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

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194<br />

K. Janardhan Reddy<br />

endoplasmic reticulum <strong>and</strong> vacuoles. In the cytosol, calcium concentration is very low<br />

<strong>and</strong> maintained in the range <strong>of</strong> 0.1-0.2 µM (Evans et al., 1991). Calcium is bound as<br />

calcium pectate in the middle lamella <strong>and</strong> is essential for strengthening <strong>of</strong> cell walls <strong>and</strong><br />

tissues in plants. Calcium is also required for root elongation. Calcium stabilizes cell<br />

membranes by bridging phosphate <strong>and</strong> carboxyl groups <strong>of</strong> phospholipids (Cladwell<br />

<strong>and</strong> Haug, 1981) <strong>and</strong> proteins (Legge et al., 1982) in the membrane. When calcium<br />

supply is low, 50% <strong>of</strong> total calcium can be bound as pectate in tomato (Armstrong <strong>and</strong><br />

Kirkby, 1979b). Calcium stimulates α-amylase activity in germinating seeds (Bush et al.,<br />

1986).<br />

In the cytosol, calcium binding proteins known as calcium modulated proteins<br />

like calmodulin (Snedden <strong>and</strong> Fromm, 2001), calmodulin binding proteins (Reddy et al.,<br />

2002) <strong>and</strong> calcium dependent but calmodulin independent protein kinases are the targets<br />

<strong>of</strong> calcium signals (Roberts <strong>and</strong> Harmon, 1992; Harmon et al., 2001). Calmodulin is an<br />

ubiquitous protein present universally in all eukaryotic cells. It is a polypeptide with<br />

148 amino acids <strong>and</strong> four binding sites for calcium <strong>and</strong> it is heat stable <strong>and</strong> insensitive<br />

to pH changes. Calmodulin activates a number <strong>of</strong> enzymes like NAD kinase, adenylate<br />

cyclase <strong>and</strong> membrane bound ATPase.<br />

Calcium requirement is lower in monocots than in dicots. For example 2.5 µM<br />

supply <strong>of</strong> calcium in rye grass <strong>and</strong> 100 µM <strong>of</strong> calcium in tomato are required for maximum<br />

growth (Table 5).<br />

Table 5. Effect <strong>of</strong> calcium on relative growth rates <strong>and</strong><br />

calcium content in shoots <strong>of</strong> Rye grass <strong>and</strong> tomato*<br />

Calcium conc. Calcium content (mg/gm dry wt.) Relative growth rate<br />

(µM) Rye grass Tomato Rye grass Tomato<br />

0.8 0.6 2.1 42 03<br />

2.5 0.7 1.3 100 19<br />

10 1.5 3.0 94 52<br />

100 1.7 12.9 94 100<br />

*Adapted from Lonergan <strong>and</strong> Snowball (1969)<br />

Calcium deficiency is very rare if the pH is maintained properly. The deficiency<br />

symptoms first appear in growing tips <strong>and</strong> young leaves because <strong>of</strong> low mobility <strong>of</strong><br />

Ca 2+ . The tips <strong>and</strong> young young leaves become chlorotic followed by necrosis <strong>of</strong> leaf<br />

margins. Calcium deficiency induces premature shedding <strong>of</strong> fruit <strong>and</strong> buds. Excess<br />

calcium in the growth medium interferes with Mg 2+ absorption. High Ca 2+ usually causes<br />

alkaline pH which in turn precipitates many <strong>of</strong> the micronutrients making them unavailable<br />

to plants.

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