Propagation of Action Potential over the Trap-lobes of Aldrovanda vesiculosa

In the trap-lobes of Aldrovanda vesiculosa, an action potentialwas generated in a cell located at the base of a sensory hairstanding on the margin of central portion of the paired lobes,which spread over this portion within about 40 msec. The electricalcoupling ratio for two adjacent cells in the middle layer ofthe lobes was 0.8. This showed that an action potential generatedin a cell of the trap-lobes must spread electrotonically toadjacent cells. An analysis of an equivalent circuit for thecell injected with current and its neighboring cells in themiddle layer of the lobes showed that the resistances of theplasmalemma, tonoplast and junction between two cells were 11.1,4.7 and 0.56 M, respectively. Numerous plasmodesmata in thejunctional walls of the cells were found by electron microscopy.The low resistance of the junction between cells must be dueto the presence of the plasmodesmata which allows an electrotonictransmission of action potential from cell to cell. (Received January 23, 1982; Accepted April 13, 1982)     

Membrane Potentials in Excitable Cells of Aldrovanda vesiculosa Trap-Lobes

The resting membrane potential in excitable cells of Aldrovandatrap-lobes is composed of diffusion and electrogenic potentials.The diffusion potential, about –100 mV in artificial pondwater, was determined from the external K⁺ and Na⁺ concentrations.The permeability ratio, P_Na/P_K of the membrane was estimatedto be about 0.3. The electrogenic potential hyperpolarized themembrane to about –140 mV. The peak value of the actionpotential increased by +26 mV with a tenfold increase in theexternal Ca²⁺ concentration. The action potential was blockedby an application of the Ca²⁺ chelater or the Ca channel blocker,LaCl₃. Cells showed additional Ca²⁺ influx (7.8 pmole/cm² impulse)during membrane excitation. These facts suggest that the transientincrease in Ca²⁺ influx causes the action potential presentin cells of Aldrovanda trap-lobes. ¹ Present address: Jerry Lewis Neuromuscular Research Center,School of Medicine, University of California Los Angeles, LosAngeles, CA90024, U.S.A. ² Present address: Biological Laboratory, Kyoritsu Women's University,Hachioji 193, Japan. (Received September 21, 1983; Accepted September 7, 1984)     

Movements of K+ during Shutting and Opening of the Trap-Lobes in Aldrovanda vesiculosa

K⁺ movements during the shutting and subsequent opening of trap-lobesin Aldrovanda vesiculosa were measured using ⁸⁶Rb as a tracerfor K⁺. Immediately after the shutting, a large amount of ⁸⁶Rbpre-loaded in the trap-lobes was detected in the hollow spaceinside the shut trap. This may indicate that much of the K⁺in the active motor cells leaks out during the shutting, resultingin turgor loss in the cells. ⁸⁶Rb(K⁺) uptake in the trap wasactive. During the opening process, enhanced ⁸⁶Rb uptake wasobserved. The time course of this uptake was similar to thatof the opening of the trap-lobes, and both courses were acceleratedby IAA. Enhanced K⁺ uptake may restore the turgor in activemotor cells. The quantity of K⁺ that moved during the shuttingor opening was estimated as 20% of that in the active motorcells in the open state of the trap-lobes. The K⁺ efflux acrossthe membranes of the active motor cells may be caused by a largeincrease in bulk flow triggered by an action potential, andwas estimated as 6,200 pmol.cm^–2. ¹ This paper is dedicated to the memory of Professor Joji Ashidawho established the physiology of rapid movement in Aldrovandavesiculosa. (Received July 22, 1982; Accepted November 11, 1982)     

Ecophysiological Characterization of Dormancy States in Turions of the Aquatic Carnivorous Plant Aldrovanda vesiculosa

