alpha - amino nitrogen, potassium and sodium i.e. decrease of the amount of sugar which cannot be isolated in the production process but turns into molasses.) Sugar molasses mean value in variant 1 (treated with fungicide Thiram 42-S (600 ml/100 kg)) in tolerant hybrids in samples analyzed after 30 days storing was by 2.11 4.00% higher than in samples analyzed immediately after digging. The difference in hybrids sensitive to pathogenic fungi R. solani ranged from 2.72 9.59%. On Eutric Cambisols (Table 10) the lowest sugar values in molasses were recorded in variant 3 (seed treated with fungicide Thiram 42 S (200 ml/100 kg seed) + inoculated with Pseudomonas fluorescens bacterium). Sugar molasses mean value in the control variant of tolerant hybrids to pathogenic fungi R. solani in samples analyzed after 30 days storing was by 2.21 3.14% higher than in samples analyzed immediately after digging. In sensitive hybrids the difference was 5.70 7.51%. Apparently, on both soil types the percentage of sugar in molasses was higher in the second year of investigation. 5. Sugar yield The highest sugar yield on Mollic Gleysols (Table 11) during 2-year trial and in all the hybrids tested was obtained in variant 2 (seed inoculated with Pseudomonas fluorescens bacterium), while variant 3 (seed treated with fungicide Thiram 42 S (200 ml/100 kg seed) + inoculated with Pseudomonas fluorescens bacterium) showed the highest sugar yield value on Eutric Cambisols (Table 12). On Mollic Gleysols, in the control variant of tolerant hybrids sugar yield of the samples analyzed immediately after digging was by 1.73 3.77% higher in comparison to the samples analyzed after 30 days storing. On Eutric Cambisols the difference was 0.83 1.67%. Sugar yield on Mollic Gleysols in variants 2 and 3 of tolerant hybrids in samples analyzed immediately after digging was by 0.31 1.65% higher in comparison to the samples analyzed after 30 days storing. On Eutric Cambisols the difference was 0.08 1.46%. Sensitive cultivars in the control variants on both soil types obtained significantly higher aforesaid differences in comparison to the tolerant ones (Mollic Gleysols 5.30 12.82%; Eutric Cambisols 5.90 - 12.10%). However, the difference in sugar yield in variants 2 and 3 in samples analyzed immediately after digging and after 30 days storing was lower than with tolerant hybrids (Mollic Gleysols 0.07 0.87%; Eutric Cambisols 0.47 1.28%). Sugar yield in the second year of investigation on both soil types and all the hybrids tested was lower than in the first investigation year. This could be due to the aforesaid weather conditions in the period of sugar beet vegetation. Sugar yield was in very significant positive correlation with root yield (r=0.944; P<0.01) and sugar content (r=0.920; P<0.01). CONCLUSION Inoculation of sugar beet seed with the bacterium Pseudomonas fluorescens affected root decay agent treated with pathogenic fungi Rhizoctonia solani and showed highly significant influence to all the elements of sugar beet yield and quality. The highest values of the elements investigated (the least in parameter Sugar in molasses) on Mollic Gleysols soil type were