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Inoculation with the Native Rhizobium Gallicum 8A3 Improves Osmotic Stress Tolerance in Common Bean Drought-Sensitive Cultivar

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Inoculation with the Native Rhizobium Gallicum 8A3 Improves Osmotic Stress Tolerance in Common Bean Drought-Sensitive Cultivar Acta Agriculturae Scandinavica Section B Á Soil and Plant Science, 2012; 62: 179Á187 ORIGINAL ARTICLE Inoculation with the native Rhizobium gallicum 8a3 improves osmotic stress tolerance in common bean drought-sensitive cultivar SAMEH SASSI-AYDI, SAMIR AYDI, & CHEDLY ABDELLY Laboratoire des Plantes Extre´mophiles, Centre de Biotechnologie de Borj Cedria, BP 901, 2050 Hammam Lif, Tunisia Abstract Symbiotic nitrogen fixation potential in common bean is considered to be low in comparison with other grain legumes. However, it may be possible to improve the nitrogen fixation potential of common bean using efficient rhizobia. In order to improve osmotic stress tolerance of a drought-sensitive common bean cultivar (COCOT) consumed in Tunisia, plants were inoculated either by the reference strain Rhizobium tropici CIAT 899 or by inoculation with rhizobia isolated from native soils Rhizobium gallicum 8a3. Fifteen days after sowing, osmotic stress was applied by means of 25 mM mannitol (low stress level) or by 75 mM mannitol (high stress level). Fifteen days after treatment plants were harvested and different physiological and biochemical parameters were analysed. Results showed no significant differences between the studied symbioses under control conditions. However after exposure to osmotic stress our results showed better tolerance of COCOT to osmotic stress when inoculated with the native R. gallicum 8a3. This can be partially explained by better water-use efficiency in both leaves and nodules, better relative water content in nodules and better efficiency in utilization of rhizobial symbiosis as compared with COCOT-CIAT 899 symbiosis. Hence, the present study suggested the better use of native soil isolated strains for the inoculation of common bean in order to improve its performance and nitrogen fixation potential under stressful conditions. Keywords: Common bean, improvement, mannitol, nitrogen fixation, osmotic stress. Abbreviations: ARA, Acetylene reduction activity; DAS, Days after sowing; DW, Dry weights; EURS, Efficiency in utilization of the rhizobial symbiosis; FW, Fresh matter weight; LRWC, Relative water content of leaves; NAR, Net assimilation rate; NDW, Nodule dry weights; NF, Nitrogen fixation; Nn, Nodule number; NRWC, Relative water content of nodules; NWUE, Water-use efficiency in nodules; PWUE, Plant water use efficiency; SLA, Specific leaf area; SNF, Symbiotic nitrogen fixation; SWUE, Water-use efficiency in shoots; TSS, Total soluble sugars; TW, Turgid fresh matter weight. Introduction plant genotype which together influence the symbio- tic performance (Sadiki and Rabih 2001, Mhadhbi Common bean (Phaseolus vulgaris L.), a traditional et al. 2004). Therefore, inoculation with efficient crop originating from Latin America, is the most rhizobia might improve symbiotic nitrogen fixation important food legume for human consumption (SNF) and productivity of common bean. worldwide, especially in Africa, where its cultivation In the Mediterranean zone, little or no rainfall as a staple food extends into marginal areas. Sym- occurs during extended periods of the year. Tunisia biotic nitrogen fixation (SNF) potential in common is mostly located in the semi-arid, arid and Saharan bean is considered to be low (Pereira and Bliss 1987, climatic zones where the annual rainfall varies Isoi and Yoshida 1991) in comparison with other from 300 to less than 100 mm (Le Houe´rou 1990). grain legumes. However, it may be possible to Water deficits (commonly known as drought) can be improve the SNF potential of common bean (Bliss defined as the absence of adequate moisture neces- 1993, Hardarson et al. 1993). Yield potential of sary for plants to grow normally and complete their legumes depends on the rhizobia association and life cycle (Zhu 2002). The lack of adequate moisture Correspondence: Samir Aydi, Laboratoire des Plantes Extre´mophiles, CBBC, BP 901, 2050 Hammam Lif, Tunisia. Fax: 00216 79 412 638; Email: [email protected]; [email protected] (Received 24 April 2011; revised 16 May 2011; accepted 17 May 2011) ISSN 0906-4710 print/ISSN 1651-1913 online # 2012 Taylor & Francis http://dx.doi.org/10.1080/09064710.2011.597425 180 S. Sassi-Aydi et al. leading to water stress is a common occurrence in Mn, 4 mM Bo, 1.5 mM Cu, 1.5 mM Zn, 0.1 mM for rainfed areas, brought about by infrequent rains and micronutrients. Medium pH was maintained at 7.0 by -3 poor irrigation (Wang et al. 2005). Common bean adding 0.2 g dm CaCO3. It was aerated with a flow appears to be particularly sensitive to this stress of 400 cm3 minÁ1 of filtered air via a compressor and (Kirda et al. 1989) with considerable reduction in N2 ‘spaghetti tube’ distribution system. Plants were fixation (Ladrera et al. 2007, Sassi et al. 2008b) as a grown in a temperature-controlled glasshouse with consequence of changes in nitrogenase activity and night/day temperatures of c. 20/28 8C, relative nodule biomass (Ga´lvez et al. 2005). humidity 90/75% and a 16 h photoperiod. The Water deficiency and drought directly affect no- irradiance was supplied by mercury vapour lamps dule activity and function (Davey and Simpson (OSRAM HQI-T400W/DH). 