Redox Reactions of Biologically Active Molecules upon Cold Atmospheric Pressure Plasma Treatment of Aqueous Solutions.

Cold atmospheric pressure plasma (CAPP) is widely used in medicine for the treatment of diseases and disinfection of bio-tissues due to its antibacterial, antiviral, and antifungal properties. In agriculture, CAPP accelerates the imbibition and germination of seeds and significantly increases plant productivity. Plasma is also used to fix molecular nitrogen. CAPP can produce reactive oxygen and nitrogen species (RONS). Plasma treatment of bio-tissue can lead to numerous side effects such as lipid peroxidation, genotoxic problems, and DNA damage. The mechanisms of occurring side effects when treating various organisms with cold plasma are unknown since RONS, UV-Vis light, and multicomponent biological tissues are simultaneously involved in a heterogeneous environment. Here, we found that CAPP can induce in vitro oxidation of the most common water-soluble redox compounds in living cells such as NADH, NADPH, and vitamin C at interfaces between air, CAPP, and water. CAPP is not capable of reducing NAD+ and 1,4-benzoquinone, despite the presence of free electrons in CAPP. Prolonged plasma treatment of aqueous solutions of vitamin C, 1,4-hydroquinone, and 1,4-benzoquinone respectively, leads to their decomposition. Studies of the mechanisms in plasma-induced processes can help to prevent side effects in medicine, agriculture, and food disinfection.

Cold atmospheric pressure He-plasma jet and plasma ball interactions with the Venus flytrap: Electrophysiology and side effects.

Cold atmospheric pressure radio frequency plasma (CAPP) can play an important role in agriculture, medicine, biophysical and bioelectrochemical applications, disinfection and sterilization, synthesis of different compounds, nitrogen fixation, and treatment of surfaces. Here we found that reactive oxygen and nitrogen species, UV-Vis photons, and high-frequency strong electromagnetic fields with an amplitude of a few kV produced by a cold plasma jet can interact with bio-tissue and damage it if the plasma treatment is long enough. The electrophysiological effects of CAPP treatment of bio-tissue and electrical signals transmission were measured in the Venus flytrap. The plasma ball does not produce any visible side effects on the Venus flytrap, but induces electrical signals in bio-tissue with very high amplitude. Plasma (Kirlian) photography shows the existence of a blue aura around the plasma ball due to a corona discharge. Understanding the mechanisms of interactions between CAPP and bio-tissue and preventing side effects can contribute to the application of plasma technology in medicine and agriculture. The use of cold plasma in medicine and agriculture should be monitored for side effects from strong high-frequency electro-magnetic fields, UV photons, and reactive oxygen and nitrogen species to protect against undesirable consequences.

Cyclic voltammetry of volatile memristors in the Venus flytrap: short-term memory.

Plants have sensory, short-term and long-term memory. Possible candidates for memory in plants are memristors; resistors with memory. Memristors have been found in seeds, plants, flowers and fruits. The electrostimulation of plants by bipolar periodic waves can induce electrical responses with fingerprints of volatile or non-volatile memristors. Here, we show that the electrostimulation of the Venus flytrap (Dionaea muscipula Ellis) by unipolar sinusoidal or triangular periodic electrical trains induces electrical responses in plants with fingerprints of volatile memristors. The discovery of volatile generic memristors in plants opens new directions in the modelling and understanding of electrical phenomena in the plant kingdom.

Radio frequency plasma capacitor can increase rates of seeds imbibition, germination, and radicle growth.

Radio frequency capacitors can be used to accelerate seed imbibition, germination, increase the growth of plants seedlings, poration and corrugation of the bio-tissue surfaces without the side effects of RONS generated by cold plasma jets. Atomic force microscope data show that the plasma lamp produced morphological changes in the seed coat. Magnetic resonance imaging studies showed the acceleration of water uptake in seeds treated with radio frequency capacitors of plasma lamps. Plasma capacitor can accelerate radicle growing rates.

Underground electrotonic signal transmission between plants.

