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The Rhizobia Nodulating Shrubs for Revegetation of Arid Lands

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The Rhizobia Nodulating Shrubs for Revegetation of Arid Lands Arid Land Research and Management, 19:307–326, 2005 Copyright # Taylor & Francis Inc. ISSN: 1532-4982 print/1532-4990 online DOI: 10.1080/15324980500299649 The Rhizobia Nodulating Shrubs for Revegetation of Arid Lands: Isolation of Native Strains and Specificity of the Plant – Rhizobia Interaction by Cross Inoculation Tests Fernando Gonzaalez-Andre ´s Escuela Universitaria de Ingenierı´aTe´cnica Agrı´cola INEA, Valladolid, Spain Jesu´s Alegre Instituto Madrile, Madrid, Spain Jose´-Luis Ceresuela Escuela Te´cnica Superior de Ingenieros Agroonomos, Universidad Polite´cnica de Madrid, Madrid, Spain Medicago strasseri, M. citrina, Colutea arborescens, and Dorycnium pentaphyllum are legume shrubs potentially useful for the revegetation of semiarid Mediterranean ecosystems and for animal grazing. As nodulation with specific rhizobia and=or mycorrhizal fungi, may become essential for the settlement and growth of legumes, the objective was to ascertain the specificity=promiscuity of the plant–rhizobia inter- action, by detecting the occurrence of infective rhizobia in the soils to be revegetated and by cross inoculation tests. Native rhizobia for the target species, were isolated from nondisturbed native areas. The four species were inoculated with four semiarid soils located in central Spain, that we intend to revegetate. Specific rhizobia for M. strasseri and M. citrina were detected in three of the four soils. The soil that did not show infective rhizobia had received sewage-sludge from a waste-water treatment plant, and levels of heavy metals were high. None of the 4 soils had specific rhizobia for C. arborescens and D. pentaphyllum. The cross inoculation tests revealed that M. strasseri and M. citrina belonged to the same symbiotic grouping as alfalfa. On the other hand C. arborescens and D. pentaphyllum only nodulated with their own specific native rhizobia, and the rhizobia from C. arborescens and D. pentaphyl- lum were not infective with any of the tested species. For M. strasseri the effective- ness of the native rhizobia was significantly higher than the effectiveness of the infective rhizobia from the soils to be revegetated. Consequently for M. strasseri, C. arborescens and D. pentaphyllum, inoculation with native strains (I.MsS1, I.CA and I.DP respectively) would be advisable. Received 17 September 2004; accepted 19 January 2005. This work has been financially supported by projects 06M=016=96, 07M=0076=98, and 07M=0023=2000 from Comunidad de Madrid, Spain. We want to thank INEA students Ana-Esther Fernaandez, Ana Cuadrado, and David Meneses for their help. Address correspondence to Fernando Gonzaalez-Andre ´s, Escuela Superior y Te´cnica de Ingenierı´a Agraria. Universidad de Leoon. Avda. de Portugal, 41. 24071 Leoon, Spain. E-mail: [email protected] 307 308 F. Gonzaalez-Andre ´s et al. Keywords shrubby legumes, N-fixation, rhizobia Shrubby plants have a double interest in the degraded semiarid Mediterranean ecosystems that usually present severe water stress, scarcity of plant available nutri- ents, and low microbiological activity (Caravaca et al., 2002). First, it has been demonstrated that drought tolerant shrubby species are of great interest for the recovery of degraded, or even desertified, Mediterranean ecosystems, because they are able to reestablish functional shurblands (Francis & Thornes, 1990; Alegre et al. 2004.). On the other hand, in the extensive livestock production systems of the Mediterranean basin, woody plants are considered important contributors to grazing animal nutrition (Dumont et al., 1995; Espejo Dı´az, 1996; Papachristou et al., 1999), because they are able to grow when herbaceous plants are dormant during the summer and=or winter period, and provide green foliage rich in nutrients to animals (Papachristou & Papanastasis, 1994). Legumes are able to fix atmospheric nitrogen in association with rhizobia. This process accounts for the ability of legumes to colonize N–deficient soils, and to enhance the fertility of the soils where they grow, because of the increment of the edaphic N content (Haystead et al., 1989; Dart, 1998). Woody legumes have become an essential part in the revegetation projects because of their ecological importance (Ndiaye & Ganry, 1997; Rodrı´guez-Echeverrı´a&Pe´rez-Fernaandez, 2003), due to the biotic N fixation in the one hand, and to the occurrence of mycorrhizas in the other, which improves nutrient acquisition and help plants to become established and cope with stress (Herrera et al., 1993; Requena et al., 2001). In some cases, the relationships between woody legumes and rhizobia has been proved highly specific (Lafay & Burdon, 1998; Zahran, 2001). However in