Transcellular proton current in Achlya bisexualis hyphae: relationship to polarized growth.

Growing hyphae of Achlya bisexualis drive an electric current through themselves, such that positive charge flows into the apical region (the anterior 300 micron) and exits distally along the hyphal trunk. They also generate a gradient of extracellular pH, such that the medium surrounding the apex is slightly alkaline whereas that along the hyphal trunk is acid. To explore the genesis of these gradients and their relationship to polarized extension, we examined the effects of changes in the composition of the growth medium. The transcellular electric current was most pronounced in medium rich in amino acids. In leaner medium, containing limited amounts of amino acids or none at all, the current was attenuated or absent. We interpret the results to mean that inward current represents H+/amino acid symport, mediated by porters that are preferentially localized in the apical region. Apical alkalinity may be due to ammonia production. Outward current, and perhaps also the generation of metabolic acid, reflects the distribution of the H+-ATPase, which is excluded from the apex but is abundant along the hyphal trunk. Thanks to the spatial segregation of transport functions, protons characteristically flow into the apical region. However, since hyphae grow apically and at the same rate despite wide variations in current pattern, the flow of electric charge through the hyphae cannot be required to polarize extension or to localize the tip.