1. Introduction
Higher plants are sessile therefore are continuously exposed to different environmental stress factors, such as drought, salinity, heavy metals, nutritional disorders, radiation without any protection. Most of these stresses produce certain common effects on plants, like induced oxidative stress by overproduction ofreactive oxygen species (ROS), besides their own specific effects (Rao, 2006). Thus, plants have developed their own specific response(s)against each of these stresses as well as cross-stress response(s). Investigating these responses is difficult under field conditions, but plant tissue culture techniques are performed under aseptic and controlled environmental conditions. These advantages of plant tissue culture allow various opportunities for researcher to study the unique and complex responses of plants against environmental stresses (Sakthivelu et al., 2008, Lokhande et al., 2011).
ROS have inevitably been factors for aerobic life since the introduction of molecular oxygen (O2) into our atmosphere by O2-evolving photosynthetic organisms. ROScansimplybe described highly reactive and partially reduced-oxygen forms. ROS, including the superoxide radical (O2˙ˉ), singlet oxygen (1O2), hydroxyl radical (OH˙), hydroperoxyl radical (HO2˙), hydrogen peroxide (H2O2) like that, are produced not only during metabolic pathway in several compartments of plants, including chloroplasts, mitochondria, peroxisomes, plasma membrane, apoplast, endoplasmic reticulum, and cell-wall but also as a result of induced environmental stress factors. When exposing of environmental stress factors, ROS levels can dramatically increase and this increase, in the later stage, leads to oxidative stress. Oxidative stress is defined a serious imbalance between the production of ROS and antioxidant defense and this situation can cause damage to cellular macromolecules, including proteins, lipids, carbohydrates and DNA (Mittler et al., 2004; Gill and Tuteja, 2010). Under steady-state conditions, the ROS are scavenged by various antioxidant defense systems: both enzymatic antioxidant(superoxide dismutase, SOD; catalase, CAT; ascorbate peroxidase, APX; glutathione reductase, GR; monodehydroascorbatereductase, MDHAR; dehydroascorbate reductase, DHAR; glutathione peroxidase, GPX;guaiacol peroxidase, POX and glutathione-S- transferase, GST) and non-enzymatic (ascorbate, glutathione, carotenoids, phenolic compounds, proline, glycine betain, sugar, and polyamines) defense systems (Foyer and Noctor, 2005; Desikan et al., 2005; Ahmad et al., 2008; Gill and Tuteja, 2010).
Plant tissue culture techniques are used to grow plants under aseptic and controlled environment for the purpose of both commercial (like mass production) andscientific (like germplasm preservation, plant breeding, physiological,and genetic) studies (www.kitchenculturekit.com). Two of these application areas are important to study ROS homeostasis in plants. The first one of these techniquesis used as a model to induce oxidative stress under controlled conditions via different stressor agents for researching
2. Oxidative stress and Reactive Oxygen Species (ROS)
Reactive Oxygen Species (ROS), is also sometimes called Active Oxygen Species (AOS), or Reactive Oxygen Intermediates (ROI), or Reactive Oxygen Derivatives (ROD),is the term used to describe highly reactive and partially reduced-oxygen forms (Desikan et al., 2005). ROS are produced in many ways in several cellular compartments, including mitochondria, chloroplast, peroxisomes, endoplasmic reticulum, cytoplasm, plasma membrane and apoplast, during normal metabolic processes and due to induction of environmental perturbations, such as drought, salinity, radiation, heavy metals, and herbicides (Desikan et al., 2005). ROS are highly reactive due to the presence of unpaired valence shell electronsand high concentration of ROS can result in non-controlled oxidation in cells, which is defined as oxidative stress, as a result of ROS-attack, cellular compartments, including DNA, protein, membrane lipids may damage (Cassells and Curry, 2001; Desikan et al., 2005).
