Protective effect of ascorbic acid and glutathione on AlCl3-inhibited growth of rice roots

The effect of AlCl₃ on the antioxidant system of rice roots and the role of applied antioxidants ascorbic acid (AsA) and glutathione (GSH) in AlCl₃-inhibited growth of rice roots were investigated. AlCl₃ treatment resulted in a rapid inhibition of root growth but had no effect on lipid peroxidation and antioxidative enzyme activities in rice roots. AlCl₃ treatment resulted in lower content of H₂O₂, AsA, and GSH than in controls. Exogenous AsA or GSH counteracted growth inhibition of rice roots induced by AlCl₃. AlCl₃ treatment increased syringaldazine peroxidase (SPOX) activities and lignin content in rice roots. Exogenous AsA or GSH prevented the decrease in H₂O₂ content and the increase in SPOX activities and lignin content in rice roots caused by AlCl₃. Results suggest that lignification induced by low AsA or GSH content may explain the mechanism of Al-inhibited growth of rice roots.     

Lipid Peroxidation Induced by Phenolics in Conjunction with Aluminum Ions

Using the whole plant and model systems, we demonstrate that the aluminum ions (Al³⁺) stimulate phenolic-dependent lipid peroxidation. Lipid peroxidation in barley (Hordeum vulgare L. cv. Donor) roots was 30 % higher under AlCl₃ treatment than without Al. Major decomposition product of lipid peroxidation was 4-hydroxynonenal (4-HNE) but not thiobarbituric acid reactive substances (TBARS), a widely used markers for lipid peroxidation. Similarly, AlCl₃ stimulated lipid peroxidation of soybean liposomes in the presence of chlorogenic acid (CGA) and H₂O₂/horseradish peroxidase system which can oxidize phenolics. Al³⁺ was found to enhance lipid peroxidation induced by oxidized CGA. Intermediates of lignin biosynthesis in plants, including p-coumaric acid, ferulic acid, sinapic acid and coniferyl alcohol, also showed similar effects. These results suggest that Al³⁺ has a potential to induce oxidative stress in plants by stimulating the prooxidant nature of endogenous phenolic compounds.     

Nickel-induced Inhibition of Wheat Root Growth is Related to H2O2 Production, but not to Lipid Peroxidation

Effects of exogenous nickel (Ni: 10 and 200 μM) on growth, mitotic activity, Ni accumulation, H₂O₂ content and lipid peroxidation as well as the activities of various antioxidative enzymes, such as superoxide dismutase (SOD), catalase (CAT), ascorbate peroxidase (APX) and glutathione peroxidase (GSH-Px) were investigated in wheat roots. A considerable Ni accumulation in the roots occurred at both the concentrations. Although Ni at 10 μM did not have any significant effect on root growth, it strongly inhibited the root growth at 200 μM. Mitotic activity in the root tips was not significantly affected by exposure of the seedlings to 10 μM Ni; however, it was almost completely inhibited at 200 μM treatment. Ni stress did not result in any significant changes in CAT and APX activities as well as lipid peroxidation. However, H₂O₂ concentration increased up to 82% over the control in the roots of seedlings exposed to 200 μM Ni. There was a significant decline in both SOD (50%) and GSH-Px (20–30%) activities in the roots when the seedlings were treated with 200 μM Ni. The results indicated that a strong inhibition of wheat root growth caused by Ni stress was not due to enhanced lipid peroxidation, but might be related to the accumulation of H₂O₂ in root tissue.     

Effects of aluminum on superoxide dismutase and peroxidase activities,and lipid peroxidation in the roots and calluses of soybeans differing in aluminum tolerance

The seedlings of two soybean genotypes, Al-tolerant PI 416937 (PI) and Al-sensitive Young, were cultured in the solution containing 0, 25 or 50 μM Al (AlCl₃·6H₂O) for 24, 36 or 48 h in the hydroponics, and the calluses induced from two genotypes were cultured in medium containing 0, 10, 50 or 100 μM Al for 5, 10 or 15 days, respectively. The effects of Al on growth of seedling roots and calluses, antioxidant enzyme activities of superoxide dismutase (SOD) and peroxidase (POD) and lipid peroxidation were investigated. Under Al stress, PI was more tolerant to Al toxicity than Young at both intact plant and tissue levels and lower concentrations of Al significantly stimulated the root and callus growth of PI. Al application enhanced the activities of SOD and POD and lipid peroxidation in both roots and calluses of two genotypes. Although the differences of SOD activities between two genotypes in response to Al toxicity depended on Al concentration and durations of treatment, SOD activities in the roots of PI were higher than those in the roots of corresponding Young in the presence of Al for 36 or 48 h. Meanwhile, the POD activities in PI roots increased as the Al levels and durations of treatment increased, significantly higher than those in the corresponding Young roots. Moreover, Al-treated PI had significantly lower lipid peroxidation than Young at both root and callus levels. These results suggest that the enhanced antioxidant-related enzyme activities and reduced lipid peroxidation in PI might be one of Al-tolerant mechanisms.     

