The membrane potentials of aged, excised barley (Hordeum vulgare L.) root cells were rapidly depolarized by the addition of salicylic acid (o-hydroxybenzoic acid) The phenomenon of allelopathy, in which one species of plant may dramatically inhibit the development of others by producing and releasing specific compounds into the environment, is well documented (18). In a few cases, the allelopathic compounds and their mode of transmission have been elucidated (11,12). In a large number of reported cases, however, the modes of action of the allelopathic compounds (allelochemicals) are not understood. This paper reports further investigations into the mode of action of an important group of allelochemicals, phenolic acids.Earlier papers by Glass (3)(4)(5) have established that one ubiquitous group of phenolics, namely benzoic and cinnamic acids, are potent inhibitors of active K and Pi absorption by excised barley roots. The effect is readily reversible and the inhibitory capacity of the benzoic acids is strongly correlated with their lipid solubilities. From these results it was suggested that phenolics act directly on the cell membrane, modifying its permeability and thereby increasing the rate of efflux of ions. To test this hypothesis the effect of phenolics on cell membrane electrical potentials has been studied. According to the Goldman (7) where E is the electropotential difference (PD) between o, the outer solution and i, the inner solution; R, the gas constant (joules degree mole); T, the absolute temperature; F, the Faraday (coulombs equivalent-'); P subscript is the permeability coefficient of a univalent cation, j+, or a univalent anion, j-; C,+°, the concentration of j+ outside, and Cj,+, the concentration of j+ inside. Predictions of E based only upon K+ and Clcharacteristics, derived from the above equation, may under certain conditions be remarkably consistent with measured values for root cell membrane potentials (14). Clearly any alterations of the permeability properties of the cell membrane with regard to K+ or Cl-will be reflected by changes in E. If the cell membrane became extremely permeable, the imbalance of ion concentration across the cell membrane, which is necessary for E, would disappear and E would fall to near zero.
MATERIALS AND METHODSSeedlings of Hordeum vulgare L. cv. Karlsberg were grown for 3 or 4 days in aerated 5 X 10-' M CaSO, solution following a 1-day germination in aerated distilled water. At the end of this period, the last 1 to 1.5 cm of roots were excised from the root tip and allowed to age in aerated 5 X 10-M CaSO4 solution. It was found that the aging process produced a very significant increase in the cell membrane potential of the order reported by Pitman et al. (14). Membrane potentials of cells in the root tips were measured with microcapillary electrodes using the apparatus described by Dunlop and Bowling (2). During determinations of membrane potentials the root tips were held in a 5 ml Plexiglas chamber which, under control conditions, contained a ...