Plants can communicate with other plants using wireless pathways above and underground. Some examples of these underground communication pathways are: (1) mycorrhizal networks in the soil; (2) the plants' rhizosphere; (3) acoustic communication; (4) naturally grafting of roots of the same species; (5) signaling chemicals exchange between roots of plants; and (6) electrical signal transmission between plants through the soil. To avoid the possibility of communication between plants using mechanisms (1)-(5), soils in both pots with plants can be connected by Ag/AgCl or platinum wires. Electrostimulation Aloe vera or cabbage plants induces electrotonic potentials transmission in the electro-stimulated plants as well as in the neighboring plants located in the same or different electrically connected pots regardless if plants are the same or different types. The amplitude and sign of electrotonic potentials in both electrostimulated and neighboring plants depend on the amplitude, rise, and fall of the applied voltage. Electrostimulation serves as an important tool for the evaluation of mechanisms of underground communication in the plant-wide web. The previously developed mathematical model of electrotonic potentials transmission within and between tomato plants, which is supported by the experimental data, is generic enough to be used for simulation study and predicting the intercellular and intracellular communication in the form of electrical signals in the electrical networks within and between a variety of plants.

Electrical signal transmission in the plant-wide web.

Plants can communicate with other plants using wireless pathways in the plant-wide web. Some examples of these communication pathways are: (1) volatile organic compounds' emission and sensing; (2) mycorrhizal networks in the soil; (3) the plants' rhizosphere; (4) naturally grafting of roots of the same species; (5) electrostatic or electromagnetic interactions; and (6) acoustic communication. There is an additional pathway for electrical signal transmission between plants - electrical signal transmission between roots through the soil. To avoid the possibility of communication between plants using mechanisms (1)-(6), soils in pots with plants were connected by Ag/AgCl or platinum wires. Electrostimulation of Aloe vera, tomato, or cabbage plants induces electrotonic potentials transmission in the electro-stimulated plants as well as the plants located in different pots regardless if plants are the same or different types. The amplitude and sign of electrotonic potentials in electrostimulated and neighboring plants depend on the amplitude, rise, and fall of the applied voltage. Experimental results displayed cell-to-cell electrical coupling and the existence of electrical differentiators in plants. Electrostimulation by a sinusoidal wave induces an electrical response with a phase shift. Electrostimulation serves as an important tool for the evaluation of mechanisms of communication in the plant-wide web.

Plasma-generated reactive oxygen and nitrogen species can lead to closure, locking and constriction of the Dionaea muscipula Ellis trap.

Reactive oxygen and nitrogen species (RONS) can influence plant signalling, physiology and development. We have previously observed that an argon plasma jet in atmospheric air can activate plant movements and morphing structures in the Venus flytrap and Mimosa pudica similar to stimulation of their mechanosensors in vivo. In this paper, we found that the Venus flytrap can be activated by plasma jets without direct contact of plasma with the lobe, midrib or cilia. The observed effects are attributed to RONS, which are generated by argon and helium plasma jets in atmospheric air. We also found that application of H2O2 or HNO3 aqueous solutions to the midrib induces propagation of action potentials and trap closing similar to plasma effects. Control experiments showed that UV light or neutral gas flow did not induce morphing or closing of the trap. The trap closing by plasma is thus likely to be associated with the production of hydrogen peroxide by the cold plasma jet in air. Understanding plasma control of plant morphing could help design adaptive structures and bioinspired intelligent materials.

Cold plasma poration and corrugation of pumpkin seed coats.

The treatment of seeds and plants by electrically generated cold atmospheric pressure plasma can accelerate seed germination and radicle growing rates. The plasma generated reactive oxygen and nitrogen species, UV photons, and high frequency electromagnetic fields can penetrate into seed coats and modify their surface properties. Atomic force microscope data shows that cold helium or argon plasma induces strong corrugation of pumpkin seed coats, produces pores and surface defects. These structural deformations and poration enhance water uptake by seeds during the imbibing process, accelerate seeds germination, and increase seed growth. The cold atmospheric pressure plasmas treatment of pumpkin seeds also decreases the apparent contact angle between a water drop and the seed surface, thereby improving the wetting properties of seeds surfaces. Magnetic resonance imaging studies show acceleration of water uptake in pumpkin seeds exposed to a cold plasma jet. Reactive nitrogen and oxygen species, high frequency electromagnetic fields and photons emitted by the plasma jets accelerate germination of pumpkin seeds both independently and synergistically. These results show that cold plasma can be used in agriculture for acceleration of seed germination, increasing growth of plants seedlings, poration and corrugation of the bio-tissue surfaces.