ROS include a wide range of oxygen-radicals, such as superoxide anion (O2·-), hydroxyl radical (OH·), perhydroxyl radical (HO2·) and hydrogen peroxide (H2O2), they become the sequential reduction of molecular oxygen. Singlet oxygen (1O2), another form of ROS, can be produced by excited-chlorophyll formation in the photosystem II (PSII) reaction center and in the antennae systems. This is the major formation mechanism of 1O2 in plant cells. Insufficient energy dissipations during the photosynthesis, the chlorophylls are excited, which then can lead the formation of chlorophyll (Chl) triplet state. Chl triplet state can react with 3O2 to give up the very reactive 1O2 (Arora et al., 2002; Reddy and Raghavendra, 2006; Gill and Tuteja, 2010). 1O2 has powerful damaging effect on the whole photosynthetic machinery, including chloroplast membrane lipids, proteins and nucleic acids. The primary means of defense within the chloroplast are the carotenolds (CARs) and a-tocopherol (vitamin E), which are located within the thylakold membranes. They are a quencher against damages of 1O2 (Knox and Dodge, 1985). Hossain et al. (2006) and Helaley and El-Hosieny (2011) reported that carotenoid contents increase under salinity stress in various plant species. O2·-, which is generally known as the first ROS to be generated, usually generate with the single electron reduction of O2. The major site of O2·- production is in the photosystem I (PSI) by Mehler Reaction. The generation of O2·- may lead to formation of OH· and 1O2. The reaction of O2·- with Fe+3 may become 1O2 (1), and reduced-form of Fe+2. O2·- can also reduce to H2O2 by SOD (2). HO2˙ is formed from O2˙ˉ by protonation in aqueous solutions. HO2˙ can cross biological membranes and subtract hydrogen atoms from polyunsaturated fatty acids (PUFAs) and lipid hydroperoxides, thus initiating lipid auto-oxidation (Halliwell and Gutteridge, 2000). Additionally, complex I, ubiquinone, and complex III in mitochondrial electron transfer chain (ETC), the other major ROS (H2O2 and O2·-) producing sites in cells (Reddy and Raghavendra, 2006; Gill and Tuteja, 2010). Xanthine oxidase generates O2·- during the catabolism of purines in the peroxisomes, andan increasing production of O2·- is caused certain herbicides, like paraquat, which is known photosynthetic inhibitors. Paraquat (also called methyl violeng) prevents the transfer of electrons from ferredoxin (Fd) in PSI, afterwards increase generation of O2·- with the transfer of electrons from molecular oxygen (Peixoto et al., 2007; Gill and Tuteja, 2010). It is also clear that environmental stress induced the production of O2·-and the other ROS (Arora et al., 2002; Reddy and Raghavendra, 2006; Gill and Tuteja, 2010). H2O2 is produced as a result of dismutation reaction of O2·. This reaction mostly catalyzed by SOD (Arora et al., 2002). H2O2 is formed in the peroxisomesas part of photorespiratory, and also produced from β-oxidation of fatty acids as a by-product.H2O2 is not a free radical, but is participates as an oxidant or a reductant in several cellular metabolic pathways (Reddy and Raghavendra, 2006). By means of transition metals, such as Fe and Cu, further reduction of H2O2 take place OH- and OH·, which are mentioned below as Haber-Weiss/Fenton Reaction (3, 4).OH· is extremely reactive and will potentially react with all biological molecules, such as DNA, proteins, and lipids. If productions of hydroxyl radicals are not eliminated by any enzymatic and non-enzymatic defense mechanisms, overproduction of its ultimately leads to cell death (Desikan et al., 2005; Gill and Tuteja, 2010). As a result of the measurement of ROSusing spectrophotometric, fluorescent dye probeand electron spin resonance (ESR) methods showed that various abiotic stress factors induced ROS formation in a wide range of plant species under
As I mentioned above, an overproduction of ROS can result in non-controlled oxidation in cells, resulting in ROS-attack, which may damage several cellular macromolecules, such as lipid membranes, proteins and DNA (Cassells and Curry, 2001; Desikan et al., 2005).The peroxidation of membrane lipids both cellular and organelles are known as the most damaging factors in all living organisms, including plants. As a result of lipid peroxidation (LPO) some products are formed by PUFAs. One of them is malondialdehyde (MDA). The reactions of MDA with thiobarbituric acid (TBA) produces color product, which is called thiobarbituric acid reactive substances (TBARS). The spectrophotometric measurement of TBARS or MDA generally used as oxidative stress biomarker and also to assess the degree of LPO. Many researchers reported that MDA content increased under several abiotic stress factors, which were induced
Another result of ROS-attack in cells is an increase in protein oxidations. Site specific amino acid modifications, fragmentation of the peptide chain, and aggregation of cross linked reaction products occur in plants as consequence of protein oxidations induced by ROS or by-products of oxidative stress. These reactions are mostly irreversible (Ahmad et al., 2008;Gill and Tuteja, 2010). Various mechanisms can cause protein oxidation, such as the formation of disulfide cross-links and glycoxidation adducts nitration of tyrosine residues, and carbonylation of specific amino acid residues (Oracz et al., 2007). The spectrophotometric measurement of protein carbonyl with dinitrophenylhydrazine (DNPH) method is widely used marker for detection of protein oxidation in biological organisms. Basu et al., (2010) reported that an increasing ratio of protein oxidations were measured in all rice varieties induced drought conditions in tissue culture.
ROS-induced genotoxic damage can induce structural changes in DNA, such as chromosomal rearrangement, strand breaks, base deletions, prymidine dimers, cross-links and base modifications, mutations and other lethal genetic effects (Cassells and Curry, 2001; Ahmad et al., 2008; Gill and Tuteja, 2010). When DNA-lesions are endogenously generated mostly via ROS, it is called spontaneous DNA damage (Ahmad et al., 2008; Gill and Tuteja, 2010). Oxidative stress, as well as effects of damaging which were referred above, also has a great potential creating variability in the plant genome by activating transposons, inducing chromosome breakage/rearrangement, and base mutation and these situations are one of the main reasons of spontaneous mutations in cells (Cassells and Curry, 2001; Gaspar et al., 2002). As I mentioned below, spontaneous mutations are one of the key factors of plant breeding.
