The reduction in growth of maize (Zea mays L.) seedling primary roots induced by salinization of the nutrient medium with 100 millimolar NaCI was accompanied by reductions in the length of the root tip elongation zone, the length of fully elongated epidermal cells, and the apparent rate of cell production: Each was partially restored when calcium levels in the salinized growth medium were increased from 0.5 to 10.0 millimolar. We investigated the possibility that the inhibition of elongation growth by salinity might be associated with an inhibition of cell wall acidification, such as that which occurs when root growth is inhibited by IAA. A qualitative assay of root surface acidification, using bromocresol purple pH indicator in agar, showed that salinized roots, with and without extra calcium, produced a zone of surface acidification which was similar to that produced by control roots. The zone of acidification began 1 to 2 millimeters behind the tip and coincided with the zone of cell elongation. The remainder of the root alkalinized its surface. Kinetics of surface acidification were assayed quantitatively by placing a flat tipped pH electrode in contact with the elongation zone. The pH at the epidermal surfaces of roots grown either with 100 millimolar NaCI (growth inhibitory), or with 10 millimolar calcium ± NaCI (little growth inhibition), declined from 6.0 to 5.1 over 30 minutes. We conclude that NaCI did not inhibit growth by reducing the capacity of epidermal cells to acidify their walls.
Half maximal inhibition of sodium ('Na ) influx into maize (Zea mays L.) root segments incubated in solutions containing from 0.25 to 100 millimolar NaCI was consistently attained with external calcium activity at 0.26 ± 0.10 millimolar. Sodium ions do not appear to compete with calcium during initial binding to sites on the plasma membrane that participate in the regulation of sodium influx under saline conditions.Increasing the activity of calcium ions in the saline root media of glycophytic crop plants can reduce sodium accumulation and partially reverse the growth inhibition induced by excess salinity in the rhizosphere (2,4,5,8,15 (2) suggested that this could be the primary root response to salt stress. The displacement ofthese essential calcium ions by sodium would disrupt membrane stability thus accelerating efflux of intracellular solutes and influx of toxic sodium from the rhizosphere; additional calcium ions in the rhizosphere would reverse these processes by competing more effectively with sodium ions for calcium binding sites on the plasma membrane (2, 8, 9). Direct support for this hypothesis was initially provided by observations of NaCl induced displacement of surface Ca2" from intact cotton root hairs and maize root protoplasts, as visualized using microfluorometric techniques (2, 9). However, these techniques may be prone to interpretation errors and a later report suggested that the predominant effect of external salinization on calcium associated fluorescence in maize protoplasts was due to the release of calcium from intracellular membranes rather than its displacement from the plasma membrane surface (10). .N.). 331 An alternative method for investigating salinity effects on sodium influx into roots and its relation to calcium binding at the plasma membrane is described here. We assume that if sodium ions are able to competitively displace calcium from plasma membrane binding sites associated with the regulation of sodium influx, then the external calcium activity required to inhibit sodium influx should increase as external salinity is increased. If, alternatively, external salinity does not competitively displace calcium, then the calcium activity required to inhibit sodium influx should remain constant as external salinity is increased. To determine which mechanism is operative, we investigated the effects of a range of external concentrations of NaCl (0.25-100 mM) on rates on sodium (22Na) influx into washed maize root segments. Sodium influx at each NaCl concentration was assayed at several different calcium concentrations from 0 to 10 mM. Salinization of solutions increases their ionic strength. Such increases in ionic strength can progressively reduce the effective concentration and the physiologically effective chemical activity of calcium ions, through ion to ion interactions (1,3,6). Thus, instead of relating sodium influx to total calcium concentrations, we investigated the relationship between sodium influx and the chemical activity of the calcium ions in each of the...
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