Mimosa pudica: Electrical and mechanical stimulation of plant movements

Thigmonastic movements in the sensitive plant Mimosa pudica L., associated with fast responses to environmental stimuli, appear to be regulated through electrical and chemical signal transductions. The thigmonastic responses of M. pudica can be considered in three stages: stimulus perception, electrical signal transmission and induction of mechanical, hydrodynamical and biochemical responses. We investigated the mechanical movements of the pinnae and petioles in M. pudica induced by the electrical stimulation of a pulvinus, petiole, secondary pulvinus or pinna by a low electrical voltage and charge. The threshold value was 1.3–1.5 V of applied voltage and 2 to 10 µC of charge for the closing of the pinnules. Both voltage and electrical charge are responsible for the electro‐stimulated closing of a leaf. The mechanism behind closing the leaf in M. pudica is discussed. The hydroelastic curvature mechanism closely describes the kinetics of M. pudica leaf movements.     

Mechanism of the Seismonastic Reaction in Mimosa pudica

The efflux of K⁺ from the pulvinar cells of Mimosa pudica was shown to increase substantially during the seismonastic reaction. This result is shown to indicate a decrease in σ (reflection coefficient) of pulvinar cell membrane for potassium salts which could account for the pulvinar cell turgor decrease during the seismonastic reaction.     

Molecular electronics in pinnae of Mimosa pudica

Bioelectrochemical circuits operate in all plants including the sensitive plant Mimosa pudica Linn. The activation of biologically closed circuits with voltage gated ion channels can lead to various mechanical, hydrodynamical, physiological, biochemical and biophysical responses. Here the biologically closed electrochemical circuit in pinnae of Mimosa pudica is analyzed using the charged capacitor method for electrostimulation at different voltages. Also the equivalent electrical scheme of electrical signal transduction inside the plant''s pinna is evaluated. These circuits remain linear at small potentials not exceeding 0.5 V. At higher potentials the circuits become strongly non-linear pointing to the opening of ion channels in plant tissues. Changing the polarity of electrodes leads to a strong rectification effect and to different kinetics of a capacitor. These effects can be caused by a redistribution of K⁺, Cl⁻, Ca²⁺ and H⁺ ions through voltage gated ion channels. The electrical properties of Mimosa pudica were investigated and equivalent electrical circuits within the pinnae were proposed to explain the experimental data.Key words: electrophysiology, plant cell electrostimulation, charged capacitor method, electrical circuits, electrical signaling, Mimosa pudica     

On the Phloem of Mimosa pudica L.

The phloem of Mimosa pudica L. was examined in view of somereports that the sieve elements in this plant show featuresnot previously described for these cells in Leguminosae. Inthe present study only a usual dicotyledonous type of sieveelement was recognized. The sieve elements pass through stagesof differentiation involving development and dispersal of P-proteinbodies, disintegration of nuclei, and appearance of plastidsstoring a starch staining red with iodine. Callose occurs onthe transverse or moderately oblique sieve plates. The phloemcontains secretory cells. They are wide and long and have transverseor more or less inclined end walls. In younger cells the endwall bears in the centre a conspicuous pit to which the protoplastsof the superimposed cells are firmly attached. In many oldercells the pit region is replaced by a perforation so that thecontents of superimposed cells may be completely merged. Itremains to be determined whether the perforation is presentin an intact plant or results from a rupture during sampling.The secretory cells accumulate material that gives a positivetest for carbohydrates and a negative test for proteins.     