other cases, a high degree of promiscuity can be achieved by both plant and microbes (Young & Haukka, 1996; Zahran, 2001; Pe´rez-Fernaandez & Lamont, 2003). For dif- ferent herbaceous and shrubby legume taxa, nodulation with specific rhizobia and=or mycorrhizal fungi, may become essential for the settlement and growth of the plants, and therefore, if the infective rhizobia are not present in the soil, inocu- lation may be necessary (Herrera et al., 1993; Jha et al., 1995; Lal & Khanna, 1996; Gonzaalez-Andre ´s & Ortiz, 1999). For the revegetation of a degraded area, two main strategies can be used. The first one is to reintroduce the preexisting species, which is called reclamation, and the second one is to introduce species from different ecosystems, which is called rehabilitation. In the last case, new stable and sustainable ecosystems are established, although having different land uses (Herrera et al., 1993). Four legume shrubs, potentially useful for reclamation or rehabilitation of soils in seasonally dry desertification-threatened areas, have been selected for the present study: Medicago strasseri Greuter, Mattha¨s & Risse; Medicago citrina (Font Quer) Greuter; Colutea arborescens L.; and Dorycnium pentaphyllum Scop. They are well adapted to the semiarid Mediterranean climatic conditions of central Spain, characterized by dry and hot summers, and cold winters. In addition, they produce forage which is useful for animal feed during the shortage from conventional pastures. Several studies, some of them are listed in the following lines, support the mentioned properties of the studied species: M. strasseri (Alegre et al., 1994; Gonzaalez-Andres & Ceresuela, 1998), M. citrina (Font Quer) Greuter (Gonzaalez-Andres & Ceresuela, 1998), C. arborescens L. (Papanastasis et al., 1998; Papachristou et al., 1999, Papachristou, Rhizobia Nodulating Mediterranean Shrubby Legumes 309 2000), Dorycnium sp. (Davies & Lane, 2003; Wills et al., 1999). In addition, when Dorycnium pentaphyllum Scop. is included in lamb diets, egg hatching and larval development for the parasite Trhichostongylus coumbriformis, decreases due to the action of the condensed tannins (Niezen et al., 2002). The starting hypothesis for this work is that prior to revegetation of land, it is necessary to assess the specificity or promiscuity of the plant–rhizobia interaction and the occurrence of symbiotically effective rhizobia in soils in order to determine if it is necessary to inoculate with selected rhizobia strains. The starting point is the isolation of infective rhizobia strains obtained from seedlings growing in non- disturbed native areas. The general objective is to ascertain the specificity or promiscuity of the plant–rhizobia interaction for the species Medicago strasseri, M. citrina, Colutea arborescens, and Dorycnium pentaphyllum. The tasks to fulfill this objective were: (a) To test several soils, in order to detect the presence of natural populations of rhizobia capable of effectively nodulating the studied species and (b) To assess the range of rhizobia that effectively nodulates the target legume spe- cies, as well as the host range of the rhizobia isolated from those species to define cross-inoculation groups for the studied species. Material and Methods A list of the rhizobial accessions included in this research, their parent hosts and ori- gin or donor institutions is presented in Table 1. Stock cultures were maintained on yeast extract mannitol agar (YMA) medium. Isolation of Nodulating Rhizobia Strains I.MsS1, I.McSC, I.McSM, I.CA, and I.DP were isolated from root nodules of the parent hosts indicated in Table 1, from seedlings growing in a native habitat in the location cited in the column ‘‘origin.’’ An area of 4 km2 was sampled, and one plant was randomly selected from each km2. One random nodule was selected from every plant and the isolates were authenticated by inoculation of seedlings of the host species. Strains I.M3 and I.M4 were each isolated from root nodules of M. citrina formed after inoculation with a 10-fold dilution from two soils intended to be revegetated (Tables 1 and 2), following the procedure described in the next subsection. They were authenticated as well. In both cases, the nodules were surface sterilized with 0.1% HgCI2, thoroughly washed, individually crushed following the crushed nodule method (Beck et al., 1993), plated onto YMA medium containing 25 mg litreÀ1 Congo Red, and incubated at 28C for 10 days. Bacteria were isolated from a single typical colony on the assumption that this would represent the domi- nant strain in any nodule (Johnston & Beringer, 1975). Isolates were characterized by color, colony appearance, extracellular polysaccharide production (EPS), and size of the first discrete colonies appearing away from the confluent growth on YMA plates (Odee et al., 1997). Colony
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