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Plant basal resistance - Universiteit Utrecht

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111<br />

General discussion<br />

2 of this thesis shows natural variation between Arabidopsis accessions in the sensitivity of<br />

jasmonic acid (JA)- and SA-inducible marker genes, pointing to variation in responsiveness of<br />

relatively late-acting post-invasive defence barriers. The same set of accessions also showed<br />

natural variation in responsiveness of chitosan-induced callose deposition, suggesting<br />

variation in responsiveness of early-acting post-invasive defence barriers. Interestingly,<br />

there was a negative correlation between the responsiveness of early-acting defences<br />

(callose) and that of late-acting defences (SA-induced PR-1 gene induction). Although a<br />

sample size of six accessions is too small to reach definite conclusions about the ecological<br />

meaning of these findings, it is tempting to speculate that some of these accessions have<br />

de-sensitised their early defences in order to interact with plant beneficial microbes, such<br />

as plant-growth promoting rhizobacteria (PGPRs). In order to compensate for this immuno-<br />

compromised defence layer, these accessions have primed later-acting defences. Further<br />

experiments should be performed to test this hypothesis. Of the possibilities to consider, a<br />

HapMap population of 360 Arabidopsis accessions could be screened for PGPR colonization<br />

and PAMP-induced callose deposition in the roots (Buckler and Gore, 2007). Identification of<br />

genes promoting PGPR colonization but suppressing PAMP-induced callose would confirm<br />

this hypothesis.<br />

CYP81D11: A SUPPRESSOR OF EARLY ACTING POST-INVASIVE DEFENCE?<br />

The QTL analysis described in Chapter 2 identified two loci regulating PAMP-induced callose<br />

deposition. One major QTL was found on chromosome III, whereas a much weaker QTL was<br />

identified at the top of chromosome I. The direction of both QTLs show opposite effects: the<br />

major QTL at chromosome III mediates a suppressive effect from Bur-0 parent and stimulatory<br />

effect from Col-0 parent, whereas the weaker QTL at chromosome I mediates a stimulatory<br />

effect from Bur-0 parent and suppressive effect from Col-0 parent. Although both QTLs still<br />

spans a multitude of genes, it is interesting that the major locus on chromosome III maps<br />

onto the cis-jasmone inducible gene CYP81D11 (At3g28740). This gene was previously linked<br />

to the regulation of indirect plant defences against insects (Bruce et al., 2008; Matthes et<br />

al., 2010). Its role in direct plant defences, however, remains unclear. To further investigate<br />

a possible function of CYP81D11 in callose deposition, a T-DNA insertion mutant and a<br />

CYP81D11 over-expression line in the genetic background of Col-0 (line CYP81D11OE 5-5.3;<br />

Bruce et al., 2008; Matthes et al., 2010), were tested for flg22-induced callose deposition,<br />

as described by Luna et al., (2011). Interestingly, this experiment revealed that CYP81D11OE<br />

plants are severity affected in PAMP-induced callose deposition (Figure 1).

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