obtained in variant 2 (seed inoculated with Pseudomonas fluorescens bacterium). On Eutric Cambisols soil type the same was obtained in variant 3 (seed treated with fungicide Thiram 42 S (200 ml/100 kg seed) + inoculated with Pseudomonas fluorescens bacterium). Tolerant hybrids exhibited low differences among the samples analyzed after digging, and after 30 days storing, in all the variants of the parameters tested. The difference obtained with sensitive hybrids was high in variant 1 (treated with fungicide Thiram 42-S (600 ml/100 kg), while in variants 2 and 3 differences were lost showing no differences comparing to the tolerant variants. REFERENCES [1] Andersen, J.B., Koch, B., Nielsen, T.H., Srensen, D., Hansen, M., Nybroe, O., Christophersen, C., Srensen, J., Molin, S. and M. Giskov: Surface motility in Pseudomonas sp. DSS73 is required for efficient biological containment of the root pathogenic microfungi Rhizoctonia solani and Pythium ultimum. Microbiology 149 (2003), 37-46. [2] Desai, J.D. and I. M. Banat: Microbial production of surfactants and their commercial potential. Microbiol. Mol. Biol. Rev. 61 (1997), 47-64. [3] FAO. World Reference Base for Soil Resources, (1998). FAO Roma. [4] Koch, A.K., Kappeli, O., Fiechter, A. and J. Reiser: Hydrocarbon assimilation and biosurfactant production in Pseudomonas aeruginosa mutants. J. Bacteriol. 173 (1991), 4212 -4219. [5] Koch, B., Nielsen, T. H., Srensen, D., Andersen, J. B., Christophersen, C., Molin, S., Giskov, M., Srensen, J. and O. Nybroe: Lipopeptide production in Pseudomonas sp. strain Dss73 is regulated by components of sugar beet seed exudates via the Gac two component regulatory system. Appl. Environ. Microbiol. 68 (2002), 4509-4516. [6] Lee, C.H., Kempf, H. J., Lim, Y. and H. Cho: Biocontrol activity of Pseudomonas cepacia AF2001 and anthelmintic activity of its novel metabolite, cepacidine A. J. Microbiol. Biotechnol. 10 (2000), 568-571. [7] Lifshitz, R., Kloepper, J. W. and M. Kozlowski: Growth promotion of canola (repeseed) seedlings by a strain of Pseudomonas putida under gnotobiotic conditions. Can. J. Microbiol. 33 (1987), 390-395. [8] Lindum, P.W., Anthoni, U., Christophersen, C., Eberl, L., Molin, S. and M. Givskov: N Acyl L homoserine lactone autoinducers control production of an extracellular lipopeptide biosurfactant required for swarming motility of Serratia liquefaciens MG1. J. Bacteriol. 180 (1998), 6384-6388. [9] Neu, T.R.: Significance of bacterial surface active compounds in interaction of bacteria with interfaces. Microbiol. Rev. 60 (1996) 151-166. [10] Nielsen, T.H., Christophersen, C., Anthoni, U. and J. Srensen: Viscosinamide, a new cyclic depsipeptide with surfactant and antifungal properties produced by Pseudomonas fluorescens DR54. J. Appl. Microbiol. 87 (1999), 80-90. [11] Nielsen, T. H., Thrane, C., Christophersen, C., Anthoni, U. and J. Srensen: Structure, production characteristics and fungal antagonism of tensin a new antifungal cyclic lipopeptide from Pseudomonas fluorescens strain 96.578. J. Appl. Microbiol. 