Keywords: Carbon partitioning, magnesium deficiency, photooxidation, phloem transport, root growth.
Magnesium (Mg) deficiency exerts a major influence on the partitioning of dry matter and carbohydrates between shoot and roots. One of the very early reactions of plants to Mg deficiency stress is the marked increase in the shoot-to-root dry weight ratio which is associated with a massive accumulation of carbohydrates in source leaves, especially of sucrose and starch. These higher concentrations of carbohydrates in Mg-deficient leaves together with the accompanying increase in shoot-to-root dry weight ratio are indicative of a severe impairment in phloem export of photoassimilates from source leaves. Studies with common bean and sugar beet plants have shown that Mg plays a fundamental role in phloem loading of sucrose. At a very early stage of Mg deficiency, phloem export of sucrose is severely impaired, an effect which occurs before any noticeable changes in shoot growth, chlorophyll concentration or photosynthetic activity. These findings suggest that accumulation of carbohydrates in Mg-deficient leaves is caused directly by Mg deficiency stress and not as a consequence of reduced sink activity. The role of Mg in the phloem loading process seems to be specific; resupplying Mg for 12 or 24 h to Mg-deficient plants resulted in a very rapid recovery of sucrose export. It appears that the massive accumulation of carbohydrates and related impairment in photosynthetic CO2 fixation in Mg deficient leaves causes an over-reduction in the photosynthetic e-transport chain that potentiates the generation of highly reactive O2 species (ROS). Plants respond to Mg deficiency stress by marked increases in antioxidative capacity of leaves, especially under high light intensity, suggesting that ROS generation is stimulated by Mg deficiency in chloroplasts. Accordingly, it has been found that Mg deficient plants are very susceptible to high light intensity. Exposure of Mg-deficient plants to high light intensity rapidly induced leaf chlorosis and necrosis, an outcome which was effectively delayed by partial shading of the leaf blade, even though the Mg concentrations in different parts of the leaf blade were unaffected by shading. The results indicate that photooxidative damage contributes to development of leaf chlorosis under Mg deficiency, suggesting that plants under high light conditions have a higher physiological requirement for Mg. Maintenance of a high Mg nutritional status of plants is, thus, essential in the avoidance of ROS generation which occurs at the expense of inhibited phloem export of sugars and impairment of CO2 fixation and, particularly under high light conditions.
Ismail Cakmak1 and Ernest A Kirkby2.
1 Sabanc? University, Faculty of Engineering and Natural Sciences, 34956 Istanbul, Turkey.
2 Institute of Integrative and Comparative Biology, Faculty of Biology, University of Leeds, Leeds LS2 9JT, UK.
24 pages, 8 figures, 1 table, 75 references.