Signaling in electrical networks of the Venus flytrap (Dionaea muscipula Ellis).

The Venus flytrap captures insects with one of the most rapid movements in the plant kingdom. There is a significant difference between properties of electrical signals generated in the Venus flytrap described in literature. Amplitudes of action potentials vary from 14 mV to 200 mV with duration of signals from 2 ms to 10 s. Here we present experimental study of potential differences between Ag/AgCl electrodes inserted to the trap, petiole, and into soil or external ECG electrodes attached to surfaces of the Venus flytrap. Diverse types of electrodes with various positions in a plant tissue or in soil show different amplitude and duration of electrical signals because potentials are measured in different electrochemical circuits. Electrical signals in the Venus flytrap were induced by mechanical stimulation of the trigger hairs or by chemical stimulation of a midrib using small drops of H2O2 or HNO3. Here we found that action potentials can propagate with speed up to 10 m/s in the trap of D. muscipula. Results are compared with equivalent electrical circuits.

Electrical signal propagation within and between tomato plants.

According to literature, electrostimulation of plants can induce plant movement, activation of ion channels, ion transport, gene expression, enzymatic system activation, electrical signaling, plant-cell damage, enhanced wound healing, and can also influence plant growth. Many plants can communicate above ground and underground between adjacent plants. Electrostimulation by square pulses induces passive electrotonic potentials propagating within and between tomato plants. The amplitude and sign of electrotonic potentials, in both the electrostimulated and neighboring tomato plants depends on the amplitude, rise and fall of the applied voltage. Electrostimulation by the pulse train, sinusoidal and triangular saw-shape voltage profile shows the existence of electrical differentiators and refractory periods in cell-to-cell electrical coupling in tomato plants. Electrical networks within one tomato plant can communicate underground with electrical circuits in another tomato plant. Here, we present the mathematical model of electrotonic potentials transmission between tomato plants which is supported by the experimental data. The information gained from this mathematical model and analytical study can be used not only to elucidate the effects of electrostimulation on higher plants, but also to observe and predict the intercellular and intracellular communication in the form of electrical signals within the electrical networks within and between tomato plants.

Sunpatiens compact hot coral: memristors in flowers.

Leon Chua postulated the theory of a memristor - a resistor with memory - in 1971, and the first solid-state memristor was built in 2008. Memristors exist in vivo as components of plasma membranes in plants, fruits, roots and seeds. A memristor is a nonlinear element; its current-voltage characteristic is similar to that of a Lissajous pattern. Here, we found memristors in flowers. Electrostimulation by bipolar periodic sinusoidal or triangular waves of an androecium, a spur, petals and a pedicel in Sunpatiens flowers induces hysteresis loops with a pinched point at low frequencies between 0.1mHz and 1mHz. At high frequencies, the pinched hysteresis loop transforms to a non-pinched hysteresis loop instead of a single line I=U/R for ideal memristors because the amplitude of electrical current depends on capacitance of a flower's tissue and electrodes, frequency and direction of scanning. The discovery of memristors in Sunpatiens (Impatiens spp.) creates a new direction in the modelling and understanding of electrophysiological phenomena in flowers.

Cold plasma interactions with plants: Morphing and movements of Venus flytrap and Mimosa pudica induced by argon plasma jet.