Two main dormancy states, innate and imposed dormancy, were characterized in turions (winter buds) of the aquatic carnivorous plant Aldrovanda vesiculosa L. (Droseraceae) kept at 3 ± 1 °C in a refrigerator over the winter. As a result of the breaking of imposed dormancy by a temperature increase (at 15 – 20 °C), some of the turions rose to the water surface within 1 – 3 d and germinated. Turion leaves contained large lacunae with a slimy reticulum and were filled by water over winter. As a result of breaking imposed dormancy, the proportion of gas volume in inner turion leaves rose from 10 – 20 % to 100 % of leaf lacunae volume. The aerobic dark respiration rate of the turions [0.74 – 1.5 μmol O₂) kg⁻¹(FM) s⁻¹] slightly increased during innate dormancy after 1 – 2 d at 20 °C, while it was almost constant during the breaking of imposed dormancy. The anaerobic fermentation rate of the turions was only 1.5 – 7 % of the oxygen respiration rate and also was constant during the breaking of imposed dormancy. In turions, the content of glucose, fructose, and sucrose was the same for the two states of dormancy, but starch content was greatly reduced for the imposed dormancy (10 – 11 vs. 32 % DM). It may be suggested that a temperature increase causes an increase of fermentation or respiration which is responsible for the evolution of gas in turion lacunae and, thus, for turion rising.     

Seed reproductive biology of the rare aquatic carnivorous plant Aldrovanda vesiculosa (Droseraceae)

Despite the unprecedented global decline in extant populations of Aldrovanda vesiculosa in the last century, little is known about the reproductive biology of this iconic aquatic carnivorous plant. This study aimed to investigate the role of seed‐based reproduction in the ecology of A. vesiculosa, with particular focus on the interplay between the regulation of seed dormancy by temperature cues and the efficacy of exogenous ethylene gas to act as a germination stimulant, the desiccation capacity and long‐term storage potential of seeds for conservation purposes. Sexual reproduction appears to be extremely limited in both natural and naturalized populations across three continents, with high variability in the success of flowering and seed set between sites and between seasons. Overall, flowering yielded few fertile fruit (6–19% of flowers producing fertile fruit) and seed viability was variable but generally low (29–88%). Fecundity appears to be influenced by seasonal climatic conditions and microhabitat characteristics. Aldrovanda vesiculosa possesses physiologically dormant seeds, with germination stimulated by exposure to ethylene gas (>90% germination) at 25 °C. Seeds appear sensitive to desiccation and sub‐zero temperature storage, with no germination and markedly reduced embryo growth after storage of seeds for >1 month at 15 °C and 15% relative humidity or after short‐term (24 h) storage at ?18 °C. In the absence of significant conservation and restoration initiatives, the continuing decline of dystrophic freshwater wetland habitats globally leaves A. vesiculosa facing extinction. As the successful long‐term storage of seeds appears unfeasible based on the approaches described in this study, other alternatives for germplasm conservation such as cryostorage of vegetative tissues or zygotic embryos must be considered for establishing long‐term ex situ collections of critical germplasm.     

Field growth characteristics of two aquatic carnivorous plants,Aldrovanda vesiculosa andUtricularia australis

Basic growth characteristics of two species of free-floating submerged carnivorous plants, the very rare and stenotopicAldrovanda vesiculosa and the very common and eurytopicUtricularia australis, were investigated in a 10/11-day field growth experiment within three nylon enclosures at two artificialAldrovanda sites in the T?eboň region, S Bohemia, Czech Republic, at the peak of a growing season. Growth ofAldrovanda was best at a meso-eutrophic site (biomass doubling time,T ₂, 8.4–10.7 days, mean growth of new leaf whorls 0.96 whorls days^?1, 1.6 developed branches per shoot) and slower at an oligo-mesotrophic site (T ₂ 17.2–21.5 days, growth of whorls 1.01 whorls days^?1, 0.1–0.5 branches per shoot). Growth ofUtricularia was similar at both sites (T ₂ 19.8–33.2 days or 9.1–16.8 days, growth of whorls 3.1 or 2.7 whorls days^?1, 1.5–2.1 or 0.8–1.4 developed branches per shoot at the former or latter site, respectively). Throughout the experiment, both species at the meso-eutrophic site allocated relatively more biomass to the production and growth of branches, than to that of new whorls. The results show thatAldrovanda, although usually considered as competitively weaker, can grow faster during the growing season peak thanUtricularia due to frequent branching and the subsequent rapid growth and separation of daughter shoots. Very rapid growth of rootless aquatic carnivorous plants in nutrient-poor habitats allows the consideration of ecophysiological adaptations that enable the plants to gain limiting mineral nutrients. These adaptations include carnivory, efficient nutrient reutilization from senescent shoots, and very high affinity for mineral nutrient uptake from water. Comparison of growth rates of rare and stenotopicA. vesiculosa and very common and eurytopicU. australis shows that differences in their rarity do not seem to be based on differences of growth rate.     