Additionally, low concentrations of ROS are key factors to maintain intercellular signal transductions in plants. Further information about ROS, there is an excellent review (Ahmad et al., 2008; Gill and Tuteja, 2010; Karuppanapandian et al., 2011) about this subjects books (Smirnoff, 2005; Rao et al., 2006; Del Rio and Puppo, 2009) published in recent years.
3. Antioxidant defence system
ROS are generated in plant cells by normal cellular metabolism or due to unfavorable environmental conditions such as drought, salinity, heavy metals, drought, herbicides, nutrient deficiency, or radiation. Their productions are controlled by various enzymatic and non-enzymatic antioxidant defense systems. Enzymatic antioxidant defense systems, including CAT, APX, POX, SOD, MDHAR, DHAR and GR and non-enzymatic antioxidant defense systems, including ascorbate, glutathione, carotenoids, phenolic compounds, proline, glycine betain, sugar, and polyamines (Ahmad et al., 2008; Gill and Tuteja, 2010; Karuppanapandian et al., 2011).
3.1. Enzymatic antioxidants
3.1.1. Superoxide dismutase (SOD; EC 1.15.1.1)
Superoxide dismutase, as a metalloenzyme, is the first enzyme of the detoxification processes, which catalyzes O2·- to H2O2 and O2. SODs are classified into three types based on their metal cofactor: Fe-SOD (localized in chloroplasts), Mn-SOD (localized in mitochondria), and Cu/Zn-SOD (localized in chloroplasts, peroxisomes, and cytosol). The activity of SOD isozymes can be detected by negative staining and can be identified on the basis of their sensitivity to KCN and H2O2. The Mn-SOD is resistant to both inhibitors; Cu/Zn-SOD is sensitive to both inhibitors whereas; Fe-SOD is resistant to KCN and sensitive to H2O2 (Ahmad et al., 2008; Gill and Tuteja, 2010; Karuppanapandian et al., 2011). There have been many reports of the increased activities of SOD under abiotic stresses induced with tissue culture techniques in a wide range of plant species, including heavy metals, such asAl, Cd, Cr, and Cu, hyperhydricity, salinity, gamma radiation, and drought (Gallego et al., 2002;Saher et al., 2004; Dewir et al., 2006; Israr et al., 2006; Erturk et al., 2007; Dasgupta et al., 2008; Sivritepe et al., 2008; Gupta and Prasad, 2010; Shehab et al., 2010;Yang et al., 2010; El-Beltagi et al., 2011; Helaly and El-Hosieny, 2011; Lokhande et al., 2011; Sen and Alikamanoglu, 2011;Xu et al., 2011; Patada et al., 2012) on the other hand, Fe-deficiency stress reduced activity of SOD (Lombardi et al., 2003). Advanced-antioxidant defense systems play an important role in plants not only to tolerate environmental stress but also to improve plants against these stresses. Enhanced activities of SOD were observed in various plants to improve tolerance against salinity (Hossain et al., 2006; Hossain et al., 2007; Chen et al., 2011;Helaly and El-Hosieny, 2011), and S-(2-aminoethy)-cysteine AEC (Kim et al., 2004) using
As a result of native polyacrylamide gel electrophoresis (native–PAGE), Chakrabarty et al., (2005) and Dewir et al., (2006) detected that Mn-SOD and Cu/Zn-SOD isoenzymes seem to play a major role in response to hyperhydricity. Additionally, Rahnama and Ebrahimzadeh (2006) and Roy et al., (2006) also reported that against salinity and gamma radiation Mn-SOD and Cu/Zn-SOD seem to play a major role in the potato and
3.1.2. Catalase (CAT, EC 1.11.7.6)
CAT is a tetrameric heme-containing enzyme that catalyzes dismutation reactions of H2O2 into H2O and O2 and is indispensable for ROS detoxification during stress conditions. CAT is also important in the removal of H2O2 generated in peroxisomes during the β-oxidation of fatty acids, photorespiration, and purine catabolism (Ahmad et al., 2008; Gill and Tuteja, 2010; Karuppanapandian et al., 2011). Various abiotic stresses induced CAT activities under
3.1.3. Guaiacol Peroxidase (POX, EC 1.11.1.7)
POX is a heme-containing enzyme, like CAT. POX prefers aromatic electron donors such as guaiacol and pyragallol to catalyze H2O2, and many researchers reported that excess POX activities were measured in a wide range of plant varieties under abiotic stress conditions induced with