Low temperature (cold) plasma finds an increasing number of applications in biology, medicine and agriculture. In this paper, we report a new effect of plasma induced morphing and movements of Venus flytrap and Mimosa pudica. We have experimentally observed plasma activation of sensitive plant movements and morphing structures in these plants similar to stimulation of their mechanosensors in vivo. Application of an atmospheric pressure argon plasma jet to the inside or outside of a lobe, midrib, or cilia in Dionaea muscipula Ellis induces trap closing. Treatment of Mimosa pudica by plasma induces movements of pinnules and petioles similar to the effects of mechanical stimulation. We have conducted control experiments and simulations to illustrate that gas flow and UV radiation associated with plasma are not the primary reasons for the observed effects. Reactive oxygen and nitrogen species (RONS) produced by cold plasma in atmospheric air appear to be the primary reason of plasma-induced activation of phytoactuators in plants. Some of these RONS are known to be signaling molecules, which control plants' developmental processes. Understanding these mechanisms could promote plasma-based technology for plant developmental control and future use for plant protection from pathogens. Our work offers new insight into mechanisms which trigger plant morphing and movement.

Electrotonic potentials in Aloe vera L.: Effects of intercellular and external electrodes arrangement.

Electrostimulation of plants can induce plant movements, activation of ion channels, ion transport, gene expression, enzymatic systems activation, electrical signaling, plant-cell damage, enhanced wound healing, and influence plant growth. Here we found that electrical networks in plant tissues have electrical differentiators. The amplitude of electrical responses decreases along a leaf and increases by decreasing the distance between polarizing Pt-electrodes. Intercellular Ag/AgCl electrodes inserted in a leaf and extracellular Ag/AgCl electrodes attached to the leaf surface were used to detect the electrotonic potential propagation along a leaf of Aloe vera. There is a difference in duration and amplitude of electrical potentials measured by electrodes inserted in a leaf and those attached to a leaf's surface. If the external reference electrode is located in the soil near the root, it changes the amplitude and duration of electrotonic potentials due to existence of additional resistance, capacitance, ion channels and ion pumps in the root. The information gained from this study can be used to elucidate extracellular and intercellular communication in the form of electrical signals within plants.

Cyclic voltammetry of apple fruits: Memristors in vivo.

A memristor is a resistor with memory that exhibits a pinched hysteretic relationship in cyclic voltammetry. Recently, we have found memristors in the electrical circuitry of plants and seeds. There are no publications in literature about the possible existence of memristors and electrical differentiators in fruits. Here we found that the electrostimulation of Golden Delicious or Arkansas Black apple fruits by bipolar periodic waves induces hysteresis loops with pinched points in cyclic voltammograms at low frequencies between 0.1MHz and 1MHz. At high frequencies of 1kHz, the pinched hysteresis loop transforms to a non-pinched hysteresis loop instead of a single line I=V/R for ideal memristors because the amplitude of electrical current depends on capacitance of a fruit's tissue and electrodes, frequency and direction of scanning. Electrostimulation of electrical circuits in apple fruits by periodic voltage waves also induces electrotonic potential propagation due to cell-to-cell electrical coupling with electrical differentiators. A differentiator is an electrical circuit in which the output of the circuit is approximately directly proportional to the rate of change of the input. The information gained from electrostimulation can be used to elucidate and to observe electrochemical and electrophysiological properties of electrical circuits in fruits.

Electrophysiology of pumpkin seeds: Memristors in vivo.

Leon Chua, the discoverer of a memristor, theoretically predicted that voltage gated ion channels can be memristors. We recently found memristors in different plants such as the Venus flytrap, Mimosa pudica, Aloe vera, apple fruits, and in potato tubers. There are no publications in literature about the existence of memristors in seeds. The goal of this work was to discover if pumpkin seeds might have memristors. We selected Cucurbita pepo L., cv. Cinderella, Cucurbita maxima L. cv Warty Goblin, and Cucurbita maxima L., cv. Jarrahdale seeds for this analysis. In these seeds, we found the presence of resistors with memory. The analysis was based on cyclic voltammetry where a memristor should manifest itself as a nonlinear two-terminal electrical element, which exhibits a pinched hysteresis loop on a current-voltage plane for any bipolar cyclic voltage input signal. Dry dormant pumpkin seeds have very high electrical resistance without memristive properties. The electrostimulation by bipolar sinusoidal or triangular periodic waves induces electrical responses in imbibed pumpkin seeds with fingerprints of memristors. Tetraethylammonium chloride, an inhibitor of voltage gated K(+) channels, transforms a memristor to a resistor in pumpkin seeds. NPPB (5-Nitro-2-(3-phenylpropylamino)benzoic acid) inhibits the memristive properties of imbibed pumpkin seeds. The discovery of memristors in pumpkin seeds creates a new direction in the understanding of electrophysiological phenomena in seeds.