Cadmium toxicity of rice leaves is mediated through lipid peroxidation

Oxidative stress, in relation to toxicity of detached rice leaves,caused by excess cadmium was investigated. Cd content inCdCl₂-treated detached rice leaves increased with increasingdurationof incubation in the light. Cd toxicity was followed by measuring the decreasein chlorophyll and protein. CdCl₂ was effective in inducing toxicityand increasing lipid peroxidation of detached rice leaves under both light anddark conditions. These effects were also observed in rice leaves treated withCdSO₄, indicating that the toxicity was indeed attributed to cadmiumions. Superoxide dismutase (SOD), ascorbate peroxidase (APOD), and glutathionereductase (GR) activities were reduced by excess CdCl₂ in the light.The changes in catalase and peroxidase activities were observed inCdCl₂-treated rice leaves after the occurrence of toxicity in thelight. Free radical scavengers reduced CdCl₂-induced toxicity and atthe same time reduced CdCl₂-induced lipid peroxidation and restoredCdCl₂-decreased activities of SOD, APOD, and GR in the light. Metalchelators (2,2-bipyridine and 1,10-phenanthroline) reducedCdCl₂ toxicity in rice leaves in the light. The reduction ofCdCl₂ toxicity by 2,2-bipyridine (BP) is closely associatedwith a decrease in lipid peroxidation and an increase in activities ofantioxidative enzymes. Furthermore, BP-reduced toxicity of detached riceleaves,induced by CdCl₂, was reversed by adding Fe²⁺ orCu²⁺, but not by Mn²⁺ or Mg²⁺.Reduction of CdCl₂ toxicity by BP is most likely mediated throughchelation of iron. It seems that toxicity induced by CdCl₂ mayrequire the participation of iron.     

Changes in ammonium ion content and glutamine synthetase activity in rice leaves caused by excess cadmium are a consequence of oxidative damage

Ammonium ion accumulation and the decrease in glutamine synthetase (GS)activity induced by CdCl₂ were investigated in relation to lipidperoxidation in detached rice leaves. CdCl₂ was effective inincreasing ammonium ion content, decreasing GS activity and increasing lipidperoxidation. Free radical scavengers (glutathione, thiourea, sodium benzoate)and an iron chelator (2,2-bipyridine) were able to inhibit the decreasein GS activity and ammonium ion accumulation caused by CdCl₂ and atthe same time inhibit CdCl₂-induced lipid peroxidation. Paraquat,which is known to produce oxygen radicals, decreased GS activity, increasedammonium ion content, and increased lipid peroxidation. GS1 appears to be thepredominant isoform present. Excess Cd caused a decrease in GS1 but not in GS2in detached rice leaves. An increase in lipid peroxidation preceded ammoniumionaccumulation and the decrease in GS1 activity. These results suggest that thedecrease in GS activity and the accumulation of ammonium ions in detached riceleaves are a consequence of oxidative damage caused by excess Cd.     

Plant shading increases lipid peroxidation and intensifies senescence-induced changes in photosynthesis and activities of ascorbate peroxidase and glutathione reductase in wheat

Plants of spring wheat (Triticum aestivum L. cv. Saxana) were grown during the autumn. Over the growth phase of three leaves (37 d after sowing), some of the plants were shaded and the plants were grown at 100 (control without shading), 70, and 40 % photosynthetically active radiation. Over 12 d, chlorophyll (Chl) and total protein (TP) contents, rate of CO₂ assimilation (P _N), maximal efficiency of photosystem 2 photochemistry (F_V/F_P), level of lipid peroxidation, and activities of antioxidative enzymes ascorbate peroxidase (APX) and glutathione reductase (GR) were followed in the 1_st, 2_nd, and 3_rd leaves (counted according to their emergence). In un-shaded plants, the Chl and TP contents, P _N, and F_V/F_P decreased during plant ageing. Further, lipid peroxidation increased, while the APX and GR activities related to the fresh mass (FM) decreased. The APX activity related to the TP content increased in the 3_rd leaves. The plant shading accelerated senescence including the increase in lipid peroxidation especially in the 1^st leaves and intensified the changes in APX and GR activities. We suggest that in the 2_nd and 3_rd leaves a degradation of APX was slowed down, which could reflect a tendency to maintain the antioxidant protection in chloroplasts of these leaves.     

Iron induction of lipid peroxidation and effects on antioxidative enzyme activities in rice leaves

Lipid peroxidation in relation to toxicity of detached rice leavescaused by excess iron (FeSO₄) was investigated. ExcessFeSO₄, which was observed to induce toxicity, enhanced the contentoflipid peroxidation but not the content of H₂O₂.Superoxidedismutase activity was reduced by excess FeSO₄. Ascorbate peroxidaseand glutathione reductase activities were increased by excess FeSO₄.Free radical scavengers, such as mannitol and reduced glutathione, inhibitedexcess iron-induced toxicity and at the same time inhibited excessiron-enhancedlipid peroxidation, suggesting that lipid peroxidation enhanced by excess ironis mediated through free radicals.     