89 (2000), 992 1001. [12] Nielsen, T. H., Srensen, D., Tobiasen, C., Andersen, J. B., Christophersen, C., Giskov, M. and J. Srensen: Antibiotic and biosurfactant properties of cyclic lipopeptides produced by fluorescent Pseudomonas spp. from the sugar beet rhizosphere. Appl. Environ. Microbiol. 68 (2002), 3416-3423. [13] Pedersen, H.C., Weiergang, I., Pontoppidan, M.M., Jrgensen, L. and A. Svingel : Danisco Seed, Seed Technology Dept., (2002). Hoejbygaardvej 31, DK 4960 Holeby, Denmark. [14] Ron, E.Z., and E. Rosenberg: Natural roles of biosurfactants. Environ. Microbiol. 3 (2001), 229-236. [15] Sorensen, D., Nielsen, T.H., Christophersen, C., Srensen, J. and M. Gajhede: Cyclic lipoundecapeptide amphisin from Pseudomonas sp. Strain Dss73. Acta Crystallogr. Sect. C Cryst. Struct. Commun. 57 (2001), 1123 1124. [16] Thrane, C., Nielsen, T.H., Nielsen, M.N., Srensen, J. and S. Olsson: Viscosinamide producing Pseudomonas fluorescens DR54 exerts a biocontrol effect on Pythium ultimum in sugar beet rhizosphere. FEMS Microbiol Ecol. 33 (2000),139-146. [17] Thrane, C., Nielsen, M.N., Srensen, J. and S. Olsson: Pseudomonas fluorescens DR54 reduces sclerotia formation, biomass development, and disease incidence of Rhizoctonia solani causing damping off in sugar beet. Microb. Ecol. 42 (2001), 438445. [18] Whipps, J.M.: Microbial interactions and biocontrol in the rhizosphere. Journal of Experimental Botany, 52 (2001), 487 511. [19] www.statsoft.com Table 1. Soil characteristics. Investigated properties in a fieldType of soilLayer (0 0.3 m)Mollic GleysolsEutric CambisolspH (H2O)7.426.46pH (KCl)6.445.98Humus (%)3.221.87P (mg/ 100 g soil)24.4422.50K (mg/ 100 g soil)29.5723.11 Table 2. Weather conditions in experimental years and years long average in Osijek. (Meteorological and hydrological service of Croatia (DHMZ). Agrometeorological bureau in Osijek; 2004, 2005). MonthPrecipitation amountsAir temperaturesNecessity1901-199120042005Necessity1901-199120042005April405612256-11.211.411.6May5063636014.216.815.417.4June5088889218.019.419.820.1July80665811718.521.222.121.8August656110521818.220.421.419.6September3546457214.016.815.817.4October4056807-11.113.011.7Total360436551622----Average-----16.716.917.1 Table 3. Percentage of the infected plants (%) as a consequence of R. solani infection in the two to four true leaves stage. Hybrid Seed treatmentSoil type AverageMollic GleysolsEutric CambisolsYear2004200520042005 Tolerant to the Rhizoctonia solani  Laetitia 121.6023.3322.7625.9023.4028.038.9710.0412.369.85310.5011.178.669.7110.01 Flores 1-22.50-23.5123.012-8.16-10.239.203-10.00-8.959.48 Gazeta 1-23.01-23.9623.492-8.56-11.4510.013-10.49-8.979.73 Solea 123.7524.4424.7426.9224.96210.8113.0112.0314.7012.64312.0614.1510.9913.6212.71 Sensitive to the Rhizoctonia solani  Belinda 129.9032.8533.4536.6033.2029.4410.3011.7012.1510.9039.9210.2110.9511.9910.77 Georgina 123.1025.7526.2529.8025.4828.859.6011.1513.9010.8839.0310.1010.2213.1010.61 Sofarizo 125.2827.0727.0530.4547.31210.7011.4012.1212.7311.74311.1012.3511.6512.4011.88 Buda 128.2032.4529.6634.7531.2728.609.2510.3512.9510.2939.109.9010.1011.0810.051 - control seed treated with fungicideThiram 42 S (600 ml/100 kg seed); 2 seed inoculated with Pseudomonas fluorescens bacterium; 3 - seed treated with fungicideThiram 