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20 Mycorrhizal Fungi and Plant Growth Promoting Rhizobacteria 355<br />

al. 2001). The main activity of these bacteria is associated with the production<br />

of auxin-type phytohormones (Dobbelaere et al. 1999). The production and<br />

significance of auxins have been investigated at the molecular level (van de<br />

Broek et al. 1999; Lambrecht et al. 2000).<br />

Several systems for studying rhizosphere colonization by PGPR have been<br />

proposed, the one described by Simons et al. (1996) appears interesting, not<br />

only for testing the PGPR ability, but also to investigate PGPR/<strong>plant</strong> root interactions.<br />

It is known that several bacterial cell <strong>surface</strong> properties could be involved<br />

in the adhesion of PGPR to roots that may be either nonspecific based on electrostatic<br />

forces, or involve specific recognition between the <strong>surface</strong>s. In this<br />

context, root <strong>surface</strong> glyco-proteins and several different bacterial exo-polysaccharides<br />

could be involved (Weller and Thomashow 1994). In some PGPR,<br />

fimbriae (pili) may function in the adherence of cells to roots, but the contribution<br />

of flagella to the colonization process apparently depends on the PGPR<br />

strain, <strong>plant</strong> species and type of soil (Kloepper 1994).A sugar-binding protein<br />

system has been described in Azospirillum related to chemostaxis and root<br />

colonization (van Bastelaere et al. 1999).<br />

Novel techniques for microbial community fingerprinting are being developed<br />

(Kozdroj and van Elsas, 2000), and these are being adapted for the<br />

genetic characterization of PGPR. For example, the approach used by Zinniel<br />

et al. (2002), based on the 16S rRNA gene amplification and sequencing, has<br />

been proposed for the genetic characterization of endophytic bacteria, while<br />

Bertrand et al. (2001) identify PGPR 16S rDNA sequence analysis. The molecular<br />

bases of rhizosphere colonization have recently been reviewed (Lugtenberg<br />

et al. 2001). Siciliano and Germida (1998) proposed the use of BIOLOG<br />

analysis and fatty acid methyl ester profiles to study PGPR behavior after<br />

inoculation, particularly the effects on root-associated microbial populations.<br />

Confocal laser scanning microscopy is being used for studying the<br />

microorganisms/<strong>plant</strong> root interactions, for example, to detect a currently<br />

used marker such as the green fluorescent protein (Lagopodi et al. 2002) or to<br />

localize colonizing bacteria by fluorescence and in situ hybridization, after<br />

staining with the fluorescent Live/Dead BacLight dye (Bianciotto et al. 2000,<br />

2001).<br />

The molecular bases of the biocontrol ability of these rhizobacteria have<br />

been investigated in the last few years (Keel et al. 1992; O’Gara et al. 1994;<br />

Cook et al. 1995; Tomashow and Weller 1995; Chin-A-Woeng et al. 2001;<br />

Moenne-Loccos et al. 2001), and systemic induced resistance has been argued<br />

as a mechanism of disease suppression by endophytes (Duijff et al. 1998), or<br />

other PGPR (Defago and Keel 1995; Chin-A-Woeng et al. 2001).

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