11.03.2015 Views

Physiology and Molecular Biology of Stress ... - KHAM PHA MOI

Physiology and Molecular Biology of Stress ... - KHAM PHA MOI

Physiology and Molecular Biology of Stress ... - KHAM PHA MOI

SHOW MORE
SHOW LESS

You also want an ePaper? Increase the reach of your titles

YUMPU automatically turns print PDFs into web optimized ePapers that Google loves.

Nutrient <strong>Stress</strong><br />

201<br />

11. COPPER<br />

Copper is a transition element <strong>and</strong> forms stable complexes. It occurs in two oxidation<br />

states (Cu + <strong>and</strong> Cu 2+ ). Divalent form <strong>of</strong> copper is readily reduced to monovalent copper<br />

which is unstable. Copper occurs in soils mainly as a divalent cation. Copper is not<br />

easily mobile in plants but it is known to move from older to younger leaves <strong>and</strong> this<br />

movement depends on copper status <strong>of</strong> the plant.<br />

Copper is an important redox component directly related to electron transfer<br />

reactions in photosynthesis, respiration, lignification <strong>of</strong> cell walls <strong>and</strong> detoxification <strong>of</strong><br />

superoxide radicals (Fox <strong>and</strong> Guerinot, 1998). Copper is associated with three different<br />

forms <strong>of</strong> proteins (S<strong>and</strong>mann <strong>and</strong> Boger, 1983) which include Blue proteins without<br />

oxidase activity (eg. Plastocyanin), Non-blue proteins which oxidize monophenols<br />

<strong>and</strong> Multi-copper proteins, containing around four copper atoms per molecule. (eg.<br />

Ascorbic acid oxidase <strong>and</strong> diphenol oxidase). Polyphenol oxidases are involved in the<br />

synthesis <strong>of</strong> lignin <strong>and</strong> alkaloids. Copper-deficient leaves <strong>of</strong> subterranean clover showed<br />

lower levels <strong>of</strong> polyphenol oxidase activity (Delhaize et al., 1985).<br />

Nearly 50% <strong>of</strong> the copper localized in chloroplasts is associated with plastocyanin.<br />

Chlorophyll <strong>and</strong> plastocyanin levels <strong>and</strong> certain enzyme activities were more<br />

with the increased copper content in dry matter <strong>of</strong> Pisum sativum (Table 11)<br />

Table 11. Relationship between copper content, some constituents <strong>of</strong> chloroplast<br />

<strong>and</strong> activities <strong>of</strong> certain <strong>of</strong> certain enzymes in Pisum sativum leaves*<br />

Copper Chlorophyll Plastocyanin Diamine Ascorbate Superoxide<br />

(mg/g dry (m moles/g (n moles/m mole oxidase oxidase dismutase<br />

wt) dry wt) chlorophyll activity (m activity activity<br />

mole/g (enzyme<br />

protein/hr)<br />

units/mg<br />

protein)<br />

2.2 4.4 0.3 0.24 220 3.6<br />

3.8 3.9 1.1 0.43 470 13.5<br />

6.9 4.9 2.4 0.86 730 22.9<br />

* Adapted from Ayala <strong>and</strong> S<strong>and</strong>mann (1988).<br />

Davies et al. (1978) observed that copper deficiency decreased the activities<br />

<strong>of</strong> polyphenol oxidase, IAA oxidase <strong>and</strong> peroxidase. Flowering was also affected by<br />

copper deficiency (Table 12).

Hooray! Your file is uploaded and ready to be published.

Saved successfully!

Ooh no, something went wrong!