There have been many reports of the changes in POX isoenzymes depending considerably upon plant species and abiotic stresses under tissue culture conditions. NaCl stress stimulated new POX isoenzyme band in Agria and Kennebec potato cultivar (50 mM) and in
3.1.4. Halliwell-Asada Cycles’ Enzymes (Ascorbate peroxidase (APX, EC 1.11.1.1), Monodehydroascorbate reductase (MDHAR, EC 1.6.5.4), Dehydroascorbate reductase (DHAR, EC 1.8.5.1) and Glutathione reductase (GR, EC 1.6.4.2))
The Ascorbate-Glutathione Cycle, sometimes called Halliwell-Asada Cycle, is another metabolic pathway that detoxifies H2O2. This is located in the cytosol, mitochondria, chloroplasts and peroxisomes in plants and it may have a more crucial role in the management of ROS during stress (Noctor and Foyer, 1998). The cycle involves the antioxidant metabolites: ascorbate, glutathione and NADPH and the enzymes linking these metabolites, involving APX, MDHAR, DHAR and GR. In the first step of this pathway, H2O2is reduced to H2O and monodehydroascorbate (MDHA) by APX using ascorbate as the electron donor. The oxidized ascorbate (MDHA) is regenerated by MDHAR. MDHAR is a flavin adenin dinucleotide (FAD) enzyme which uses NAD(P)H directly to recycle ascorbate, and dehydroascorbate (DHA). After, DHA is reduced to ascorbate by dehydroascorbate reductase (DHAR) using of glutathione (GSH) as the electron donor. As a result of this reaction oxidized glutathione (GSSG)occur. Finally GSSG is reduced to GSH by glutathione reductase (GR) using NADPH as electron donor (Noctor and Foyer, 1998; Ahmad et al., 2008; Gill and Tuteja, 2010; Karuppanapandian et al., 2011). Enhanced expression of Halliwell-Asada Cycles’ enzymes in plants has been demonstrated during different stress conditions. Saher et al., (2004) reported that hyperhydric stress increased Halliwell-Asada Cycle’s enzyme activities (APX, MDHAR, DHAR and GR) in
Chakrabarty et al., (2005) reported that after the native–PAGE analysis, five APX isoenzyme bands were observed on the gels in hyperhydric apple leaves. Three of them (APX-1, APX-4 and APX-5) only appeared in hyperhydric apple leaves. New APX and GR isoenzyme bands were also induced during the As-stress both shoots and roots, for APX, and only roots, for GR, in rice tissue culture, respectively (Shri et al., 2009).
3.1.5. Glutathione Peroxidases (GPX, EC 1.11.1.9)
GPXs are a large family of diverse isozymes that use GSH to reduce H2O2, besides this situation GPX also has more crucial role for lipid peroxidation process, and therefore helps plant cells from oxidative stress (Gill and Tuteja, 2010). Millar et al., (2003) reported that GPX includes a family of seven related proteins in cytosol, chloroplast, mitochondria and endoplasmic reticulum. Hyperhydric stress increased GPX activity in
3.1.6. Glutathione S-transferases (GST, EC 2.5.1.18)
GSTs catalyse the conjugation of electrophilic xenobiotic substrates with the tripeptide glutathione (GSH; γ-glu-cys-gly). Plant GST gene families are large and highly diverse, like GPXs. GSTs are generally cytoplasmic proteins, but microsomal, plastidic, nuclear and apoplastic isoforms has also been reported. They are known to function in herbicide detoxification, hormone homeostasis, vacuolar sequestration of anthocyanin, tyrosine metabolism, hydroxyperoxide detoxification, regulation of apoptosis and in plant responses to biotic and abiotic stresses. GSTs have the potential to remove cytotoxic or genotoxic compounds, which can react or damage the DNA, RNA and proteins (Gill and Tuteja, 2010). Enhanced activities of GST in potato tuber callus was demonstrated during 2,4-D and Dicamba treatments (Peixoto et al., 2007) and also in paraquat- tolerant poplar clones, which were improved using
3.2. Non-enzymatic antioxidants
Apart from the enzymatic defense system, several non-enzymatic antioxidant defense mechanisms also play an important role in the response of plant stress tolerance, such as ascorbate, glutathione, carotenoids, phenolic compounds, proline, glycinebetain, sugar, and polyamines.