1990). Regardless of the physiological mechanism of N2 fixation inhibition by drought stress, there is evidence that legume species have significant genetic Osmotic stress treatments variation in their ability to fix N2 under drought conditions, e.g. Pimentel et al. (1990). Several Osmotic stress was applied by means of 25 mM studies have explained the effect of water stress on mannitol (low osmotic stress level) or 75 mM plant physiology and SNF in common bean (Ramos mannitol (high osmotic stress level). The water et al. 2003, Ga´lvez et al. 2005), nevertheless, few potentials of nutrient solutions were: -0.5 MPa, studies have been conducted under hydroaeroponic 0.7 MPa and -1.2 MPa for control, 25 mM and conditions. Hydroaeroponic environment enables 75 mM mannitol respectively. Mannitol is an osmo- the comparison of different symbiotic associations tic component used generally to generate osmotic and the selection of the most tolerant symbiosis stress when added to nutrient solution. Mannitol was under stressed conditions ( Jebara et al. 2001) which added to 15-day-old plants corresponding to the are the main objectives of this study. initial period of nodules formation and the establish- In Tunisia, Mhamdi et al. (2002) showed that ment of N2 fixation. Simultaneously, nitrogen source P. vulgaris is nodulated by a diversity of species was provided to plants as 1 mL of (Rhizobium tropici ) including Rhizobium gallicum, R. leguminosarum bvs. CIAT 899 or local (Rhizobium gallicum) 8a3 strain Phaseoli and viciae, R. etli, R. giardinii, Sinorhizobium that was previously isolated from the Cap Bon region fredii, S. meliloti and S. medicae and Mnasri et al. in Tunisia, characterized at the phenotypic and (2007) showed the efficiency of the R. gallicum for molecular levels by Mhamdi et al. (1999) and kindly bean cultivation. The present work focused on the provided and maintained in culture in the Labora- enhancement of osmotic stress tolerance of a tory of Legumes Micro-organisms Interactions drought-sensitive cultivar consumed in Tunisia by (LILM), Centre of Biotechnology Borj Cedria inoculation with rhizobia isolated from native soils R. (CBBC). At the beginning of flowering, 30 DAS, gallicum 8a3 and compared with inoculation by the plants were harvested for growth parameters deter- reference strain R. tropici CIAT 899. For that mination and compared with non-stressed plants purpose, we analysed several physiological and bio- (controls). chemical traits in order (i) to look for the main traits inducing osmotic stress tolerance amelioration, (ii) to understand the likely mechanisms involved in such Dry weight and leaf area improvement and (iii) to determine useful criteria for After harvest, different plant parts were separated. genetic improvement of drought tolerance. Leaves, roots and nodules were then weighed for fresh weight determination. Leaf areas were deter- mined with a portable Area Metre (Model LI- Methods 3000A, LI COR). Dry weights (DW) of different Plant growth and conditions for imposing osmotic stress plant parts were determined after drying for 3 days at 70 8C. The biological material was bean (Phaseolus vulgaris L.) seeds of COCOT blanc (provided by M. Trabelsi, ESA Mateur, Tunisia). Seeds were surface sterilized Relative water content and pre-germinated in agar 0.9% then transferred in 1dm3 glass bottles wrapped with aluminium foil to The relative water content of leaves (LRWC) and maintain darkness in the rooting environment. The nodules (NRWC) were measured respectively in the nutrient solution contained 0.25 mM KH2PO4, second or third youngest fully expanded leaf that was 0.7 mM K2SO4, 1 mM MgSO4 ×7H2O, 1.65 mM harvested in the morning and on fresh nodules CaCl2, 22.5 mM Fe for macronutrients, and 6.6 mM harvested at the end of the treatment period. Inoculation with the native Rhizobium gallicum 8a3 181 This parameter was determined using the following Total soluble sugars determination equation: Total soluble sugars were quantified using the anthrone method. The 20 mg DW homogenate in deionized water was incubated in a water bath at RWC % 100 FW DW TW DW ð Þ¼ ½ð À Þ= ð À Þ 70 8C then centrifuged at 3000 g for 10 min. 100 mL of the supernatant was added to 4 ml of anthrone solution (0.15 g anthrone in 100mL 80% H2SO4) FW is the fresh matter weight determined within 2 h and incubated in a boiling water bath. The absor- after the harvest and TW stands for the turgid fresh bance of the samples was determined spectrophoto- matter weight (Schonfeld et al. 1988). TW was metrically at 620 nm using glucose as standard (Aydi obtained after soaking the leaves in distilled water in et al. 2010). test tubes for 4 h at room temperature (c.208C) under low light condition or after 16 h at 4 8C for Extraction and assay of leghaemoglobin nodules.
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