Population ecology of the endangered aquatic carnivorous macrophyte Aldrovanda vesiculosa at a naturalised site in North America

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The aquatic carnivorous plant Aldrovanda vesiculosa (Droseraceae) exhibits altered developmental stages in male gametophyte

Protoplasma - Aldrovanda vesiculosa (Droseraceae) is a rare aquatic carnivorous plant, distributed in Europe, Asia, Africa, and Australia. Aldrovanda populations can flower prolifically under...     

Action Potential Initiation in the Hodgkin-Huxley Model

A recent paper of B. Naundorf et al. described an intriguing negative correlation between variability of the onset potential at which an action potential occurs (the onset span) and the rapidity of action potential initiation (the onset rapidity). This correlation was demonstrated in numerical simulations of the Hodgkin-Huxley model. Due to this antagonism, it is argued that Hodgkin-Huxley-type models are unable to explain action potential initiation observed in cortical neurons in vivo or in vitro. Here we apply a method from theoretical physics to derive an analytical characterization of this problem. We analytically compute the probability distribution of onset potentials and analytically derive the inverse relationship between onset span and onset rapidity. We find that the relationship between onset span and onset rapidity depends on the level of synaptic background activity. Hence we are able to elucidate the regions of parameter space for which the Hodgkin-Huxley model is able to accurately describe the behavior of this system.     

Tonoplast Action Potential of Characeae

The plasmalemma action potential was found to be indispensableto the production of the tonoplast action potential. In a solutionlacking Ca²⁺ and containing other divalent cations such as Ba²⁺,Mg²⁺ or Mn²⁺, the plasmalemma excited in Nitella but did notin Chara. In Nitella, however, both the tonoplast action potentialand EC-coupling were abolished due to depletion of Ca²⁺ fromthe external medium. Ca²⁺ ions injected into the cytoplasmiclayer caused a transient change in both plasmalemma and tonoplastpotentials. These results suggest that a transient rise in Ca²⁺concentration during excitation of the plasmalemma may triggerthe tonoplast action potential. (Received February 14, 1986; Accepted August 29, 1986)     

Some Relations between Action Potential and Resting Potential of the Lobster Giant Axon

Experiments were performed to determine the quantitative relation existing between action potential and resting potential of the lobster giant axon. Resting potential changes were induced by either increasing the external potassium concentration or by reducing the external calcium concentration. For either treatment the action potential amplitude is proportional to the logarithm of the resting potential minus a constant. This constant is equivalent to the minimum resting potential at which a propagated spike is possible, and is larger for depolarization in low calcium than in high potassium. Thus the change in action potential per unit change in resting potential is greater in low external calcium than in high external potassium. Analog computer solutions to the Hodgkin-Huxley equations for squid axon membrane potentials show that, if the initial conditions are properly specified, the action potential is proportional to the logarithm of the potassium potential minus a constant. The experimental results and the analog computations suggest that reducing external calcium produces changes in the invertebrate axon that cannot be accounted for solely on the basis of alterations in the potassium potential.     

豚鼠离体腹腔神经节神经元自发f-EPSP和AP

应用细胞内生物电记录技术观察豚鼠腹腔神经节(CG)神经元自发快兴奋性突触后电位(f-EPSP)和动作电位(AP)的特征,分析其形成的可能机制。发现在豚鼠离体CG上存在自发f-EPSP和AP,发放频率不同。自发f-EPSP的幅度(5.67±2.66)mV(n=26),明显低于刺激内脏大神经诱发f-EPSP的幅度(13.26±6.74)mV(n=34,P<0.01),而自发的AP后超极化幅度(AHPA)(13.86±4.24)mV(n=30),明显高于刺激内脏大神经诱发的AHPA(8.99±2.79)mV(n=54,P<0.01)。六烃季铵或低Ca2 /高Mg2 Krebs液能完全阻断自发的AP,但自发的f-EPSP则不被完全阻断。结果提示豚鼠离体CG神经元有自发性电活动,这除与突触前膜ACh的随机释放有关以外,可能还有对Ca2 不敏感的其他递质介导。     

