Fulvalenes UV Anthocyanins Plants London 2013 Final 2

Instituto de Investigación Lightbourn A.C. Bionanofemtofisiología Vegetal Disruptiva Fulvalene, Rotaxane and Catenane Compounds: Ultraviolet Radiation Effects on Plants and its Bioremediation Lightbourn-Rojas L A1, León-Chan R G1, Heredia J B1, 2 1 Instituto de Investigación Lightbourn A.C. Bionanofemtofisiología Vegetal Disruptiva Centro de Investigación en Alimentación y Desarrollo A.C. (CIAD) Unidad Culiacán 2 Abstract There are several studies on climate change, which focus on different factors: the increase in atmospheric CO2, droughts, variable amount of rainfall, sudden temperature changes and the increase of ultraviolet radiation (UV) due to the deterioration of the ozone layer by atmospheric contaminants. The increase of UV radiation causes adverse responses in plants development, particularly in DNA and in photosynthetic system, especially in photosystem II, therefore the loss of photosynthetic efficiency and a foods reduction. Plants have developed several defense systems which are related to the synthesis of anthocyanins. These compounds can protect plant tissues by absorbing excess of light, UV radiation or by their antioxidant capacity, scavenging UV produced reactive oxygen species. However, sometimes these mechanisms are insufficient due to photosynthetic deficiency; therefore we have developed a new plant nutrition technology, based on fulvalene, rotaxane and catenane compounds. With this approach, an increase in the uptake, storage and availability of monochromatic ray at 563 nm can be achieved. This can induce the photosynthetic optimization and the reduction of metabolic delays in the photosynthetic system. This would help to maintain the plant metabolism and therefore the stability of the food production regardless of the adverse environmental conditions. Introduction The climate of a planet is determined by its total mass, its distance from the closest star(s) (in the case of the Earth, the Sun) and the chemical composition of its atmosphere (González et al., 2003). The latest is the most variable factor, both naturally and by the activities of the organisms that inhabit it, especially humans. In recent decades there have been a wide range of environmental changes that are affecting life on Earth, so there are a variety of studies on climate change, which focus on different factors such as the increase in atmospheric CO2 concentration, melting icecap, drought, change in rainfall patterns, wind speed, sudden temperature change, global warming and increased ultraviolet radiation, among others (Bellard et al., 2012). Plants are the organisms more exposed to environmental changes because they can not migrate from where they develop, so they are essential baseline to understanding the impact of climate change on Earth. Changes in the environment to produce a negative impact on growth and development of plants are considered as stress factors (Bray et al., 2000). This can be classified into primary stress, when the stress factor directly affects the development of plants, and secondary stress, when the stress factor provides generation of reactive oxygen species (ROS) which adversely affect the metabolism of the plant (Ahmad y Prasad 2012). The effects of climate change on living organisms can be classified into four categories (Gonzales et al., 2003): 1) geographic distribution (tendency of some species to move), 2) adaptation (micro-evolutionary changes in situ) 3) physiological (photosynthesis, respiration, growth) and 4) phenological (changes in the life cycles such photoperiod effects, cold hours, etc.). Therefore, changes in radiation and temperature are factors that have more impact on the organisms present on Earth, especially in plants, which are the basis of life energy on the Earth, because they can transform solar energy into chemical energy , which is taken by the rest of the organisms as food. The plant growth is defined as an increase in dry matter, while development is the increase in the number and size of their organs by division and/or cell expansion: leaves, sticks, swords, flowers, roots, etc. The growth rate is proportional to the product of the rate and efficiency of the catabolic activity to convert photosynthates to structural biomass, therefore, plant growth is extremely temperature sensitive (Lightbourn, 2011a; Źróbek-Sokolnik, 2012). On the other hand, solar radiation (electromagnetic radiation) is one of the most important factors in the growth and development of plants (Carrasco-Ríos, 2009), and is involved in many important processes such as photosynthesis, phototropism, photomorphogenesis, opening stomata, soil and temperature, etc. (Salisbury and Ross, 1994; Castilla, 2007). Solar radiation can be divided into three main ranges of wavelength: infrared, visible and ultraviolet radiation (UV) (Castilla, 2007). UV radiation provides a higher energy level that can adversely affect the metabolism of any organism. This type of radiation is partially absorbed by the ozone layer. However, the increasing of certain gases such as chlorofluorocarbons (CFC's) resulted in the deterioration of the protective layer, therefore lead to an increase of UV radiation at the Earth's surface (Hollósy, 2002). The stress caused by increase of UV radiation triggers a range of responses, such as alterations in gene expression and changes in cell metabolism, which has influence on growth, development and production of plants. The length and severity of stress, as well as the characteristics of the plants, their stage of development, the affected tissue and genotype would influence on the answers plants can present (Bray et al., 2000). One of the most commonly response presented on stressed plants induced by UV radiation is the synthesis of flavonoids, such as anthocyanins, which may be due to its ability to absorb UV light or to the antioxidant capacity that these compounds provide to reverse oxidative damage caused by UV radiation on cells (Mazid et al., 2011). However, these mechanisms could be insufficient due to photosynthetic deficiency induced by UV radiation. Therefore, a new technology in plant nutrition has been developed which optimizes the photosynthetic efficiency by maintaining more available the power provided by the monochromatic ray of 563 nm. Temperature In recent decades there has been a rise in temperature that is seriously affecting the biological processes on Earth. This temperature increase is widespread over the globe, such as in the Arctic, where the average temperature has increased at almost twice as fast as the global average of the last one hundred years (IPCC, 2007). The temperature is not a measure of amount or concentration of a substance or total energy. The temperature measures the molecular movement, which means, the kinetic energy of the molecules within the system. Consequently, the velocity indices of all elemental reactions increase exponentially with the temperature increments (Lightbourn, 2011a). The thermodynamics first law refers that the heat added to a system minus the work done by this, it will produce a change in the internal energy of the system: ΔE = q - W where ΔE is the change in internal energy of the system, q is the amount of heat added and W is the work carried out by the system, in this case we refer to plants as the system in question. If q < W then the internal energy of the plant decreases down to death, while if q >>> W then the heat inside the plant will increase causing denaturation of its components (Lightbourn et al., 2012). Therefore, a change of a few Celsius degrees leads to a significant change in the growth rate, and this is because there is not a range in which the temperature stops changes that affect the growth rates, so it is incorrect to talk about stress ranges by temperatures, being correct when talking about stress by temperaturetime (Lightbourn, 2011a). The stress caused in plants by temperature is classified into three types: 1) cold damage, 2) freezing damage and 3) by high temperatures. This is because each species or variety has, at any given state of its life cycle, a minimum temperature below which does not grow, the optimum temperature (or temperature range) which grows to a maximum rate and temperature above the maximum which will not grow and may even die (Salisbury y Ross, 1994; Źróbek-Sokolnik, 2012). Therefore, the temperature is defined as an environmental factor that significantly affects biological processes in all organisms because it will primarily modify the properties of the membranes, the levels of enzymatic activity, speed up chemical reactions, as well as affecting the phloem, xylem and cytoplasm solutions (Taiz y Zeiger 2002; Źróbek-Sokolnik, 2012). The membranes are in one of three stages depending on the external environment: 1) as a crystalline liquid phase, which represents the range of fluidity in which the membrane and its components work naturally, 2) as a solid gel, which represents a membrane that retains its shape but is rigid and therefore not functional, 3) and hexagonal and cubic phases, which represents the disruption of the membrane caused by extreme environments (Nilsen and Orcutt, 1996). In the first stage we have mentioned a balanced equation of thermodynamics first law, while cases two and three are derived from an imbalance between the work of the plant and the added heat (Lightbourn et al 2012). The increase in temperature causes a greater fluidity in the membranes, causing problems in cell functions, mainly in mitochondria and chloroplasts (Allakhverdiev et al., 2008). Therefore, the deleterious effects that high temperatures cause on plants mainly occur in photosynthetic functions