Two of them, ascorbate and glutathione are crucial metabolites in plants which are considered as most important intracellular defense against ROS induced oxidative damage. Ascorbate can directly scavenge 1O2, O2·- and ·OH and by regenerate a-tocopherol from tocopheroxyl radical. It also acts as co-factor of violaxanthin de-epoxidase, thus sustaining dissipation of excess excitation energy. Glutathione, like ascorbate, plays a pivotal role in several physiological processes, including regulation of sulfate transport, signal transduction, conjugation of metabolites, detoxification of xenobiotics and the expression of stress-responsive genes. Both of them are also main components of the Halliwell-Asada Cycle (Gill and Tuteja, 2010). Shehab et al., (2010) and El-Beltagi et al., (2011) reported that ascorbate and glutathione contents were increased under PEG-induced drought stress and low doses gamma radiation in rice and
Carotenoids are a lipid soluble antioxidant, which are considered as potential scavengers of ROS and lipid radicals. They are known major antioxidants in biological membranes for protection of membrane stability against lipid peroxidation, including quenching or scavenging ROS like 1O2. Carotenoids have several major functions such as preventing membranes for lipid peroxidation. One of them, they act as energetic antenna, absorb light at wavelength between 400 and 550 nm and transfer it to the Chl. Second, they protect the photosynthetic apparatus by quenching a triplet sensitizer (Chl3), 1O2 and other harmful free radicals which are naturally formed during photosynthesis. Third, they are important for the PSI assembly and the stability of light harvesting complex proteins as well as thylakoid membrane stabilization (Gill and Tuteja, 2010). Helaly and El-Hosieny, (2011) reported that carotenoid contents increased in
Accumulating osmotic adjustment, sometimes is called osmoprotectant, in their structures is another crucial mechanism in many plant species in response to environmental stress, including proline (amino acids), glycinebetain (quaternary ammonium compounds) and sugars (mannitol, D-ononitil, trehalose, sucrose, fructan). Proline and glycinebetain act as osmoprotectants by stabilizing both the quaternary structure of proteins and the structure of membranes, Proline also acts a metal chelator, an inhibitor of LPO, and OH· and 1O2 scavenger (Arshaf and Harris, 2004). Enhanced osmoprotectant contents have been demonstrated in plants during different stress conditions by many researchers. Patada et al. (2012) reported that glycinebetain, proline and reduced sugar contents increased in embryonic sugarcane callus under PEG and NaCl treatment. In another study, Lokhande et al. (2010) observed that glycinebetain, proline and soluble sugar contents enhansed in
Phenolic compounds, which are often referred to as secondary metabolites and functions of most of them have still poorly understood, including flavonoids, tannins, anthocyanins, hydroxycinnamate esters, and lignin, are abundant in plant tissues. Many secondary metabolites play widely important role from as defensive agents against pathogens to general protection against oxidative stress using as electron donors for free radical scavenging (Grace, 2005). Phenylalanine ammonia lyase (PAL) activity is one of the main enzymes in the synthesis of phenolic compounds, and phenolic contents were increased under PEG-induced drought stress in rice callus culture (Shehab et al., 2010). PAL activities also increased in
Franck et al., (2004) and Ghnaya et al. (2011) reported that polyamine contents increased in hyperhydric
Additionally, measuring free radical scavenging or quenching capacities in cells are the other techniques the detection of total non-enzymatic antioxidant activity. Various methods have been used for measuring total antioxidant activities in biological systems. The increasing ratios of total antioxidant capacity were measured under different abiotic stress conditions induced with tissue culture techniques using 2,2-azino-bis (3-ethyl-benzothiazoline-6-sulfonic acid (ABTS) (Cui et al., 2010); 2,2-diphenyl-1-picrylhydrazyl (DPPH) (Hossain et al., 2006; Cui et al., 2010; Basu et al., 2010; Zamora et al., 2010) and ferric reducing antioxidant power (FRAP) (Sotiropoulos et al., 2006; Chatzissavvidis et al., 2008) methods, respectively.