42 S (200 ml/100 kg seed) + inoculated with Pseudomonas fluorescens bacterium. Table 4. Percentage of the decayed plants (%) as a consequence of R. solani infection in the two to four true leaves stage. Hybrid Seed treatmentSoil type AverageMollic GleysolsEutric CambisolsYear2004200520042005 Tolerant to the Rhizoctonia solani  Laetitia 115.3016.4515.7917.9416.3724.484.595.325.815.4836.046.674.795.165.67 Flores 1-15.90-16.5716.242-4.71-5.945.333-5.55-4.915.23 Gazeta 1-16.17-16.9016.542-4.73-6.995.863-5.00-4.344.67 Solea 116.7017.0317.5119.2617.6325.336.246.688.136.5936.737.986.047.317.02 Sensitive to the Rhizoctonia solani  Belinda 126.8129.4629.8833.9530.0324.064.815.655.945.1234.895.354.815.425.12 Georgina 121.2523.2623.8426.7123.7724.214.965.806.355.3334.625.215.366.905.52 Sofarizo 122.6424.5624.8827.0124.7724.364.836.027.205.6034.905.175.416.115.40 Buda 125.8029.0126.4530.9028.0425.165.835.906.405.8235.416.215.576.225.851 - control seed treated with fungicideThiram 42 S (600 ml/100 kg seed); 2 seed inoculated with Pseudomonas fluorescens bacterium; 3 - seed treated with fungicideThiram 42 S (200 ml/100 kg seed) + inoculated with Pseudomonas fluorescens bacterium. Table 5. Root yield in both investigation years on soil type Mollic Gleysols (t/ha). Mollic GleysolsHybridSeed treatmentRoot yield (t/ha)20042005 Tolerant to the Rhizoctonia solani  Laetitia 177.3166.12282.1971.27381.8072.45 Flores 1-69.802-74.233-75.18 Gazeta 1-69.142-68.383-70.12 Solea 178.0364.20279.1971.20378.2468.38 Sensitive to the Rhizoctonia solani  Belinda 178.1871.32282.0577.16380.9274.11 Georgina 190.1285.462104.2496.80399.6191.13 Sofarizo 188.0585.41297.6890.20391.9088.10 Buda 189.1780.10295.6289.90391.2084.16 LSD0.05 0.853 1.260 LSD0.01 1.450 2.077 1 - control seed treated with fungicideThiram 42 S (600 ml/100 kg seed); 2 seed inoculated with Pseudomonas fluorescens bacterium; 3 - seed treated with fungicideThiram 42 S (200 ml/100 kg seed) + inoculated with Pseudomonas fluorescens bacterium Table 6. Root yield in both investigation years on soil type Eutric Cambisols (t/ha). Eutric CambisolsHybridSeed treatmentRoot yield (t/ha)20042005 Tolerant to the Rhizoctonia solani  Laetitia 170.2164.90276.3568.34380.1071.12 Flores 1-61.352-67.903-68.48 Gazeta 1-61.182-64.903-65.12 Solea 164.1056.40271.6360.22371.2065.17 Sensitive to the Rhizoctonia solani  Belinda 174.0067.38275.3173.38378.7075.11 Georgina 188.2584.12294.3087.55397.9190.90 Sofarizo 186.1883.10292.1286.01392.3587.30 Buda 184.2482.21289.1886.12391.4887.50 LSD0.05 1.143 1.010 LSD0.01 2.190 2.040 1 - control seed treated with fungicideThiram 42 S (600 ml/100 kg seed); 2 seed inoculated with Pseudomonas fluorescens bacterium; 3 - seed treated with fungicideThiram 42 S (200 ml/100 kg seed) + inoculated with Pseudomonas fluorescens bacterium Table 7. Sugar content in both investigation years on soil type Mollic Gleysols (%). Mollic GleysolsHybridSeed treatmentSugar content (%)20042005ABAB Tolerant to the Rhizoctonia solani  Laetitia 113.8413.4712.3011.81214.9014.9212.7612.68314.1114.0512.5412.60 Flores 1--13.4613.292--13.8913.733--13.7113.60 Gazeta 1--13.2113.532--13.8813.813--13.6013.55 