and thylakoid membranes. The three more sensitive photosynthetic sites to heat stress are the carbon assimilation process, the ATP generation and the photosystems, mainly photosystem II complexes (PSII), which are the most labile part of the photosynthetic system to the heat effects (Salisbury and Ross, 1994; Allakhverdiev et al., 2008). Also, the synthesis of various thylakoid membrane proteins is extremely reduced during elevated temperature exposure (Süss and Yordanov, 1986), i.e. the apoprotein reaction center of photosystem II (P680), the sub-units α and β of the ATPase synthase, cytochrome ƒ, cytochrome b559 and the apoprotein from the center of the CP47 antenna complex (Santarius, 1973). This causes a great disruption of PSII leading the plant inability to produce energy (ATP) from photosynthesis. In general, the photosynthetic activity is stable until 30 °C but dramatically decreases above this temperature until a complete inhibition is reached at 40 °C (Carpentier, 2005). Moreover, the increase chloroplasts membrane fluidity allows the ATP molecules to go through more easily from citosol to chloroplasts (Carpentier, 2005). ATP is used in chloroplast to carry out the synthesis of carbohydrates in mature plants, but in plants in growth phase, the ATP is used predominantly for proteins and nucleotides synthesis, for which is required greater amount of energy (Salisbury and Ross, 1994), so that in the photosynthetic system inefficiency caused by stress is necessary to obtain energy from other sources to continue the plant development until reserves will allow it. Therefore, the morphological symptoms presented in heat stressed plants may include sunburn on leaves, branches and stems, senescence and abscission of leaves, stem growth inhibition and roots, damage and fruit discoloration, less production, cell size reduction, stomatal closure and reduction of transpiration to prevent dehydration (Waheed et al.; 2007 Mitra and Bhatia, 2008). The observed increase in global average temperatures since the midtwentieth century is mainly due to the observed increase in concentrations of anthropogenic greenhouse gases, which was found at 70% from 1970 to 2004; this increase is due to the higher emission of these gases in relation of its decomposition. Human activities result in emissions of four long-lived greenhouse gases: carbonic anhydride mostly known as carbon dioxide (CO2), methane (CH4), nitrous oxide (N2O) and chlorofluorocarbons (CFC's, group of gases containing fluorine, chlorine or bromine). Increased CO2 concentrations are due primarily to fossil fuel use, the CH4 is predominantly due to agriculture and fossil fuel use and N2O increased is mainly due to agricultural activities (IPCC, 2007). The increase in some of these compounds causes the degradation of the ozone layer, allowing a greater emission of ultraviolet rays at the Earth surface, which also inferred in plants development (Björn and McEnzie, 2008). Ultraviolet Radiation (UV) The UV radiation corresponds to a wavelength range of 200 to 380 nm. This radiation is low when the sun elevation above the horizon is low and to low altitude. UV radiation represent about 9% of the solar total radiation energy and can be divided into UV-A (320 to 280 nm) skin tanning radiation, UV-B (290 to 320 nm) responsible of skin cancer, and UV-C (200-290 nm) which is potentially hazardous but almost completely absorbed by the ozone layer (Castilla, 2007, Prado et al., 2012). The absorption coefficient of ozone decreases rapidly at wavelengths above 280 nm and approaches zero around 330 nm (Hollósy, 2002). Although the stratospheric ozone determines the amount of UV radiation that reach the Earth surface, its level is significantly affected by variations in latitude and altitude. The level of UV radiation over tropical latitude is higher than in temperate regions due to lesser atmospheric UV absorption determined by the solar angle and the ozone layer itself, which is thinner in equatorial regions (Jaakola and Hohtola, 2010; Prado et al., 2012). In 1987, it was established the Montreal Protocol on "Substances that deplete the Ozone Layer" to carry out the reduction of these compounds in the atmosphere, mainly CFCs; however, the deterioration continues increasing to 0.6 % yearly (Prado et al., 2012). In general, every 1 % of reduction in the ozone layer results in an increase of 1.3 to 1.8 % of UV-B radiation (Hollósy, 2002). Most studies concerning that the effects of UV radiation increases are focus on UV-B radiation, however, the presence of UV-C radiation has been show, even within plastic greenhouses (León-Chan, 2012), so there are some studies that provide UV-C to study their effects on plants (Mahdavian et al., 2008; Sarghein et al., 2008; Katerova et al., 2009). UV radiation can inhibit photosynthesis by altering gene expression and by damaging