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Plant basal resistance - Universiteit Utrecht

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121<br />

General discussion<br />

There are different ways by which rhizobacteria can protect plants. These include<br />

competition for nutrients (Kamilova et al., 2005), siderophore-mediated competition for iron<br />

(Schippers et al., 1987), signal interference (Lin et al., 2003), antibiosis (Haas and Keel, 2003)<br />

and ISR (Zamioudis and Pieterse, 2011). Interestingly, transcriptome analysis of DIMBOA-<br />

exposed P. putida revealed enhanced expression of the phzF gene (Chapter 4, Table I),<br />

which encodes an enzyme in the biosynthesis of the broad-spectrum antibiotic phenazine<br />

(Blankenfeldt et al., 2004). These results suggest that DIMBOA exudation from cereal roots<br />

has the potential to boost phenazine production by natural Pseudomonas bacteria in the<br />

rhizosphere, which has been linked to suppression of take-all disease (Thomashow and<br />

Weller, 1988). Moreover, preliminary evidence suggests that root colonization by P. putida<br />

KT2440 primes emission of wounding-inducible volatiles aboveground (Figure 3). These<br />

volatiles have well-known functions in the recruitment of natural enemies of herbivores<br />

(Turlings and Ton, 2006), and some can also serve as long-distance signals to prime defence<br />

in neighbouring plants (Heil and Ton, 2008). The study described in Chapter 4 demonstrates<br />

that BXs can act as belowground signals to recruit and select for plant-beneficial P. putida<br />

bacteria. This discovery is exploitable for plant breeder and biotech companies to sustainably<br />

manage pests and diseases by selecting for cereal crops that exude increased amounts<br />

of DIMBOA from their roots. Furthermore, it was recently reported that mycorrhization<br />

increases DIMBOA levels in maize roots (Song et al., 2011). It is known that mycorrhization<br />

causes major changes in the rhizobacterial community (Linderman, 1988). Considering<br />

the results in Chapter 4, DIMBOA may be a driving factor behind this plant-beneficial<br />

mycorrhizoshere effect.<br />

Classical breeding strategies can be used to select varieties with an enhanced<br />

capacity to exudate DIMBOA in the roots. If these breeding schemes can be combined with<br />

the selection for increased BX content aboveground, these varieties will not only be able to<br />

better in recruiting a disease-suppressing rhizosphere, but they will also be more resistant<br />

to pests and diseases aboveground. Less targeted approaches can be used to identify similar<br />

traits in non-cereal crops. With the next-generation sequencing technologies becoming<br />

more cost-efficient, it will become possible to carry out large-scale QTL analyses in order<br />

to associate crop performance in terms of growth and stress <strong>resistance</strong> with metagenomic<br />

profiles in the rhizosphere (Poland et al., 2011; Bisseling et al., 2009). Such an approach will<br />

lead to the identification of novel plant genes that promote the establishment of a disease-<br />

suppressing and growth-promoting rhizosphere.<br />

AKNOWLEDGEMENTS<br />

We thank Georg Jander and Adewale Martins Adio from the Boyce Thompson Institute<br />

for <strong>Plant</strong> Research (Cornell University; New Yourk, United States of America) for analysing

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