Solea 116.0415.8813.9213.81217.0116.9114.9614.93316.8616.7214.8014.69 Sensitive to the Rhizoctonia solani  Belinda 115.0714.2813.5812.44217.1617.0314.6014.51316.8416.7914.5414.38 Georgina 115.8615.0112.4811.51216.2716.1913.7613.62316.1416.1313.0112.89 Sofarizo 115.4714.2113.1012.22216.9816.8614.5214.46316.0616.1014.4014.37 Buda 115.6214.7714.0713.18217.2817.2015.5715.51316.4516.3915.4915.46 LSD0.05 0.213 0.305 0.192 0.143 LSD0.01 0.380 0.550 0.423 0.260 1 - control seed treated with fungicideThiram 42 S (600 ml/100 kg seed); 2 seed inoculated with Pseudomonas fluorescens bacterium 3 - seed treated with fungicideThiram 42 S (200 ml/100 kg seed) + inoculated with Pseudomonas fluorescens bacterium A samples analyzed immediately after digging; B samples analyzed after 30 days storing Table 8. Sugar content in both investigation years on soil type Eutric Cambisols (%). Eutric CambisolsHybridSeed treatmentSugar content (%)20042005ABAB Tolerant to the Rhizoctonia solani  Laetitia 113.2112.9711.4611.31213.9613.9012.1212.07314.0114.0712.3012.22 Flores 1--13.1012.862--13.4413.313--13.8013.68 Gazeta 1--12.4411.992--12.9712.923--13.0113.04 Solea 115.9115.6813.0812.84216.4416.3913.9713.85316.8916.8114.2514.15 Sensitive to the Rhizoctonia solani  Belinda 114.2613.5512.9012.14215.9315.9413.8313.81316.8116.7714.2114.08 Georgina 115.1014.3712.0711.46215.8615.7213.1113.05316.0315.9413.4813.44 Sofarizo 114.9914.1012.5612.02215.6015.5514.0113.76315.9215.9814.5614.57 Buda 115.5314.9714.1013.35216.8416.7715.3615.31317.2017.0915.6815.52 LSD0.05 0.091 0.122 0.110 0.166 LSD0.01 0.173 0.204 0.242 0.330 1 - control seed treated with fungicideThiram 42 S (600 ml/100 kg seed); 2 seed inoculated with Pseudomonas fluorescens bacterium 3 - seed treated with fungicideThiram 42 S (200 ml/100 kg seed) + inoculated with Pseudomonas fluorescens bacterium A samples analyzed immediately after digging; B samples analyzed after 30 days storing Table 9. Sugar in molasses in both investigation years on soil type Mollic Gleysols (%). Mollic GleysolsHybridSeed treatmentSugar in molasses (%)20042005ABAB Tolerant to the Rhizoctonia solani  Laetitia 11.581.641.761.8221.301.371.561.6131.481.521.621.65 Flores 1--1.611.662--1.401.453--1.491.51 Gazeta 1--1.501.562--1.371.393--1.4 21.44 Solea 11.421.451.791.8421.251.261.571.6031.311.341.611.63 Sensitive to the Rhizoctonia solani  Belinda 11.281.391.471.5121.161.171.301.3931.191.211.411.48 Georgina 11.561.691.701.8121.321.331.511.5331.411.441.561.60 Sofarizo 11.451.581.781.8721.281.301.391.4231.331.351.501.54 Buda 11.391.481.461.6021.141.161.191.2131.231.261.271.30 LSD0.05 0.021 0.024 0.036 0.026 LSD0.01 0.037 0.043 0.061 0.053 1 - control seed treated with fungicideThiram 42 S (600 ml/100 kg seed); 2 seed inoculated with Pseudomonas fluorescens bacterium 3 - seed treated with fungicideThiram 42 S (200 ml/100 kg seed) + inoculated with Pseudomonas fluorescens bacterium A samples analyzed immediately after digging; B samples analyzed after 30 days storing Table 10. Sugar in molasses in both investigation years on soil type Eutric Cambisols (%). Eutric CambisolsHybridSeed treatmentSugar in molasses (%)20042005ABAB Tolerant to