the parts of the photosynthetic machinery (Smith et al., 2009). Sites that are affected by this type of light are the light collector complex II (LHCII), the PSII reaction center and PSI acceptor. However, most studies have demonstrated that PSII is more sensitive to UV radiation as compared to PSI; this is due to the chemical changes which produces the UV radiation on amino acids with double bonds of the PSII proteins (Carrasco-Ríos, 2009). The aromatic amino acids such as phenilalanine, tryptophan and tyrosine, as well as cysteine, cystine, and histidine, would provide to proteins the characteristic of absorbing UV and thereby the property to be modified (Hollósy, 2002). The amino acid histidine is present in the PSII D1 protein, which contains chlorophylls linked to it (Taiz and Zeiger, 1998), therefore, the conformational changes causing by UV on these proteins will liberate chlorophylls, facilitating their photo-oxidation (Mahdavian et al ., 2008). The UV radiation also induces the loss of enzymes activity involved in the Calvin cycle, especially on the 1,5 diphosphate carboxylase (Rubisco) which catalyzes the CO2 incorporation. Another effect of UV radiation is the production of reactive oxygen species (ROS), which also act on the denaturation of proteins; moreover, ROS are involved in the lipoperoxidation processes from the plasma membrane (Björn and McEnzie, 2008, Carrasco-Ríos, 2009). However, ROS can also initiate signals stress responses to UV, including enzyme activation, gene expression, programming cell death, etc. (Jaspers and Kangasjärvi, 2010). The DNA is also sensitive to UV-B and UV-C, because these photons promote π-π transitions in nitrogenous bases that constitute the nucleotides, altering the normal establishment of chemical bonds. This mainly causes the formation of cycle butane pyrimidine dimers (CPD) produced by the dimerization of adjacent pyrimidines (TT, TC, CT or CC), and produce other compounds known as (6-4) photo products (Björn and McEnzie, 2008; Carrasco Rivers, 2009). The biological effects of these lesions are variable, since in some cases replication in the lesion is stopped, while in other cases the replication continuous, thereby promoting mutations. Therefore, these products are the leading causes of cancer (Björn and McEnzie, 2008). Plant Responses to Ultraviolet Radiation The plants exhibit different responses to environmental stresses like UV radiation exposure. Under these conditions, plants would show the presence of wax cuticle deposition and trichomes that function as radiation reflectors; leaf area reduction and increased of sheet thickness to reduce the damaged area; changes on stomatal density, reduced of stems elongation, changes in branching pattern, the synthesis of secondary metabolites with the ability to absorb UV light, as well as alterations in plant-pathogen plant-predator interactions and gene expression (Prado et al., 2012). The synthesis of secondary metabolites of the phenylpropanoid pathway has been widely studied as a defense mechanism to counteract the deleterious effects of UV radiation produced in plants. Among these compounds there are phenolic acids, insoluble polyphenols and flavonoids such as anthocyanins. Today anthocyanins are the most studied flavonoids usually referred as the compounds capable of reducing the photo-oxidative damage. These compounds stay mainly in the epidermis cells, and are also being responsible for a variety of colorations in plant tissues (Castañeda-Ovando et al., 2009). Some authors mention that the tissues most exposed to the sun show higher anthocyanin content, presenting a high variability of the content of these compounds from leaves or fruits of the same plant (Salisbury and Ross 1994; Lightbourn et al., 2008; Steyn, 2012). Here, the hypothesis about the anthocyanin function in plants are showed: 1) protection of chloroplasts from excessive light, especially in plants in development process or that have developed under shaded conditions (Oren-Shamir, 2009); 2) protection against UV radiation, by the ability of these compounds to absorb this type of radiation and, 3) the antioxidant capacity that these compounds have, several times greater than some of its antioxidant vitamins analogues. This antioxidant capacity could reduce the damage caused by ROS generated by UV radiation (Hatier and Gould, 2009). Some researches have shown an anthocyanins increase by effect of UV radiation on different plant sources and in different development stages (Mahadavian et al., 2008; Saghein et al., 2008, Guo and Wang 2010; Leon Chan, 2012). Therefore, there are investigations about the induction of some enzymes involved in anthocyanin synthesis by radiation effect, finding an increase in the activity of some of them, among which are the phenylalanyl ammonia lyase (PAL), chalcone synthase (CHS), chalcone