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20 Mycorrhizal Fungi and Plant Growth Promoting Rhizobacteria 359<br />

8 Interactions Involved in Nutrient Cycling and Plant<br />

Growth Promotion<br />

It is well known that soil microorganisms are able to change the bioavailability<br />

of mineral <strong>plant</strong> nutrients, and this ability has been shown by soil bacteria<br />

probed to have PGPR activity and, in some cases, be used as <strong>plant</strong> inoculants<br />

(Barea et al. 1997; Probanza et al. 2002). Several experiments have described<br />

the improvement of <strong>plant</strong> growth and nutrition by means of synergistic interactions<br />

between PGPR and AM fungi (Barea 2000). It has also been suggested<br />

that a certain level of selectivity (“specificity”) is involved in these interactions<br />

(Azcón 1989). From the perspective of sustainability, the re-establishment<br />

of nutrient cycles after any process of soil degradation is of interest, as<br />

is the understanding of the microbial interactions responsible for the subsequent<br />

management of such natural resources, either for a low-input agricultural<br />

technology (Bethlenfalvay and Linderman 1992; Gianinazzi and<br />

Schüepp 1994; Jeffries and Barea 2001), or for the re-establishment of the natural<br />

vegetation in a degraded area (Miller and Jastrow 1994, 2000; Requena et<br />

al. 2001). Most of the information on this topic concerns N and P cycling<br />

(Barea 2000).<br />

In spite of this, Rhizobium sp. (general term) are not considered among the<br />

PGPR types; due to the relevance of their interaction with AM fungi, it would<br />

be interesting to make some comments on this. A great deal of work has been<br />

carried out on the tripartite symbiosis legume (general term) – mycorrhiza-<br />

Rhizobium (Azcón-Aguilar and Barea 1992; Barea et al. 1992; Barea 2000). The<br />

inoculation of AM fungi has been shown to improve nodulation and N 2 fixation.<br />

Using the isotope 15 N has made it possible to ascertain and quantify the<br />

amount of N which is actually fixed in a particular situation, and measure the<br />

contribution of the AM symbiosis to the process (Barea et al. 1992). The physiological<br />

and biochemical mechanisms underlying the AM fungi x Rhizobium<br />

interactions to improve legume productivity have also been discussed. In<br />

spite of the main AM effect in enhancing Rhizobium activity mediated by a<br />

generalized stimulation of host nutrition, more localized effects may occur at<br />

the root or nodule level (Barea et al. 1992). Interactions can also take place at<br />

either the pre-colonization stages, when both microorganisms interact as rhizosphere<br />

inhabitants, or during the development of the tripartite symbiosis<br />

(Azcón-Aguilar and Barea 1992). The influence of host and/or bacterial genotypes<br />

in these interactions has also been discussed, suggesting a certain level<br />

of specificity (Azcón et al. 1991; Ruiz-Lozano and Azcón 1993; Monzón and<br />

Azcón 1996).<br />

Multimicrobial interactions including AM fungi, Rhizobium sp. and PGPR<br />

have also been tested (Requena et al. 1997). Target microorganisms were isolated<br />

from a representative area of a desertification-threatened semi-arid<br />

ecosystem in the south-east of Spain. Microbial isolates were characterized<br />

and screened for effectiveness in soil microcosms. Anthyllis cytisoides L., an

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