4. Plant tissue culture
Plant tissue culture, as an alternatively known cell, tissue and organ culture or
4.1. Induced-oxidative stress conditions in plant tissue culture and antioxidant defense systems
Nowadays, one of the most serious problems is the influence of environmental stress factors on plants which are exacerbated day-by-day through anthropogenic effects. Thus, plant growth, development and the yield performance of plants is adversely affected. These negative results forced humans to find new solutions to minimize these problems.
In recent years,
4.2. Antioxidant activities of in vitro selected plants under abiotic stress conditions
embryogenic callus | 0, and 20% (w/v)PEG 8000 | SOD, CAT, APX, TBARS, TSS and TRS, Pro., PC, and GB | spectrophotometric | Patada et al. (2012) | |
callus | 0,500,and1000 mM Mannitol | SOD, CAT, PPO, H2O2, POX, MDA, PC, CARs, and Pro. | spectrophotometric | Torabi and Niknam (2011) | |
adventitious roots | 0, 1, 3, 5, 7, and 9% (w/v) Sucrose | DPPH, ABTS, Pro., H2O2, MDA, Flavonoid, Phenol and chlorogenic acid, Hypericin and Residual Sugars | spectrophotometric | Cui et al. (2010) | |
seedlings | 0, and 20% (w/v) PEG-6000 | SOD, CAT, POX, H2O2, LOX, MDA, PP PO, DPPH, Antocyanin Flavonoid, and Phenol | spectrophotometric | Basu et al. (2010) | |
shoots | PEG-8000 | DPPH, APX, CAT, POX, GR, Proline, H2O2, MDA, Ascorbate, Flavonoid, PP, and Phenol | spectrophotometric | Zamora et al. (2010) | |
callus | 0, 5, 10, 15, and 20% PEG | H2O2, MDA, GSH, AsA, SOD, APX, CAT GR, PAL, TSS and AA, and Phenol | spectrophotometric | Shehab et al. (2010) | |
shoot tips | 0, 1, 2, and 4% PEG 8000 | SOD, CAT, POX, APX, GR, MDA, PP, and Pro. | spectrophotometric | Sivritepe et al. (2008) | |
shoot tips | 0, and 576 mM Mannitol 0, and 562.5 mM Sorbitol | SOD, CAT, POX, FRAP, H2O2, Pro., PP, and MDA | spectrophotometric and isoenzyme variations | Molassiotis et al. (2006) | |
shoots | 0, and 300 mM Mannitol | POX, MDA, and H2O2 | spectrophotometric and isoenzyme variations | Radić et al.(2006) | |
shoot tips | 0, and 40% PEG 6000 | SOD, CAT, APX, GR, and MDA | spectrophotometric | Chai et al. (2005) |
callus | 100 mM | SOD, and CAT | spectrophotometric | Kusvuran et al. (2012) | ||
callus | 150 mM | SOD, CAT, APX, TBARS, TSS and TRS, Pro., PC, and GB | spectrophotometric | Patada et al. (2012) | ||
callus | 0, 100, 300, and 600 mM | SOD, CAT, PPO, H2O2, POX, MDA, PC, CARs, and Pro. | spectrophotometric | Torabi and Niknam,(2011) | ||
mature embryos | 0, 50, 100, 150, 200, and 250mM | PP, SOD, POX, and CAT | spectrophotometric and isoenzyme variations | Sen and Alikamanoglu, (2011) | ||
axillary shoots | 0, 200, 400, and 600 mM | SOD, CAT, APX, TSS, Pro., GB, and MDA | spectrophotometric | Lokhande et al. (2011) | ||
callus | 0, 100, 200 and400 mM | SOD, CAT, APX, TSS, Pro., and GB | spectrophotometric | Lokhande et al. (2010) | ||
callus | 0, 50, 100 and 200 mM | SOD, POX, CAT, APX, H2O2, and NADPH-oxidase | spectrophotometric | Yang et al. (2010) | ||
seedlings | 0, 20, 40, 60, 80 and 160 mM | MDA, PP, CARs, PC, and Proline, | spectrophotometric | Ayala Astorga and Alcarez-Melendez, (2010) | ||
shoots | 0, 15, 30, 45, 60, 75, and 100 mM | SOD, CAT, POX, MDA, Pro., Phenol, PP, and Sugar | spectrophotometric | Garg, (2010) | ||
suspension cells | 0, 50, 100,and 150 mM | MDA, SOD, and ROS mes. | isoenzyme variations and transcription analysis | Azevedo et al. (2009) | ||
callus | 0, 50, 100, 150, 200, and 250mM | Sucrose, Treholase Pro., Total Flavonoid and GB | spectrophotometric | Zhao et al. (2009) | ||
shoot apices and callus | 0, 20, 40, 60, 80, 100, 120 and 140 mM | SOD, CAT, POX, and PC | spectrophotometric | Sajid and Aftab,(2009) | ||
callus | 0, 20, 40, 60, 80, 100 mM | SOD, POX, PC, and Pro. | spectrophotometric and isoenzyme variations | Kumar et al. (2008) | ||
shoot apexes | 0, 0.5, and 1% | SOD, POX and CAT | spectrophotometric | Dasguptan et al. (2008) | ||
shoot tips | 0, 30, and 60 mM (NaCl and CaCl2) | FRAP, CAT, POX, and PP | spectrophotometric and isoenzyme variations | Chatzissavvidis et al. (2008) | ||
shoots | 0, 35, 100, and 200 mM (NaCl) 0, 5, and 10 mM (CaCl2) | Sugar, and Proline | spectrophotometric | Sotropoulos, (2007) | ||
Sweet chery rootstock Gisela 5 | shoot tips | 0, 50, 100, and 150 mM | SOD, POX, CAT, APX, GR, MDA and Proline | spectrophotometric | Erturk et al. (2007) | |