isomerase (CHI), dyhydroflavonol 4-reductase (DFR), anthocyanin synthase (ANS), flavonone-3-hydroxylase (F3H), etc. (Tsukasa et al. 2000; Hao et al., 2009; Guo and Wang 2010). With these facts, it has been proven the involvement of these compounds on UV protection effects; besides obtaining more information about the expressing routes and thereby creating new alternatives that will promote the survival of plants against UV radiation stress (Guo et al., 2008). Other Environmental Impacts by Ultraviolet Radiation The changes in plants metabolism due to the increase of UV radiation not only reduce the life of plants affected, because the reduction of food production would cause great problems in the life of other organisms, particularly in humans; as well as other issues like the balance of certain cycles, such as the carbon cycle that can be severely affected by the changes in damaged plants and their capture mechanism, as well as photosynthesis, carbon storage and respiration, among others (Zepp et al., 2007). The classification of the UV radiation impact on the Earth's surface is related to direct effects such as photosynthesis inhibition and the carbon cycle balance; and indirect effects such as the alteration of the chemical composition of plant foods which will also affect the decomposition of organic plant matter in the soil by other organisms (Smith et al., 2009). The increase of anthocyanins in plant tissues can cause changes in plants interactions with other organisms. These interactions include attracting pollinators and frugivores as well as herbivores and parasites repellents (Lev-Yadum and Gould, 2009). It has also been observed some reduction of pathogenic attack, therefore the increase of these compounds in certain case could be beneficial (Paul et al., 2012). However, the production of these compounds may result in a high energy cost to the plants, and it can become even greater considering the deficiencies in energy production of plants whose have been damaged by UV radiation; so there is a difficulty to the plant in order to sustain this defense systems, as well as to reduce the production of plants with this stress (Wargent and Jordan, 2013). This reduction in food production may become even a greater problem, considering the increase in world population that is estimated at 9 billion of people by 2050 (The Royal Society, 2009). The leaves commonly that contain anthocyanins absorb more light in the visible green and yellow region spectrum regarding those leaves which do not contain it; however, is unknown this energy function (Hatier and Gould, 2009). This energy could be used in repair mechanisms or photosynthesis optimization, whereby the plant can continue to perform its metabolic activities with the least possible deficiency; nevertheless, this mechanism may occur deficient with nutritional techniques that have to now, since the increase in certain nutrients appears not to be improved in plants that are damaged by UV light, because it only considers some basic elements and excluding that plants would use many more elements to carry out their functions (Lightbourn et al., 2011; Singh et al., 2012). On the other hand, there is an increase in the use of plastic greenhouses in order to reduce the risk of pathogen attack, and also to reduce high UV radiation levels, therefore preventing from reaching and damaging the plants. However, when the light passes from one medium to another, it suffers inverted cycloid deviations namely tautocrony producing dicroism and birefringence that affects the polarity and intensity of the light beam incident and refracted which consequently alters the photosynthetic phenomenon. This can be quantified as a function of light energy, starting from the basic conceptions and traditional until the logic formality given by the mathematical complex of variations calculation in cycloid curves of mathematical analysis (Lightbourn, 2010). Reducing Plant Damage It has been demonstrated that with the application of a high dose of photosynthetically active radiation (PAR) in conjunction with low levels of UV-B radiation, Rubisco levels are unaffected; therefore, a sufficient amount of PAR radiation diminishes the UV radiation adverse effects (Hollósy, 2002). Furthermore inhibition of PSII activity in intact leaves during heat stress at 40 °C is mitigated if performed during a low-intensity illumination. This is because light is responsible to activate the adaptation mechanisms of the photosynthetic apparatus in the presence of stress temperature, especially photosystems repair, which require a low light intensity to carry out the protein phosphorylation and stimulation of diverse enzymes activity. In contrast, a strong light accelerates the PSII deterioration (Carpentier, 2005; Allakhverdiev et al., 2008). Actually a new technology has developed in plant