Using a highly sensitive vibrating electrode, the pattern of naturally occurring electric currents around I-day-old primary roots of Lepidium sativum L. growing vertically downward and the current pattern following gravistimulation of the root has been examined. A more or less symmetrical pattern of current was found around vertically oriented, downward growing roots. Current entered the root at the root cap, the meristem, and the beginning of the elongation zone and left the root along most of the elongation zone and in the root hair zone. After the root was tilted to a horizontal position, we observed current flowing acropetally at the upper side of the root cap and basipetally at the lower side within about 30 seconds in most cases. After a delay of several minutes, acropetally oriented current was also found flowing along the upper side of the meristematic zone. The apparent density of the acropetal current in the root cap region increased and then decreased with time. Gravitropic curvature was first visible approximately 10 minutes after tilting of the root to the horizontal position. Since the change in the pattern of current in the root cap region precedes bending of the root and is different for the upper and lower side, a close connection is suggested between the current and the transduction of information from the root cap to the elongation zone following graviperception in the cap.Measurements with an extracellular vibrating electrode have revealed characteristic patterns of self-generated electric current traversing individual plant and animal cells (5,6,19,21). Growing pollen tubes and root hairs, for example, produce an electric current which enters at the growing tip and leaves at the basal, nongrowing region of the cell (19,22). Electric currents and fields have also been observed around entire organs, such as roots, hypocotyls, and coleoptiles (10-12, 17, 20). Suggested roles for these currents in roots are in ion uptake (13) segments tilted to the horizontal position were reported to undergo a change in surface potential within as short a time as 2 to 3 mi (17).We shall attempt to show that a vertical, freely growing root has a steady pattern of current and that the pattern changes when the root is placed horizontally. Such a change suggests that the current could be involved in the transduction of gravitropic stirnulus. MATERIALS AND METHODSGrowth Conditions and Holding Apparatus for Seedlings. Seeds of Lepidium sativum L. were soaked 30 min in tap water and then placed on vertical moist filter paper in a closed container. The seeds were oriented with their micropyles down so that the roots would not curve during germination. After 20 h at a temperature of 23 + 2°C, the primary roots were 6.0 ± 1.2 mm long. Single seeds were then transferred and fastened with a drop ofwarm agar medium to a small L-shaped Plexiglas holder which, after a further 3.5 h, was clamped in a micromanipulator, avoiding as much as possible any shaking of the seed. Using the micromanipulator, the root was placed...

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Membrane potentials were measured in lateral statocytes of vertically and nonvertically growing roots of Lepidium sativum L. using conventional glass-microelectrode techniques. Statocytes in vertically growing roots showed a stable resting potential of-118±5.9 mV without spontaneous fluctuations. Upon tilting the root 45° from the vertical, an electrical asymmetry was observed. Statocytes on the physically lower side of the root depolarized by approx. 25 mV. This depolarization occurred following a latent period of 8 s reaching a minimum (approx.-93 mV) after 170 s. This depolarization is the earliest event in graviperception ever recorded. After this depolarization, the cell repolarized within 60 s to a potential approx. 10 mV more positive than the original resting potential. Statocytes on the upper flank showed a slow hyperpolarization (t 1/2h=half time for hyperpolarization=168 s) reaching a final, stable potential at a level 10 mV more negative. These effects of gravistimulation were statenchyma-specific, since cells in the cortex and rhizodermis showed no similar effects. The gravi-electrical responses were observed in 25% of all roots tested. Roots which showed no gravi-electrical response had a reduced elongation growth, lacked gravity-induced bending and lacked the typical structural polarity in punctured statocytes. This observed transition from a symmetrical pattern of resting potential in the statenchyma to an asymmetrical pattern following gravistimulation supports the results observed with external current measurements (Behrens et al., Plant Physiol. 70, 1079-1083, 1982) and extends these results to the cellular level and to considerably improved temporal resolution. The asymmetry in the gravi-electrical response extends the graviperception model of Sievers and Volkmann (Planta 102, 160-172, 1972) which comprises an asymmetrical sedimentation of the amyloplasts on the distal endoplasmic reticulum of statocytes. This generates an intraorgan signal which then must be transmitted to the growth zone.

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Electrical transmembrane potential differences and resistances in different tissues of intact root tips of Lepidium sativum L. were investigated in a humid atmosphere by conventional glass-microelectrode techniques with the reference electrode at the surface (apoplast) of the root. The resting potential (inside negative) in cells of the root cap rose from-80 mV in external cell layers (secretion cells) to approx.-140 mV in central cells (statocytes). Measurements of the electric input resistance within the apoplast of the root tip (calyptra, meristem and elongation zone) yielded a preference for longitudinal contact (resistance per length of tissue approx. 3.4 GOhm m(-1)) compared with transversal contact (approx. 14 GOhm m(-1)). Similarly, the symplastic coupling expressed as the characteristic length (L) where a signal is reduced to 1/c compared with the origin yielded L y =390 μm in the longitudinal (y) direction and L x =140 μm in the transversal (x) direction. Cable analytical treatment of the symplastic input resistances (approx. 10 MOhm) resulted in low membrane resistances in the y-direction at the ends of cells compared with the membrane resistances in the x-direction (approx. 0.2 Ohm m(2)) of the lateral membranes in the approximately cylindrical cells. This anisotropy is discussed in terms of model calculations. The resistivity of the symplast was calculated to be about 2.5 Ohm m. The input current-voltage relationship displayed a slight curvature with increasing slope for the more negative membrane potential typical of membranes with electrogenic pumps. Even after massive electrical stimulation in the range from-50 to-150mV carried out to trace current-voltage curves, electrical excitations (action potentials) were not detected in the cells investigated.