Mechanical and electrical anisotropy in Mimosa pudica pulvini

Thigmonastic or seismonastic movements in Mimosa pudica, such as the response to touch, appear to be regulated by electrical, hydrodynamical and chemical signal transduction. The pulvinus of Mimosa pudica shows elastic properties, and we found that electrically or mechanically induced movements of the petiole were accompanied by a change of the pulvinus shape. As the petiole falls, the volume of the lower part of the pulvinus decreases and the volume of the upper part increases due to the redistribution of water between the upper and lower parts of the pulvinus. This hydroelastic process is reversible. During the relaxation of the petiole, the volume of the lower part of the pulvinus increases and the volume of the upper part decreases. Redistribution of ions between the upper and lower parts of a pulvinus causes fast transport of water through aquaporins and causes a fast change in the volume of the motor cells. Here, the biologically closed electrochemical circuits in electrically and mechanically anisotropic pulvini of Mimosa pudica are analyzed using the charged capacitor method for electrostimulation at different voltages. Changing the polarity of electrodes leads to a strong rectification effect in a pulvinus and to different kinetics of a capacitor discharge if the applied initial voltage is 0.5 V or higher. The electrical properties of Mimosa pudica''s pulvini were investigated and the equivalent electrical circuit within the pulvinus was proposed to explain the experimental data. The detailed mechanism of seismonastic movements in Mimosa pudica is discussed.Key words: electrophysiology, plant electrostimulation, pulvinus, Mimosa pudica, charged capacitor method, electrical circuits, ion channels     

Mimosa pudica may possess antidepressant actions in the rat.

In Mexico, aqueous extracts from dried leaves of Mimosa puolica are employed to alleviate depression. In this study, the behavioral actions of aqueous extracts of M. pudica at various concentrations were tested. Rats having received saline (0.9%; 0.30 ml; I.P.), clomipramine, desipramine or several dosages of aqueous extracts from M. pudica (ml = 2.0 mg/kg; m2 = 4.0 mg/kg; m3 = 6.0 mg/kg; m4 = 8.0 mg/kg) during a 30-day period were submitted to the forced swimming test and to the test for differential reinforcement of low rates of response at 72 sec (DRL-72s). Any possible anxiolytic action resulting from several doses (ml = 2.0 mg/kg; m2 = 4.0 mg/kg; m3 = 6.0 mg/kg; m4 = 8.0 mg/kg) of extracts of M. pudica were compared with those caused by diazepam (1.3 mg/kg, I.P.) in the elevated plus-maze test. Results showed that clomipramine (1.25 mg/kg, I.P.), desipramine (2.14 mg/kg, I.P.) and M. pudica (6.0 mg/kg and 8.0 mg/kg, I.P.) reduced immobility in the forced swimming test and increased the rate of reinforcers received in the DRL-72s test; these data suggest that M. pudica produces antidepressant effects in the rat. Diazepam increased the open-arms exploration time in the elevated plus-maze test, but M. pudica did not show any comparable action at any tested dose. M. pudica therefore produced an antide-pressant-like profile similar to two tricyclic antidepressants.     

Molecular characterization of cytosolic cysteine synthase in Mimosa pudica

Journal of Plant Research - In the cysteine and mimosine biosynthesis process, O-acetyl-l-serine (OAS) is the common substrate. In the presence of O-acetylserine (thiol) lyase (OASTL, cysteine...     

Structural and ultrastructural characteristics of the vascular apparatus of the sensitive plant (Mimosa pudica L.)

Summary 1. In motor organs ofMimosa pudica xylem contains living fibriform elements limited by a thick lignified highly pitted wall, whereas in other parts of the plant (stem, petiole, rachis), xylem and protoxylem vessels are closely associated with parenchyma cells which possess wall ingrowths. These ingrowths, at the apex of which the plasmalemma and the tonoplast touch, are localized like those of transfer cells of C type described byGunning andPate. Nevertheless, xylem parenchyma cells differ from cells of C type in several characteristics. Moreover, in motor organs, phloem contains cells characterized by wall ingrowths, less abundant on the parts adjacent to the sieve tubes; these cells which are localized near collenchyma cells of primary phloem, look like transfer cells of A type defined byGunning andPate; they are absent from internodes, petioles and rachides. 2. In motor organs, three types of vascular cells (companion cells, living xylem fibriform elements and protoxylem parenchyma cells) are characterized by reduced vacuolar volumes and well developed membrane systems, as compared with homologuous cells belonging to other parts of the plant. 3. A symplastic continuity holds from the middle of motor organs to their cortex: it is provided by the presence, in xylem and phloem respectively, of living fibriform elements and collenchyma cells bearing numerous pit fields containing large numbers of plasmodesmata. Several ultrastructural features suggest that the vascular apparatus ofMimosa pudica would be the site of intensive lateral transfer at different levels, specially in motor organs. Possible functions of certain structures observed are discussed in relation to some hypotheses relative to excitatory conduction pathways.     

