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corrhizal inoculation INTRODUCTION Tomato (Lycopersicon esculentum Mill.) is one of the most important vegetable plants in the world (Nicola et al., 2009; Panthee and Chen, 2010; FAOSTAT, 2014). It is widely cultivated in tropical, subtropical and temperate climates and thus ranks third in terms of world vegetable production (FAOSTAT, 2014). Diseases are a major limiting factor for tomato production. The most important diseases are those caused by pathogens that include fungi, bacteria and viruses. These diseases are a major limiting factor and can be severe, reducing tomato yield and quality. Due to the drawbacks of the chemical plant protection method, consumer pressure and the introduction of international food safety standards (IFSS), a need has arisen for farmers to change their production and marketing practices. As required by the IFSS, agricultural producers may consider it necessary to use alternative pest management methods, especially in ecological production. This production system takes into account biological methods, understood not only as direct protection, but also as the use of the natural processes of self-regulation. One of these methods is biological control, which provides environmentally safe ways to reduce the adverse effects of pathogens on crops. Potential biological control agents include arbuscular mycorrhizal fungi (AM Fungi) (Al-Askar and Rashad, 2010). AM Fungi are a promising source for improving sustainable food production and increasing human © 2020 Institute of Agrophysics, Polish Academy of Sciences A. JAMIOŁKOWSKA et al. 254 nutritional needs (Hart et al., 2015). AM Fungi used in horticulture are an effective means of improving plant growth (Oseni et al., 2010). Mycorrhiza is a natural phenomenon offering many benefits for farmers and, therefore, may be useful in sustainable plant development (Conversa et al., 2007; Candido et al., 2015). AM Fungi are a major component of most plant rhizospheres and they play an important role in reducing the occurrence of plant diseases. Different studies indicate that AM Fungi increase the nutrient uptake of plants. This improves the resistance of plants to pathogens through compensation for the symptoms of infection, and the morphological and biochemical changes in plants, as well as the increase in the number of antagonistic microorganisms in the rhizosphere (Kapoor, 2008; Song et al., 2015). Mycorrhizal fungi stimulate plant defence reactions at the initial stages of symbiosis (Paszkowski, 2006). The modulation of plant defence during the establishment of mycorrhiza causes a pre-conditioning of tissues by effectively activating the defence mechanisms of the plants, which is known as the priming phenomenon (Pozo and Azcón-Aguilar, 2007). It plays a major role in mycorrhiza-induced resistance (Jung et al., 2012). Song et al. (2015) found that plant mycorrhization increases tomato plant resistance to early infection by Phytophthora infestans by activating the jasmonic acid (JA) signaling pathway. Numerous studies have confirmed that AM Fungi enhance plant resistance against various pathogens (Harrier and Watson, 2004; Pozo et al., 2005; Bi et al., 2007). Mycorrhizal colonization improved tomato resistance to wilt diseases caused by Fusarium oxysporum f. sp. lycopersici and increased tomato resistance to the foliar disease of early blight caused by Alternaria solani (Akköprü and Demir, 2005; Fritz et al., 2006). The appropriate management of AM Fungi has the potential to provide sustainability in agricultural systems for crop disease management (Zeng, 2006; Liu et al., 2007). However, the mechanism of disease resistance induced by AM Fungi remains elusive. It is well known that transcriptional reprogramming occurs in the host-plant upon mycorrhizal colonization (Jung et al., 2012; Lopez-Ráez et al., 2010). Mycorrhizal fungi control not only soil-borne pathogens but evidence has been presented which shows the suppression of leaf attacking pathogens. Fritz et al. (2006) found that the protective effect of mycorrhiza against the development of tomato leaf spots caused by A. solani has an effect equivalent to induced systemic resistance. Common mycorrhizal networks between tomato plants provide the protection of neighbours against early blight (Song et al., 2015). In recent times, the system of the classification of fungi forming or considered to form arbuscular mycorrhiza placed them in phylum Glomeromycota and class Glomeromycetes with four orders (Archaeosporales, Diversisporales, Glomerales, and Paraglomerales), ten families and thirteen genera (Walker et al., 2007; Palenzuela et al., 2008; Oehl et al., 2011; Błaszkowski, 2012). Considering the above-mentioned aspects, it is believed that there is a need to study and select the species of mycorrhizal fungi which – when inoculated into