Abstract
Novel molecular pinball machines of the plasma membrane control cytosolic Ca(2+) levels that regulate plant metabolism. The essential components involve: 1. an auxin-activated proton pump; 2. arabinogalactan glycoproteins (AGPs); 3. Ca(2+) channels; 4. auxin-efflux "PIN" proteins. Typical pinball machines release pinballs that trigger various sound and visual effects. However, in plants, "proton pinballs" eject Ca(2+) bound by paired glucuronic acid residues of numerous glycomodules in periplasmic AGP-Ca(2+). Freed Ca(2+) ions flow down the electrostatic gradient through open Ca(2+) channels into the cytosol, thus activating numerous Ca(2+)-dependent activities. Clearly, cytosolic Ca(2+) levels depend on the activity of the proton pump, the state of Ca(2+) channels and the size of the periplasmic AGP-Ca(2+) capacitor; proton pump activation is a major regulatory focal point tightly controlled by the supply of auxin. Auxin efflux carriers conveniently known as "PIN" proteins (null mutants are pin-shaped) pump auxin from cell to cell. Mechanosensitive Ca(2+) channels and their activation by reactive oxygen species (ROS) are yet another factor regulating cytosolic Ca(2+). Cell expansion also triggers proton pump/pinball activity by the mechanotransduction of wall stress via Hechtian adhesion, thus forming a Hechtian oscillator that underlies cycles of wall plasticity and oscillatory growth. Finally, the Ca(2+) homeostasis of plants depends on cell surface external storage as a source of dynamic Ca(2+), unlike the internal ER storage source of animals, where the added regulatory complexities ranging from vitamin D to parathormone contrast with the elegant simplicity of plant life. This paper summarizes a sixty-year Odyssey.