The role of electrical phenomena in tip growth, with special reference to the developmental plasticity of filamentous fern gametophytes.

Cell expansion in many plant structures, including algal rhizoids, fungal hyphae, root hairs, and pollen tubes, is restricted to their apical tips. Endogenous electric fields are seen to accompany polarized growth in all tip-growing cells studied to date. The extensive studies on absorptive tip-growing structures have established that positive currents enter their elongating tips, with a portion of the entry current being carried by a localized calcium influx into the extreme tip. The resulting tip-to-base gradient in calcium concentration appears to be responsible for maintaining polarized growth in these systems, although it is uncertain whether this calcium effect is mediated via either electrophoretic or cytoskeletal mechanisms. In contrast, the few electrical measurements made on photosynthetic cells suggest that the orientation of their transcellular fields is transiently or permanently reversed relative to the fields in absorptive structures. In darkness, microelectrode measurements indicate that the apical tip of the fern filament is 5 mV electronegative relative to the base of the apical cell. This cellular dipole is perceived with the vibrating probe as a focused outward current that departs from the tip region and a more diffuse inward current that enters the lateral sides of the apical cell. The tip current is predominantly composed of protons, as can be identified with various cation-selective electrodes. This proton current is thought to help maintain localized wall expansion in the filament tip. Blue light mediates the major morphogenetic transition in fern gametophytes, i.e. the transition from the tip-growing filament to the planar prothallus. All the above electrical and ionic parameters change in the few minutes of irradiation before the filament tip starts lateral swelling. The plasma membrane at the extreme tip begins to hyperpolarize within 3 s, while the basal region shows a delayed, but greater response. The cellular dipole that had existed in darkness is thus abolished in 10 to 15 min after the start of irradiation. With the vibrating probe a more diffuse pattern of positive currents is observed to emerge from the tip as well as the subapical regions of the apical cell. Simultaneously, proton efflux increases in the subapical region; the resulting decrease in cell wall pH should help plasticize the lateral walls, which may, in turn, facilitate the process of lateral swelling over the next few hours.(ABSTRACT TRUNCATED AT 400 WORDS)