(Questions)(n) on phloem biology. 1. Electropotential waves, Ca2+ fluxes and cellular cascades along the propagation pathway

Abstract

This review explores the relationships between electrical long-distance signalling, Ca(2+) influx coincident with propagation of electropotential waves, and cellular responses to Ca(2+) influx including the consequences for sieve-tube conductivity and mass flow. Ca(2+) influx is inherent to electropotential waves and appears to constitute the key link between rapid physical signals and resultant chemical cascades in sieve tubes and adjacent cells. Members of several channel groups are likely involved the regulation of Ca(2+) levels in sieve elements. Among them are hyperpolarization-activated, depolarization-activated, and mechanosensitive Ca(2+) channels located in the plasma membrane and Ca(2+) dependent Ca(2+) channels that reside in ER-membranes of sieve elements. These channels collectively determine intracellular Ca(2+) levels in sieve elements and their neighbour cells. The latter cells react to Ca(2+) elevation by inducing diverse functional responses dependent on the cell type. If the Ca(2+) concentration in sieve elements surpasses a threshold level, dual sieve-plate occlusion by proteins and callose deposition is triggered. Occlusion is reversed when Ca(2+) levels subside. Electrical messages may regulate the degree of sieve plate hydraulic conductivity in intact plants by partial sieve-plate occlusion that has a major impact on volume flow through sieve tubes. Furthermore, complete but temporary occlusion of sieve tubes may modify mass flow patterns in intact plants.