Osmoregulation and contractile vacuoles of protozoa

Protozoa living in fresh water are subjected to a hypotonic environment. Water flows across their plasma membrane since their cytosol is always hypertonic to the environment. Many wall-less protozoa have an organelle, the contractile vacuole complex (CVC), that collects and expels excess water. Recent progress shows that most, if not all, CVCs are composed of a two-compartment system encircled by two differentiated membranes. One membrane, which is often divided into numerous vesicles and tubules, contains many proton-translocating V-ATPase enzymes that provide an electrochemical gradient of protons and which fuses only with the membrane of the second compartment. The membrane of the second compartment lacks V-ATPase holoenzymes, expands into a reservoir for fluid storage, and is capable of fusing with the plasma membrane. It is this second compartment that periodically undergoes rounding ("contraction"), setting the stage for fluid expulsion. Rounding is accompanied by increased membrane tension. We review the current state of knowledge on osmolarity, ion concentrations, membrane permeability, and electrophysiological parameters of cells and their contractile vacuoles, where these criteria are helpful to our understanding of the function of the CVC. Effects of environmental stresses on the CVC function are also summarized. Finally, other functions suggested for CVCs based on molecular and physiological studies are reviewed.