Memristors: Memory elements in potato tubers.

A memristor is a nonlinear element because its current-voltage characteristic is similar to that of a Lissajous pattern for nonlinear systems. This element was postulated recently and researchers are looking for it in different biosystems. We investigated electrical circuitry of red Irish potato tubers (Solanum tuberosum L.). The goal was to discover if potato tubers might have a new electrical component - a resistor with memory. The analysis was based on a cyclic current-voltage characteristic where the resistor with memory should manifest itself. We found that the electrostimulation by bipolar sinusoidal or triangle periodic waves induces electrical responses in the potato tubers with fingerprints of memristors. Tetraethylammonium chloride, an inhibitor of voltage gated K(+) channels, transforms a memristor to a resistor in potato tubers. Our results demonstrate that a voltage gated K(+) channel in the excitable tissue of potato tubers has properties of a memristor. Uncoupler carbonylcyanide-4-trifluoromethoxy-phenyl hydrazone decreases the amplitude of electrical responses at low and high frequencies of bipolar periodic sinusoidal or triangle electrostimulating waves. The discovery of memristors in plants creates a new direction in the understanding of electrical phenomena in plants.

An analytical model of memristors in plants.

The memristor, a resistor with memory, was postulated by Chua in 1971 and the first solid-state memristor was built in 2008. Recently, we found memristors in vivo in plants. Here we propose a simple analytical model of 2 types of memristors that can be found within plants. The electrostimulation of plants by bipolar periodic waves induces electrical responses in the Aloe vera and Mimosa pudica with fingerprints of memristors. Memristive properties of the Aloe vera and Mimosa pudica are linked to the properties of voltage gated K(+) ion channels. The potassium channel blocker TEACl transform plant memristors to conventional resistors. The analytical model of a memristor with a capacitor connected in parallel exhibits different characteristic behavior at low and high frequency of applied voltage, which is the same as experimental data obtained by cyclic voltammetry in vivo.

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.

Morphing structures of the Dionaea muscipula Ellis during the trap opening and closing.

The Venus flytrap is a marvelous plant that has intrigued scientists since the times of Charles Darwin. This carnivorous plant is capable of very fast movements to catch a prey. We found that the maximal speed of the trap closing in the Dionaea muscipula Ellis is about 130,000 times faster than the maximal speed of the trap opening. The mechanism and kinetics of this movement was debated for a long time. Here, the most recent Hydroelastic Curvature Model is applied to the analysis of this movement during closing and opening of the trap with or without a prey. Equations describing the trap movement were derived and verified with experimental data. Chloroform and ether, both anesthetic agents, induce action potentials and close the trap without the mechanical stimulation of trigger hairs. We tested this by dropping 10 µL of ether on the midrib inside the trap without touching any of the mechanosensitive trigger hairs. The trap closed slowly in 10 s. This is at least 20 times slower than the closing of the trap mechanically or electrically. The similar effect can be induced by placing 10 µL of chloroform on the midrib inside the trap, however, the lobes closing time in this case is as fast as closing after mechanical stimulation of the trigger hairs.

Memristors in plants.

We investigated electrical circuitry of the Venus flytrap, Mimosa pudica and Aloe vera. The goal was to discover if these plants might have a new electrical component--a resistor with memory. This element was postulated recently and the researchers were looking for its presence in different systems. The analysis was based on cyclic current-voltage characteristic where the resistor with memory should manifest itself. We found that the electrostimulation of plants by bipolar sinusoidal or triangle periodic waves induces electrical responses in the Venus flytrap, Mimosa pudica and Aloe vera with fingerprints of memristors. Tetraethylammonium chloride, an inhibitor of voltage gated K(+) channels, transforms a memristor to a resistor in plant tissue. Our results demonstrate that a voltage gated K(+) channel in the excitable tissue of plants has properties of a memristor. This study can be a starting point for understanding mechanisms of memory, learning, circadian rhythms, and biological clocks.