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Plant basal resistance - Universiteit Utrecht

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Chapter 1<br />

genetic resources for Arabidopsis (i.e. fully genotyped recombinant mapping populations<br />

or association mapping populations) to field experimentation. For instance, Arabidopsis<br />

mapping populations in which defence traits segregate could be grown under different<br />

field conditions with varying degrees of disease pressure by one or multiple pathogens<br />

and/or herbivores. Subsequent fitness evaluation may reveal defence-regulating QTLs that<br />

provide selective benefits under specified environmental conditions. With more and more<br />

Arabidopsis accessions being genome-sequenced, another promising approach arises from<br />

genome-wide association mapping approaches, which are based on associations between<br />

phenotypes and DNA sequence variants within individuals or isogenic populations (Nordborg<br />

and Weigel, 2008; Atwell et al., 2010). Particularly if defence phenotypes can be related to<br />

ecological stress parameters from the accessions’ geographical origins, this technique has<br />

the potential to assign measurable ecological significance to defence regulatory alleles.<br />

SECONDARY DEFENCE METABOLITES: BIOSYNTHETIC ORIGINS AND CHEMICAL<br />

CLASSIFICATION<br />

Irrespective of the type of <strong>resistance</strong> response expressed in plants, secondary metabolites<br />

are ubiquitous “tools” in plant defence. Figure 4 provides a generic overview of the<br />

biosynthesis pathways involved in the production of defence secondary compounds in plants.<br />

Unlike primary metabolites, which play a role in the process of photosynthesis, respiration,<br />

solute transport, nutrient assimilation and differentiation, secondary metabolites have no<br />

recognised role in plant processes that are essential for growth. The distribution of secondary<br />

metabolites across the plant kingdom is diverse and varies between plant species and taxa.<br />

Based on chemical structure, plant secondary metabolites can be divided into four major<br />

groups: terpenes, phenolics, nitrogen- and sulphur-containing compounds and oxyilipins.<br />

Figure 4 shows a generic overview of the different biochemical pathways controlling these<br />

plant compounds.<br />

Terpenes<br />

This class of metabolites is immensely diverse and includes more than 30,000 lipophilic<br />

compounds (Kennedy and Wightman, 2011). Their structure includes one or more 5-carbon<br />

isoprene (C 5 H 8 ) units, which are synthesized in plants by both the mevalonate and dexy-d-<br />

xylulose pathways (Rohmer, 1999). Classifications of terpenoids are based on the number<br />

of isoprene units they contain. Hemiterpenes incorporate 1 isoprene unit, monoterpenes<br />

incorporate 2 units, sesquiterpenes incorporate 3 units, diterpenes incorporate 4 units,<br />

sesterpenes comprise 5 units, triterpenes include 6 units, and tetraterpenes incorporate 8<br />

units. Terpenes exhibit a broad range of ecological roles in the plant kingdom. Their roles<br />

include antimicrobial properties, attraction of pollinator, parasitoic or predator insects, and<br />

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