nutrition that consists on clusters of selenium, nickel, titanium and polioxomolibdato. This is in fulvalenic rotaxane-catenananica base, generating orthogonal sequence dendrimers that are intra-tilacoidic nanosomes, which enables to optimize the photosynthetic efficiency by capturing, storing and maintaining more available power provided by the monochromatic beam of 563 nm. Therefore, induces the photosynthesis optimization, serving as a light reserve at chloroplasts level. This helps to maintain the plant metabolism, which results in phytotaxic stability, and therefore, stability of production, irrespective of any adverse conditions such as UV radiation stress (Lightbourn 2011b). Furthermore this technology contains zinc as the principal element, which is required for growth hormones synthesis (cytokinins and auxins), besides participate in chlorophyll production and possibly prevent its destruction (Salisbury and Ross, 1994), thereby further reduce the damage caused by high temperatures and UV radiation. This compound is applied by foliar absorption and involves an innovation in signaling and synchronization cell, because provide continuity in photosynthetic energy uptake and transfer, due to that clusters absorb and store energy. Furthermore, owing of these clusters not interrupt the metabolism on account of darkness, there are no delays in the formation and maintenance of plant tissue which means the total dejection of metabolic delays and consequences translated into structural failures, metabolic, energetic and homeostatic that directly affect the quantity and quality of biomass (Lightbourn, 2011b). Besides the damage due to overheating and/or UV radiation, may be metabolic delays, which are because when the active photosynthetic cycle has been initiated in diurnal phase and this is interrupted by reasons such lack of light or decreased of light intensity, results in a total restart of metabolic energy mechanism. These interruptions or decrease of PAR radiation may be due to changes in weather, such as air pollution or sudden cloud, as well as the use of greenhouses that being a barrier which causes changes in the light beam and can also accumulate substances that block the arrive of PAR radiation to the foliage plants. i.e., in crops which begin its photosynthetic processes and are interrupted by blockage or decreased of solar radiation due to cloudy, the plants must wait for a recurrence of the initial conditions of radiation to restart the total process because the plants do not continue its metabolism in the point where is stopped. Therefore, this involves a long metabolic delay of many hours or even days, this depend of the event length, which is reflected in a reduction of the creation and repair of maintenance and production tissues. The implementation of this technological innovation made by Lightbourn Metabolic Engineering and Lightbourn Biochemical Model, diametrically change the situation because when the clusters of selenium, titanium, nickel and polioxomolibdate are applied with an amphiphilic colloids nutrition, they are directly positioned on the chloroplast tilacoidic structures and this acting as absortors, storers and interconverters in a intermolecular triple switch, that able to realize consecutive and reversible monoelectrics reductions, schematized as shown in the following figures: Photosynthetic Phenomena Optimizing of 563 nm Monochromatic Beam to Optimized Induced Photosynthesis I1 = Ultraviolet radiation I2 = Visible light I3 = H+ induce the interconversion among the states 5, 6 and 7, the 5 state do not absorbs in the visible region, the 6 state (yellow-green) absorbs in 401 nm (01), the 7 state absorbs in 563 nm (02) purple. The (H+) controls the reversible interconversion between 8 and 9, in response to ultraviolet (I1) and visible (I2) stimuli. The intramolecular triple switch modulates the ratio between the two forms and the absorbance (O) of ⑨ through of photoinduced protonic transfer, the truth table and the logical sequence of the circuit shows as performed intramolecular communication. ó1 TETRAHIAFULVALENE (TTF) BIPINIDINIUM (BIPY) ABSORBANCE Low Low High Low High Low High Low Low High High High The fluorescence of Pyrazolina interval is high when the (H+) is low and viceversa. The fluorescence of the derived antrcinic is high when (Se+) or (K+) is high, the emission is low when the concentration of both is low. (Low = 0) (High = 1) Rotaxane (2) Nanoscaled incorporating a Ni (II) Tris-Bipyridine stopper and two Bipyridium electroactive units. This unit is able to perform two consecutive monoelectric reductions and reversible in the presence of Triethylenediamine solution. Cited Literature Ahamad P, Prasad MNV, 2012. Environmental Adaptations and Stress Tolerance of Plants in the Era of Climate Change. 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