the rhizosphere of tomato – will contribute to better plant nutrition, their improved growth and health status. Hence, the objectives of this study were to: I – determine the influence of AM Fungi on root colonization, II – investigate the variability of leaf nutrient content, growth and yield of tomato plants under the influence of the studied species of mycorrhizal fungi, III – determine which mycorrhizal fungus (MF) species will be better to use in the ecological production of tomato. MATERIALS AND METHODS The research work was carried out at an ecological farm located in Grądy in the Lublin province, Poland (51°05′ N latitude, 22°12′ E longitude) in the years 2015-2017. The object of the studies included three cultivars of tomato hybrids (Lycopersicon esculentum Mill.) important for commercial production: ‘Antalya F1’ (Yuksel Tohomculuk Company in Turkey), ‘Esmira F1‘ (Rijk Zwaan Company in Holland), ‘Pelikan F1’ (Seed and Nursery Company in Ożarów Mazowiecki in Poland) and two species of mycorrhizal fungi (MF): Claroideoglomus etunicatum (CE) (syn. Glomus etunicatum W.N. Becker & Gerd) and Rhizophagus intraradices (RI) (syn. Glomus intraradices N.C. Schenck & G.S. Sm.). MF spores were provided from the collection belonging to the Department of Agricultural Microbiology, the Institute of Soil Science and Plant Cultivation – National Research Institute in Puławy, Poland. The experiment was established in a plastic tunnel with the dimension of 6×30×2.80 m. The experiment was carried out on albic luvisols (cutanic) soil (Kabała et al., 2019). In each year, organic nitrogen fertilizer (Bioilsa 12.5 – NaturalCrop Poland) was used before planting plants in a permanent place using a dose of 500 kg ha. Tomatoes were grown in soil with the following parameters: pH in H2O 5.83-6.66 (KQ/PB-47); levels of available minerals: P-39.0-63.0 mg kg f.w. of soil (KQ/PB-51); K-314.0-477 mg kg f.w. of soil (KQ/PB-52); Mg-60.0-89.0 mg kg f.w. of soil (KQ/ PB53); Ca-462.0-619.0 mg kg f.w. of soil (KQ/PB-52); total nitrogen: 23.4-63.5 mg kg f.w. of soil) (Chemical and Agricultural Station in Lublin). Cucurbitaceae plants (cucumber) were the preceding crops for each year of the study. Tomato seedlings were produced in accordance with the generally accepted rules for this vegetable using horticultural soil (peat moss) and quartz sand at a ratio of 2:1 (Welbaum, 2015). Four week old seedlings (at the stage of three true leaves) were planted in a plastic tunnel in the first week of May where the distance between the rows was 0.80 m, and the spacing between the plants in the row was 0.50 m (2.5 plants m). Before planting the plants in a permanent place, mycorrhizal inoculum was applied using 25-30 spores of MF in physiological saline solution (5 ml) to the MYCORRHIZA AS AN ALTERNATIVE FOR THE ECOLOGICAL PRODUCTION OF TOMATO 255 rhizosphere of each seedling (to a depth of 5 cm). In each year of the studies, the field experiment was randomized in a complete block design as a two-factor experiment with five replicates and three treatments. The experimental treatments were: plants inoculated with C. etunicatum (CE), plants inoculated with R. intraradices (RI), plants without mycorrhizal inoculation as control (C). The abovementioned treatments were used for each of the tomato cultivars. Black film was used for mulching in the rows in order to protect plants from weeds. Plants were irrigated using drip irrigation depending on their requirements. No fertilization or chemical protection was applied during the vegetation period. The colonization of roots by the tested mycorrhizal fungi (MF) was measured at the stage of full fructification of the tomato plants (BBCH 89). The evaluation of root colonization (%) by MF was carried out using the Phillips and Hayman method (Phillips and Hayman, 1970). The percentage of root colonization was calculated when five plants were taken from each experimental combination. The lateral roots of these plants were selected, washed under tap water and cut into 0.5 cm fragments. Washing was performed, so that the mycelium and the spores, which are attached to the roots, would not be washed away. After cleaning, the root fragments were transferred into 10% KOH solution and boiled at 90 ̊C for 1 h. After cooling, the root fragments were washed in distilled water and transferred into 1% HCl at 90 ̊C for 20-30 min to acidify the roots. Next, the root fragments were washed in distilled water 2-3 times. They were stained with 0.05% trypan blue solution. The stained root fragments were placed in De-stain solution, then placed on a slide for the observation of the mycelium of MF under the microscope. The colonization of the roots with MF was estimated using the following formula (Phillips and Hayman, 1970) :

Mycorrhizal inoculation as an alternative for the ecological production of tomato (Lycopersicon esculentum Mill.)