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Abiotic and biotic stresses and changes in the lignin ... - ResearchGate

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In red p<strong>in</strong>e (P<strong>in</strong>us res<strong>in</strong>osa), comb<strong>in</strong>ed nitrogen-phosphorus-potassium (NPK)<br />

fertilization reduced <strong>the</strong> lign<strong>in</strong> content of branches (Blodgett et al., 2005) but had an<br />

opposite effect on <strong>the</strong> ma<strong>in</strong> stem of Picea abies (Kostia<strong>in</strong>en et al., 2004).<br />

Populus plants receiv<strong>in</strong>g high concentrations of nitrogen (10 mM NH 4 NO 3 ) had<br />

less lign<strong>in</strong>, reduced β-O-4 l<strong>in</strong>kage, <strong>in</strong>creased frequency of ρ-hydroxyphenyl units <strong>and</strong> a<br />

tendency toward low S/G ratios (Pitre et al., 2007).<br />

At <strong>the</strong> molecular level, <strong>the</strong> expression of <strong>the</strong> pot171 gene, which encodes a<br />

prote<strong>in</strong> similar to CCoAOMT, is negatively regulated <strong>in</strong> response to nitrogen <strong>in</strong> Populus<br />

trichocarpa x deltoid hybrids (Cooke et al., 2003).<br />

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Calcium (Ca): The effect of Ca on <strong>the</strong> biosyn<strong>the</strong>sis of phenylpropanoids is not<br />

sufficiently clear. Some studies have shown that Ca <strong>in</strong>creases <strong>the</strong> activity of guaiacol-<br />

POD <strong>and</strong> PAL (Castañeda <strong>and</strong> Pérez, 1996; Kolupaev et al., 2005); however, a<br />

reduction <strong>in</strong> <strong>the</strong> amount of phenolic compounds has also been reported (de Obeso et al.,<br />

2003). O<strong>the</strong>r authors observed a reduction <strong>in</strong> PAL <strong>and</strong> POD activity (soluble <strong>and</strong> cell<br />

wall-bound), as well as reductions <strong>in</strong> <strong>the</strong> levels of phenolic compounds, <strong>in</strong>clud<strong>in</strong>g lign<strong>in</strong><br />

(Teixeira et al., 2006). No substantial <strong>changes</strong> <strong>in</strong> <strong>the</strong> amounts of <strong>the</strong>se compounds was<br />

observed (Tomas-Barberan et al., 1997).<br />

In Populus tremula x Populus tremuloides, Ca deficiency was responsible for<br />

wood deformation, which <strong>in</strong>cluded smaller diameter of xylem vessels, shorter fibre<br />

length <strong>and</strong> reduced levels of S units <strong>in</strong> lign<strong>in</strong> (Lautner et al., 2007). Therefore, it seems<br />

that Ca is necessary for <strong>the</strong> biosyn<strong>the</strong>sis of lign<strong>in</strong> <strong>in</strong> plants. It is important to note that<br />

several PODs are l<strong>in</strong>ked to <strong>the</strong> galacturonic doma<strong>in</strong>s of pect<strong>in</strong> <strong>in</strong> <strong>the</strong> presence of Ca<br />

(Penel <strong>and</strong> Grepp<strong>in</strong>, 1996) <strong>and</strong> such a l<strong>in</strong>k occurs only <strong>in</strong> <strong>the</strong> pect<strong>in</strong> cha<strong>in</strong> that is crossl<strong>in</strong>ked<br />

to Ca, known as Ca-pectates (Carp<strong>in</strong> et al., 2001). S<strong>in</strong>ce <strong>the</strong> middle lamella <strong>and</strong>

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