Laser-Interferometric Re-examination of Rapid Conductance of Excitation in Mimosa pudica

The correlation between electrical excitation and turgor changesin plants has been investigated with a novel combination ofan electrometer and a laser-interferometer. With a resolutionof about 10 nm, no significant correlation could be detectedbetween the passage of action potentials and physical movementin the excitation conducting stem of Mimosa pudica. Apart frommarginal observations, the results render an hydraulic conductanceof excitation unlikely; they rather confirm the primary roleof electrical events in rapid conductance of excitation in higherplants. Key words: Laser-interferometer, turgor movement, action potential, osmotic relations     

Transport processes in stimulated and non-stimulated leaves of Mimosa pudica

Summary Using energy-dispersive X-ray microanalysis, the concentrations of ions, especially potassium and chlorine, were determined in different tissues of primary and tertiary pulvini of Mimosa pudica. It was shown that stimulating the leaf was followed by ion displacements which were most striking in the outer extensor cells, resulting in turgor loss. Since Ca concentration remains relatively constant in cell walls of collapsed cells, the changes of K concentration are best described by the K:Ca ratio. After stimulation the K:Ca ratio dropped in the outer extensor of the primary pulvinus from 775.3 to 2.37 in the cytoplasm, and from 542.2 to 9.25 in the cell wall. Changes in chlorine content were less striking in the primary pulvinus. The KCl ratios in some cases were lower than 1.0, which indicates that Cl content can increase, while K content is diminished. In the non-stimulated tertiary pulvini the outer extensor cells show high concentrations of Cl, but much lower Cl concentrations were found after stimulation. In contrast to the primary pulvinus the K content of the tertiary pulvini is very low. In the vascular tissues of both primary and tertiary pulvini stimulation is followed by a release of K and Cl out of the sieve element cytoplasm into the apoplast. K then appears accumulated in the cell walls of the collenchymatous tissue. These displacements lead to the assumption that the collenchymatous apoplast temporarily functions as a reservoir for K and to a lesser extent for Cl. With regard to the mechanism of leaf movement after stimulation, the accumulation of ions in the apoplast seems to be initiated by the decrease of water potential triggered by an apoplastic accumulation of unloaded sucrose (Fromm and Eschrich 1988a). The resulting turgor release in the outer extensor is accompanied by an efflux of ions.Supported by the Deutsche Forschungsgemeinschaft     

Calmodulin Implication in the Glycine Uptake by Mimosa pudica Pulvinar Cells

Previous experiments have evidenced that calcium is functionallyimplicated in glycine uptake by pulvinar motor cells of Mimosapudica L. The present data show that compounds having anticalmodulinproperties, compound 48/80 and the sulfonamide W-7, inhibitedthe amino acid uptake suggesting that a step in this processmay be regulated by calmodulin. H⁺ excretion by the tissuesand transmembrane potential of the motor cells were not modifiedby these compounds, thus showing that the inhibition of aminoacid uptake was not an indirect consequence of a decrease inthe proton motive force energizing the glycine H⁺ cotransport.Therefore, the data argue for the implication of calmodulinin a specific Ca²⁺-regulated reaction. (Received March 2, 1994; Accepted May 6, 1994)     

Change in the actin cytoskeleton during seismonastic movement of Mimosa pudica

The seismonastic movement of Mimosa pudica is triggered by a sudden loss of turgor pressure. In the present study, we compared the cell cytoskeleton by immunofluorescence analysis before and after movement, and the effects of actin- and microtubule-targeted drugs were examined by injecting them into the cut pulvinus. We found that fragmentation of actin filaments and microtubules occurs during bending, although the actin cytoskeleton, but not the microtubules, was involved in regulation of the movement. Transmission electron microscopy revealed that actin cables became loose after the bending. We injected phosphatase inhibitors into the severed pulvinus to examine the effects of such inhibitors on the actin cytoskeleton. We found that changes in actin isoforms, fragmentation of actin filaments and the bending movement were all inhibited after injection of a tyrosine phosphatase inhibitor. We thus propose that the phosphorylation status of actin at tyrosine residues affects the dynamic reorganization of actin filaments and causes seismonastic movement.     