seedlings | 0, 25, 50, 100, and 200 mM | H2O2, GSH, Ascorbate, SOD, CAT, GR, PP, G6PHD, NADP-ICDH, and FNR | spectrophotometric,immunofluores., isoenz. and transcript. | Valderrama et al. (2006) | ||
shoot tips | 0, and 240 mM NaCl 0, and 220 mM KaCl | SOD, CAT, POX, FRAP, H2O2, FRAP, PP, Proline, and MDA | spectrophotometric and isoenyme variations | Molassiotis et al. (2006) | ||
seeds and callus | 0, 50, 100, 150; and 200 mM | CAT, POX, PPO, PC, and Proline | spectrophotometric and isoenyme variations | Niknam et al. (2006) | ||
internodes | 0, 50, 100 and 150 mM | SOD and POX | spectrophotometric and isoenyme variations | Rahnama and Ebrahimzadeh (2006) | ||
cell suspension | 0, 50, 100, 150, 200, 300 and 400 mM | SOD, POX, CAT, Proline and MDA | spectrophotometric | Ferreira and Lima-Costa (2006) | ||
shoots | 0, 150, 300, 450 and 600 mM | POX, MDA, and H2O2 | spectrophotometric and isoenyme variations | Radić et al. (2006) | ||
nodes | 0, 50, 75 and 100 mM | SOD, CAT, POX and APX, | spectrophotometric and isoenyme variations | Rahnama and Ebrahimzadeh (2005) | ||
Eucalyptus camadulensis Dehnh. clones | shoots | 0, 50, 100 mM | Proline, PP | spectrophotometric | Woodward and Bennett (2005) |
callus | Cd | 0, 150 and 250μM | APX, CAT, POX, Thiols, and Phytochelatins | spectrophotometric | Iori et al. (2012) | ||
thin cell layers | Zn | 0-1 mM | POX, MDA, PP, CARs, and Polyamines | spectrophotometric | Ghnaya et al.(2011) | ||
callus | Cu | 0, 0.05, 0.1, 0.2, 0.6, 0.8, and 1 mM | SOD, CAT, POX, MDA, PP, ROS mes. (H2O2 and O2-• ) | spectrophotometric | Xu et al. (2011) | ||
seeds | Cd | 0, 0.05, 0.1, 0.2 and 0.4 mM | SOD, CAT, POX, PPO, and PAL | spectrophotometric and isoenzyme variations | Zheng et al.(2010) | ||
embryos | Pb | 0, 100, 200, 400 and 800μM | SOD, CAT, POX, and PAL | spectrophotometric and isoenyme variations | Jiang et al. (2010) | ||
embryos | Zn | 0, 0.25, 0.5, 1, 2 and 3 mM | SOD, CAT, POX, and PAL | spectrophotometric and isoenyme variations | Luo et al. (2010) | ||
seeds | As(III) and As(V) | 0, 50 and 100μM As(III) 0, 100 and 500μM As(V) | SOD, APX, GR, GSSG, POX, and MDA | spectrophotometric and isoenyme variations | Shri et al. (2009) | ||
embryos | Ni | 0, 100, 200, 400 and 800 μM | SOD, CAT, POX, and PAL | spectrophotometric and isoenyme variations | Yan et al. (2008) | ||
seeds | Cd | 0, 50, 100, 200 and 300 μM | CAT, POX and MDA | spectrophotometric | Kumar et al.(2008) | ||
suspension cultures | Cd and Zn | 0, 12.5, 25, 50, 100 and 200μM (Cd) 0, 50, 100, 200, 400 and 800μM (Zn) | Thiol, AA, and Polyamines | spectrophotometric | Thangavel et al.(2007) | ||
Medicago sativa L. cv. Aragon | Cd and Hg | 0, 3, 10 and 30μM | H2O2 , Ascorbate, GSH, APX, and SOD, | spectrophotometric, isoenz., fluor., and transcrit. | Villasante et al.(2007) | ||
callus | Cd | 0, 10, 25, 50, 100 and 250μM | SOD, APX, GR, GSSG, and GSH | spectrophotometric | Israr et al. (2006) | ||
shoots | B | 0.1, 0.5, 1, 3, and 6 mM | SOD, CAT, POX , PP, and FRAP | spectrophotometric | Sotiropoulos et al. (2006) | ||
callus | Cd | 150 μM | MDA, GSH, GSSG, Phytochelatins, and ROS mes., | spectrophotometric and fluorescein dye | Gallego et al. (2005) | ||
callus | Cd+3, Al+3, Cr+3 | 150 μM | POX, SOD, CAT, APX GR, TBARS, GSH, GSSG, Ascorbate, ROS mes, and Dehydroascorbate, Phytochelatins | spectrophotometric | Gallego et al. (2002) | ||
callus | Cd | 0,0.01, 0.02, 0.05, 0.1, 0.2, 0.5 and 1 mM | SOD and CAT | spectrophotometric and isoenyme variations | Fornazier et al. (2002) |
callus | gamma radiation (60Co) | 0, 5, 10, 15 and 20Gy | MDA, AsA, GSH, SOD, PAL, TSS, AA, Phenol, Flavonoid, H2O2 and O2-.mes. PAL, APX, CAT, and POX | spectrophotometric | El-Beltagi et al. (2011) | |
callus | hyperhydricity | different culture systems | MDA, AsA, SOD, APX, CAT, and POX | spectrophotometric and isoenyme variations | Gupta and Prasad, (2010) | |
calli | herbicides | paraquat, 2,4-D and dicamba | SOD, CAT, GR and GST | spectrophotometric | Peixoto et al. (2007) | |
callus | gamma radiation (60Co) | 0, 20, 50, 100 and 200 Gy | SOD, and POX | isoenyme variations | Roy et al. (2006) | |
Apple “M9 EMLA” | nodal segments | hyperhydricity | bioreactor culture | SOD, APX, GR, GPX, MDHAR, DHAR, and ROS mes. | spectrophotometric, flour. and isoenyme variations | Chakrabarty et al. (2006) |
