NATURE OF THE ACTION CURRENT IN NITELLA : VI. SIMPLE AND COMPLEX ACTION PATTERNS

The experiments indicate that the protoplasm of Nitella consists of an aqueous layer W with an outer non-aqueous surface layer X and an inner non-aqueous surface layer Y. The potential at Y is measured by the magnitude of the action curve and the potential at X by the distance from the top of the action curve to the zero line. These potentials appear to be due chiefly to diffusion potentials caused by the activity gradients of KCl across the non-aqueous layers X and Y. The relative mobilities of K(+) and Cl(-) in X and in Y can be computed and an estimate of the activity of KCl in W can be made. In the complete resting state the mobilities of K(+) and Cl(-) in X are not very different from those in Y. The action curve is due to changes in Y which suddenly becomes very permeable, allowing potassium to move from the sap across Y into W, and thus losing its potential. A gradual loss may be due to changes in ionic mobility in Y. When recovery is incomplete and Y has not yet regained its normal potential a stimulus may cause a loss of the potential at Y giving an action curve of small magnitude. The magnitude may vary in successive action curves giving what is called a complex pattern in contrast to the simple pattern observed when recovery is complete and all the action curves are alike. Complex patterns occur chiefly in cells treated with reagents. Untreated cells usually give simple patterns. A variety of complex action patterns is discussed. It is evident that the cells of Nitella show much more variation than such highly specialized cells as muscle and nerve which give stereotyped responses. In some cases it may be doubtful whether the all-or-none law holds.