Memory elements in the electrical network of Mimosa pudica L.

The fourth basic circuit element, a memristor, is a resistor with memory that was postulated by Chua in 1971. Here we found that memristors exist in vivo. The electrostimulation of the Mimosa pudica by bipolar sinusoidal or triangle periodic waves induce electrical responses with fingerprints of memristors. Uncouplers carbonylcyanide-3-chlorophenylhydrazone and carbonylcyanide-4-trifluoromethoxy-phenyl hydrazone decrease the amplitude of electrical responses at low and high frequencies of bipolar sinusoidal or triangle periodic electrostimulating waves. Memristive behavior of an electrical network in the Mimosa pudica is linked to the properties of voltage gated ion channels: the channel blocker TEACl reduces the electric response to a conventional resistor. Our results demonstrate that a voltage gated K⁺ channel in the excitable tissue of plants has properties of a memristor. The discovery of memristors in plants creates a new direction in the modeling and understanding of electrical phenomena in plants.     

Signal transduction in Mimosa pudica: biologically closed electrical circuits

Biologically closed electrical circuits operate over large distances in biological tissues. The activation of such circuits can lead to various physiological and biophysical responses. Here, we analyse the biologically closed electrical circuits of the sensitive plant Mimosa pudica Linn. using electrostimulation of a petiole or pulvinus by the charged capacitor method, and evaluate the equivalent electrical scheme of electrical signal transduction inside the plant. The discharge of a 100 µF capacitor in the pulvinus resulted in the downward fall of the petiole in a few seconds, if the capacitor was charged beforehand by a 1.5 V power supply. Upon disconnection of the capacitor from Ag/AgCl electrodes, the petiole slowly relaxed to the initial position. The electrical properties of the M. pudica were investigated, and an equivalent electrical circuit was proposed that explains the experimental data.     

Light- and Auxin-Induced Leaflet Opening in Detached Pinnae of Mimosa pudica

Leaflet pairs from detached pinnae of Mimosa pudica opened afterthe pinnae had been irradiated with light (2 W·m^–2)of 726 or 403 nm, whereas they remained almost closed with lightof 585 or 656 nm. Light-induced leaflet opening was observedonly in the daytime, from 6:00 to 16:00. Application of IAAat more than 30 µg/ml to the cut end of the pinna rachisesmade the leaflets open even in darkness with almost constantlag times of about 100 min which were independent of the concentration.NAA and 2,4-D also made the leaflet open at lower concentrationsthan IAA. Auxin-induced leaflet opening showed diurnal variation.Application of IAA for 2 to 6 min, depending on the concentration,was enough to open the leaflets. Autoradiography showed thatIAA was transferred from the cut end of a rachis throughouta pinna within 4 min. ¹Present address: Biological Institute, Faculty of Science,Kobe University, Kobe 657, Japan. (Received September 24, 1982; Accepted March 4, 1983)     

Mechanosensitivity below Ground: Touch-Sensitive Smell-Producing Roots in the Shy Plant Mimosa pudica

