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Disease cycle of potato late blight - MSpace at the University of ...

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from HMG CoA by a reaction c<strong>at</strong>alyzed by <strong>the</strong> enzyme 3-hydroxy-3-methylglutaryl-CoA<br />

reductase (HMGR). This enzyme is encoded by a small family <strong>of</strong> genes with different<br />

p<strong>at</strong>tern <strong>of</strong> expression, ranging from constitutive to tissue-specific induction. After two<br />

remaining consecutive steps IPP is released and isopentenyl diphosph<strong>at</strong>e (DMAPP) is<br />

formed by <strong>the</strong> isomeriz<strong>at</strong>ion <strong>of</strong> IPP (Rodriguez-Concepcion and Boron<strong>at</strong> 2002;<br />

Eisenreich et al. 2004; Phillips et al. 2008) (Fig. 1.3).<br />

The DOXP-MEP p<strong>at</strong>hway, also called <strong>the</strong> non-mevalon<strong>at</strong>e route, consists <strong>of</strong> eight<br />

reactions c<strong>at</strong>alyzed by nine enzymes, seven <strong>of</strong> which are characterized structurally<br />

(Hunter et al. 2003). Briefly, 1-deoxy-D-xylulose 5-phosph<strong>at</strong>e (DXP) obtained by<br />

condens<strong>at</strong>ion <strong>of</strong> pyruv<strong>at</strong>e and D-glyceraldehyde 3-phosph<strong>at</strong>e (GAP) in a reaction<br />

c<strong>at</strong>alyzed by <strong>the</strong> enzyme 1-deoxy-D-xylulose 5-phosph<strong>at</strong>e synthase (DXS) undergoes a<br />

reorganiz<strong>at</strong>ion associ<strong>at</strong>ed with a reduction step. The next step <strong>of</strong> <strong>the</strong> DOXP-MEP<br />

p<strong>at</strong>hway is <strong>the</strong> conversion and reduction <strong>of</strong> DXP to 2C-methyl-D-erythritol 4-phosph<strong>at</strong>e<br />

(MEP) by <strong>the</strong> enzyme 1-deoxy-D-xylulose 5-phosph<strong>at</strong>e reductoisomerase (DXR). MEP is<br />

subsequently transformed to IPP and DMAPP by five-independent steps by <strong>the</strong> action <strong>of</strong><br />

<strong>the</strong> enzymes 2-C-methyl-D-erythritol 4-phosph<strong>at</strong>e cytidylyltransferase (MCT), 4-<br />

(cytidine 50-diphospho)-2-Cmethyl-D-erythritol kinase (CMK) and 2-C-methyl-D-<br />

erythritol 2,4-cyclodiphosph<strong>at</strong>e synthase (MDS). In two final steps c<strong>at</strong>alyzed by (E)-4-<br />

hydroxy-3-methylbut-2-enyl diphosph<strong>at</strong>e synthase (HDS) and reductase (HDR), IPP and<br />

DMAPP are formed (Rodriguez-Concepcion and Boron<strong>at</strong> 2002; Eisenreich et al. 2004;<br />

Phillips et al. 2008) (Fig. 1.4).<br />

17

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