the Rhizoctonia solani  Laetitia 11.601.651.811.8521.471.511.661.7031.381.421.621.65 Flores 1--1.621.662--1.511.543--1.431.46 Gazeta 1--1.591.642--1.491.513--1.421.46 Solea 11.491.531.781.8221.381.411.631.6531.331.361.601.64 Sensitive to the Rhizoctonia solani  Belinda 11.331.411.481.5721.241.281.301.3331.201.231.231.26 Georgina 11.581.671.731.8621.451.481.611.6331.341.361.571.59 Sofarizo 11.471.561.801.9121.341.371.511.5531.311.341.421.45 Buda 11.441.531.491.5821.251.291.351.3831.181.211.231.26 LSD0.05 0.029 0.018 0.030 0.036 LSD0.01 0.050 0.035 0.058 0.061 1 - control seed treated with fungicideThiram 42 S (600 ml/100 kg seed); 2 seed inoculated with Pseudomonas fluorescens bacterium 3 - seed treated with fungicideThiram 42 S (200 ml/100 kg seed) + inoculated with Pseudomonas fluorescens bacterium A samples analyzed immediately after digging; B samples analyzed after 30 days storing Table 11. Sugar yield in both investigation years on soil type Mollic Gleysols (t/ha). Mollic GleysolsHybridSeed treatmentSugar yield (t/ha)20042005ABAB Tolerant to the Rhizoctonia solani  Laetitia 110.9610.698.498.31211.6311.559.799.80311.5711.479.619.54 Flores 1--11.1610.892--12.3212.123--12.0712.16 Gazeta 1--9.6311.282--11.9511.873--11.1411.05 Solea 111.1810.9911.0310.83212.7412.7012.0112.04312.4912.3411.8611.72 Sensitive to the Rhizoctonia solani  Belinda 113.2012.4611.9711.03214.5014.4813.8813.76314.3114.2913.5413.54 Georgina 114.0113.1710.9810.01215.4615.3913.7613.71314.9714.9613.2213.25 Sofarizo 112.3711.039.158.11214.8414.8510.9910.92313.7113.6610.7310.68 Buda 114.0012.9711.8310.55215.2915.1713.2613.19314.6814.4912.5812.56 LSD0.05 0.065 0.081 0.062 0.091 LSD0.01 0.139 0.155 0.125 0.152 1 - control seed treated with fungicideThiram 42 S (600 ml/100 kg seed); 2 seed inoculated with Pseudomonas fluorescens bacterium 3 - seed treated with fungicideThiram 42 S (200 ml/100 kg seed) + inoculated with Pseudomonas fluorescens bacterium A samples analyzed immediately after digging; B samples analyzed after 30 days storing Table 12. Sugar yield in both investigation years on soil type Eutric Cambisols (t/ha). Eutric CambisolsHybridSeed treatmentSugar yield (t/ha)20042005ABAB Tolerant to the Rhizoctonia solani  Laetitia 110.6810.518.208.11211.5511.549.439.43311.8411.879.589.50 Flores 1--10.9010.812--11.6511.623--11.9911.94 Gazeta 1--9.159.002--10.7110.683--10.8710.79 Solea 110.1610.0310.6610.55211.5111.4611.5111.47311.7911.6211.8411.83 Sensitive to the Rhizoctonia solani  Belinda 112.3711.5311.4610.80214.0113.8512.8412.76314.2613.9113.2113.20 Georgina 113.4612.7110.199.09214.6814.6012.0511.99314.9114.8412.5712.50 Sofarizo 111.1310.229.098.31212.8512.789.679.68313.4113.409.919.84 Buda 113.2712.1210.9510.20214.2214.0912.8312.77314.9814.7913.3113.33 LSD0.05 0.103 0.231 0.090 0.135 LSD0.01 0.217 0.399 0.161 0.257 1 - control seed treated with fungicideThiram 42 S (600 ml/100 kg seed); 2 seed inoculated with Pseudomonas fluorescens bacterium  "68:BD\^`fjtv   $ & 4 6 8 : Ƚhx5mH sH hB5mH sH hL5mH sH hs5mH sH hI5mH sH h756mH sH hc56mH sH hPhs56mH sH h75mH sH hc5mH sH hPhs5mH sH 7j6 8 : @  i v V Y 4  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