The roots of the shy plant Mimosa pudica emit a cocktail of small organic and inorganic sulfur compounds and reactive intermediates into the environment, including SO₂, methanesulfinic acid, pyruvic acid, lactic acid, ethanesulfinic acid, propanesulfenic acid, 2-aminothiophenol, S-propyl propane 1-thiosulfinate, phenothiazine, and thioformaldehyde, an elusive and highly unstable compound that, to our knowledge, has never before been reported to be emitted by a plant. When soil around the roots is dislodged or when seedling roots are touched, an odor is detected. The perceived odor corresponds to the emission of higher amounts of propanesulfenic acid, 2-aminothiophenol, S-propyl propane 1-thiosulfinate, and phenothiazine. The mechanosensitivity response is selective. Whereas touching the roots with soil or human skin resulted in odor detection, agitating the roots with other materials such as glass did not induce a similar response. Light and electron microscopy studies of the roots revealed the presence of microscopic sac-like root protuberances. Elemental analysis of these projections by energy-dispersive x-ray spectroscopy revealed them to contain higher levels of K⁺ and Cl⁻ compared with the surrounding tissue. Exposing the protuberances to stimuli that caused odor emission resulted in reductions in the levels of K⁺ and Cl⁻ in the touched area. The mechanistic implications of the variety of sulfur compounds observed vis-à-vis the pathways for their formation are discussed.Plant roots are known to exude a diversity of both small and macromolecular chemicals that mediate antimicrobial, antiquorum sensing, allelopathic, and other effects (De-la-Peña et al., 2012). However, the machinery associated with the synthesis and extrusion of these compounds is not well understood. One of the most intriguing but least studied of these is the emission of volatile and reactive organosulfur compounds such as the foul and toxic gas carbonyl sulfide (COS) and volatile carbon disulfide (CS₂). Both are reportedly released by numerous plants and are proposed to make a significant contribution to the environmental sulfur burden (Haines et al., 1989). As a case in point, the Central American rainforest plant Stryphnodendron exelsum (Mimosaceae) is a sufficiently strong sulfur emitter that its location in the forest can be determined by odor (Haines et al., 1989). Furthermore, 40 taxa from nine genera within the subfamily Mimosoideae revealed that 29 taxa from six genera produced CS₂ and 19 of the 40 taxa produced COS (Piluk et al., 2001). It has been proposed that the COS and CS₂ are derived from a putative Cys lyase-mediated cleavage of djenkolic acid, an amino acid previously isolated from the plant (Piluk et al., 1998), but this has not been confirmed.We used Mimosa pudica (Leguminosae), a perennial shrub endemic to Brazil but now pantropical in its distribution (Howard, 1988), as a model to begin investigations of how this and related plants emit these highly reactive and corrosive compounds without themselves incurring tissue damage. Its various colloquial names, such as sensitive plant, touch-me-not, shy plant, and humble plant, among many others (Holm, 1977), derive from its seismonastic movements: in response to touch, water, shaking, wind, or warming, its leaves quickly close, slowly opening after an average of about 10 min (Song et al., 2014). It also displays nyctinasty, with its leaves closing or “sleeping” with the onset of darkness. These curious characteristics coupled with its small size have made the plant a convenient and popular attraction in schools, greenhouses, and other learning environments, where it is used to illustrate seismonasty.Our studies show that by using direct analysis in real time high-resolution mass spectrometry (DART-HRMS; Cody et al., 2005), it is possible to detect the compounds emitted by plant roots in situ. Using this method, it was revealed that both M. pudica plants germinated aseptically on agar and those germinated in soil emitted a variety of small molecules into the atmosphere at levels that were not detectable by human subjects. However, an odor detectable by humans could be sensed when the plant root was disturbed, with odor emission being dependent on the nature of the stimulus. Analysis of the chemical contributors to the odor revealed that, although the array of compounds observed to be produced by the roots was the same both before and after stimulation, emission of a subset of organosulfur compounds was increased when the roots were stimulated. Light and scanning electron microscope imaging studies revealed the presence of sac-like protuberances dotted along M. pudica seedling root shafts that collapsed when the roots were exposed to stimuli that elicited odor emission. The detection by energy-dispersive x-ray spectroscopy (EDS) of relatively high levels of K⁺ and Cl⁻ prior to root stimulation, on the one hand, and reductions in the levels of these species, on the other hand, implicate the involvement of these ions in the observed mechanostimulatory behavior.