inflorescences | hyperhydricity | bioreactor culture | SOD, APX, CAT, POX, GR, GST, GPX, MDHAR,DHAR, GSH, GSSG, LOX, and MDA | spectrophotometric and isoenyme variations | Dewir et al. (2006) | |
shoots | Fe-deficiency | MS (+Fe) control, MS (-Fe),MS (5 mM NaHCO3 + 0.5 gl-1 CaCO3) (pH 6.9) and MS (10 mM NaHCO3 + 0.5 gl-1 CaCO3 ) (pH 7.3) | CAT, POX, H2O2 and FRAP | spectrophotometric and isoenyme variations | Molassiotis et al. (2005) | |
shoots | hyperhydricity | changing concentration of agar from 0.8% to 0.58% | SOD, CAT, APX, POX, GR, MDHAR, DHAR, LOX, PAL, H2O2 lignin, AsA,PP, Ethylen, and MDA | spectrophotometric | Saher et al. (2004) | |
shoots | hyperhydricity | changing from 0.8% agar to 0.25% gelrit) | Proline, GPX, Ethylene, and Polyamines | spectrophotometric | Franck et al. (2004) | |
seeds | Fe-deficiency | 0, 13 and 27.8mgl-1FeSO4 | APX, CAT, MDA, GSH, PP, and EPR | spectrophotometric and isoenyme variations | Mohamed and Aly, (2004) | |
shoots | Fe-deficiency | control, MS (+Fe) and MS (+ 1mM KHCO3) | PP,CARs, CAT, and SOD | spectrophotometric and transcript. | Lombardi et al. (2003) |
shoot tips | gamma radiation (137Cs) | drought (PEG) | SOD, APX, CAT and POX | spectrophotometric and isoenyme variations | Sen and Alikamanoglu, (2012) | |
protoplasts | gamma radiation (60Co) | NaCl | SOD, APX, CAT, POX, GR, H2O2, MDA, Pro, PP, TS, GB, and Phenols | spectrophotometric | Helaly andEl-Hosieny, (2011) | |
nods | gamma radiation (137Cs) | drought (PEG) | SOD, APX, CAT and POX | spectrophotometric | Alikamanoglu et al.,(2011) | |
embryogenic callus | gamma radiation (60Co) | NaCl | SOD, CAT, POX and Pro. | spectrophotometric | Chen et al. (2011) | |
nodes | gamma radiation (137Cs) | NaCl | SOD, APX, CAT and POX | spectrophotometric | Alikamanoğlu et al.,(2009) | |
plantlets | sucrose-induced | atrazine | SOD, APX, CAT, DHAR, MDAR, GR, H2O2, 1O2, O2.-, CARs, PP, and ROS-scavenging systems | spectrophotometric, fluorescent dying, and transcriptomic analy. | Ramel et al., (2009) | |
Poplar clones | leaf petioles | paraquat | GR, APX, GST, and LOX | spectrophotometric and transcriptomic analy. | Bittsánszky et al. (2008) | |
callus | NaCl | SOD, APX, GR, and Pro. | spectrophotometric | Hossain et al., (2007) | ||
callus | NaCl and drought | SOD, CAT, and Pro. | spectrophotometric | Lu et al., (2007) | ||
shoot | EMS | NaCl | SOD, APX, DHAR, MDAR, GR, H2O2, DPPH, PP, AsA PP, CARs, and Pro. | spectrophotometric and isoenzyme variations | Hossain et al., (2006) | |
callus | NaCl | Pro., and TSS | spectrophotometric | Gandonou et al. (2006) | ||
callus | drought (PEG) | Pro., and Charbonhydrates | spectrophotometric | Hassan et al., (2004) | ||
callus | gamma radiation (60Co) | AEC res. | SOD, APX and AA | spectrophotometric and proteomic analysis | Kim et al. (2004) | |
seeds | drought (PEG) | CAT, GR and Pro. | spectrophotometric | Safarnejad, (2004) | ||
cell suspension | paraquat and Cd | SOD, APX, CAT and POX | spectrophotometric and isoenzyme variations | Kopyra and Gwozdz, (2003) | ||
callus | drought (Mannitol) | Pro., and TSS | spectrophotometric | Mohamed et al., (2000) |
5. Conclusion
The overproduction of ROS in plants is stimulated by environmental stressors as well as many metabolic reactions, such as photosynthesis, photorespiration, and respiration. All of these ROS are toxic to biological molecules and generally lead to non-controlled oxidation in cellular macromolecules, such as lipid autocatalytic peroxidation, protein oxidation or DNA-lesions. These irreversible damages of cellular macromolecules cause many cases in plants from mutations to cell death. Plants possess sophisticated-antioxidant defense mechanisms, including antioxidant enzymes and molecules that can protect cells from oxidative damage and maintain ROS homeostasis. Besides causing damage, ROS can also participate in signal transduction. Despite all these knowledge about ROS, how to maintain balance between these oxidant and antioxidant properties in plants have still poorly been understood. Plant tissue culture techniques are performed under aseptic and controllable environmental conditions for this reason allows various opportunities to study details of this balance. Controlled stress in
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