Quantification of Transmembrane Currents during Action Potential Propagation in the Heart

The measurement, quantitative analysis, theory, and mathematical modeling of transmembrane potential and currents have been an integral part of the field of electrophysiology since its inception. Biophysical modeling of action potential propagation begins with detailed ionic current models for a patch of membrane within a distributed cable model. Voltage-clamp techniques have revolutionized clinical electrophysiology via the characterization of the transmembrane current gating variables; however, this kinetic information alone is insufficient to accurately represent propagation. Other factors, including channel density, membrane area, surface/volume ratio, axial conductivities, etc., are also crucial determinants of transmembrane currents in multicellular tissue but are extremely difficult to measure. Here, we provide, to our knowledge, a novel analytical approach to compute transmembrane currents directly from experimental data, which involves high-temporal (200 kHz) recordings of intra- and extracellular potential with glass microelectrodes from the epicardial surface of isolated rabbit hearts during propagation. We show for the first time, to our knowledge, that during stable planar propagation the biphasic total transmembrane current (I_m) dipole density during depolarization was ∼0.25 ms in duration and asymmetric in amplitude (peak outward current was ∼95 μA/cm² and peak inward current was ∼140 μA/cm²), and the peak inward ionic current (I_ion) during depolarization was ∼260 μA/cm² with duration of ∼1.0 ms. Simulations of stable propagation using the ionic current versus transmembrane potential relationship fit from the experimental data reproduced these values better than traditional ionic models. During ventricular fibrillation, peak I_m was decreased by 50% and peak I_ion was decreased by 70%. Our results provide, to our knowledge, novel quantitative information that complements voltage- and patch-clamp data.     

A Threshold Equation for Action Potential Initiation

In central neurons, the threshold for spike initiation can depend on the stimulus and varies between cells and between recording sites in a given cell, but it is unclear what mechanisms underlie this variability. Properties of ionic channels are likely to play a role in threshold modulation. We examined in models the influence of Na channel activation, inactivation, slow voltage-gated channels and synaptic conductances on spike threshold. We propose a threshold equation which quantifies the contribution of all these mechanisms. It provides an instantaneous time-varying value of the threshold, which applies to neurons with fluctuating inputs. We deduce a differential equation for the threshold, similar to the equations of gating variables in the Hodgkin-Huxley formalism, which describes how the spike threshold varies with the membrane potential, depending on channel properties. We find that spike threshold depends logarithmically on Na channel density, and that Na channel inactivation and K channels can dynamically modulate it in an adaptive way: the threshold increases with membrane potential and after every action potential. Our equation was validated with simulations of a previously published multicompartemental model of spike initiation. Finally, we observed that threshold variability in models depends crucially on the shape of the Na activation function near spike initiation (about −55 mV), while its parameters are adjusted near half-activation voltage (about −30 mV), which might explain why many models exhibit little threshold variability, contrary to experimental observations. We conclude that ionic channels can account for large variations in spike threshold.     

Cell Surface Deformation during an Action Potential

The excitation of many cells and tissues is associated with cell mechanical changes. The evidence presented herein corroborates that single cells deform during an action potential. It is demonstrated that excitation of plant cells (Chara braunii internodes) is accompanied by out-of-plane displacements of the cell surface in the micrometer range (～1–10 μm). The onset of cellular deformation coincides with the depolarization phase of the action potential. The mechanical pulse: 1) propagates with the same velocity as the electrical pulse (within experimental accuracy, ～10 mm s^?1), 2) is reversible, 3) in most cases is of biphasic nature (109 out of 152 experiments), and 4) is presumably independent of actin-myosin-motility. The existence of transient mechanical changes in the cell cortex is confirmed by micropipette aspiration experiments. A theoretical analysis demonstrates that this observation can be explained by a reversible change in the mechanical properties of the cell surface (transmembrane pressure, surface tension, and bending rigidity). Taken together, these findings contribute to the ongoing debate about the physical nature of cellular excitability.