Differences in Cowpea Root Growth Triggered by Salinity and Dehydration are Associated with Oxidative Modulation Involving Types I and III Peroxidases and Apoplastic Ascorbate

The aim of this work was to investigate the balance between the activities of ascorbate peroxidase (APX) and phenol peroxidases (POD) and cowpea root growth in response to dehydration and salt stress. Root growth and indicators of oxidative response were markedly changed in response to salinity and dehydration. Salt treatment strongly inhibited root elongation, which was associated with an increase in lignin content and a significant decrease in the concentrations of apoplastic hydrogen peroxide (H₂O₂) and ascorbate. In conditions of extreme salinity, cytosol–APX activity was significantly decreased. In contrast, cell-wall POD activity was greatly increased, whereas lipid peroxidation was unchanged. These results indicate that POD could be involved in both H₂O₂ scavenging and the inhibition of root elongation under high salinity. In contrast, dehydration stimulated primary root elongation and increased lipid peroxidation and apoplastic ascorbate content, but it did not change APX and POD activities or H₂O₂ concentration. When cowpea roots were subjected to salinity followed by dehydration, the water and pressure potentials were decreased, and lipid peroxidation was markedly increased, highlighting the additive nature of the inhibitory effects caused by salt and dehydration. The proline concentration was markedly increased by dehydration alone, as well as by salt followed by dehydration, suggesting a possible role for proline in osmotic adjustment. Salinity and dehydration induce contrasting responses in the growth and morphology of cowpea roots. These effects are associated with different types of oxidative modulation involving cytosolic-APX and cell-wall POD activities and apoplast H₂O₂ and ascorbate levels.     

Arsenic-induced root growth inhibition in mung bean (<Emphasis Type="Italic">Phaseolus aureus</Emphasis> Roxb.) is due to oxidative stress resulting from enhanced lipid peroxidation

Arsenic (As) toxicity and its biochemical effects have been mostly evaluated in ferns and a few higher plants. In this study, we investigated the effect of As (10.0 and 50.0 μM) on seedling growth, root anatomy, lipid peroxidation (malondialdehyde and conjugated dienes), electrolyte leakage, H₂O₂ content, root oxidizability and the activities of antioxidant enzymes in mung bean (Phaseolus aureus Roxb.). Arsenic significantly enhanced lipid peroxidation (by 52% at 50.0 μM As), electrolyte leakage and oxidizability in roots. However, there was no significant change in H₂O₂ content. Arsenic toxicity was associated with an increase in the activities of superoxide dismutase (SOD), guaiacol peroxidase (GPX) and glutathione reductase (GR). In response to 50.0 μM As, the activities of SOD and GR increased by over 60% and 90%, respectively. At 10.0 μM As, the activity of ascorbate peroxidase (APX) increased by 83%, whereas at 50.0 μM it declined significantly. The catalase (CAT) activity, on the other hand, decreased in response to As exposure, and it corresponded to the observed decrease in H₂O₂ content. We conclude that As causes a reduction in root elongation by inducing an oxidative stress that is related to enhanced lipid peroxidation, but not to H₂O₂ accumulation.     

Impact of aluminium,fluoride and fluoroaluminate complex on ATPase activity ofNostoc linckia andChlorella vulgaris

This study demonstrates a pH-dependent inhibition of Mg²⁺- and Ca²⁺- ATPase activities ofNostoc linckia andChlorella vulgaris exposed to AlCl₃, AlF₃, NaF and AlCl₃ + NaF together. AlF₃ and the combination of AlCl₃ + NaF were more inhibitory to both the enzymes as compared with AlCl₃ and NaF. Toxicity of the test compounds increased with increasing acidity. Interaction of AlCl₃ + NaF was additive onN. linckia andC. vulgaris, respectively, at pH 7.5 and 6.8, and synergistic at pH 6.0 and 4.5. In the presence of 60 and 100 m PO₄ ^3- an increased NaF concentration (in the AlCl₃ + NaF combination) was required to produce the same degree of inhibition in ATP synthesis and ATPase activity. Toxicity of fluoroaluminate was reduced in the presence of EDTA and citrate. Except for beryllium to some extent, combinations of cadmium, cobalt, iron, manganese, tin and zinc with fluoride were not as effective as aluminium in inhibiting the ATPase activity. The presence of a 100 kDa protein band in SDS-PAGE of both control as well as AlCl₃ + NaF-treated samples suggested that AlF₄ ^– inhibits the ATPase activity by acting as a functional barrier without affecting the structure of the enzyme.     

Flooding-induced membrane damage, lipid oxidation and activated oxygen generation in corn leaves

Flooding effects on membrane permeability, lipid peroxidation and activated oxygen metabolism in corn (Zea mays L.) leaves were investigated to determine if activated oxygens are involved in corn flooding-injury. Potted corn plants were flooded at the 4-leaf stage in a controlled environment. A 7-day flooding treatment resulted in a significant increase in chlorophyll breakdown, lipid peroxidation (malondialdehye content), membrane permeability, and the production of superoxide (O ₂ ^- ) and hydrogen peroxide (H₂O₂) in corn leaves. The effects were much greater in older leaves than in younger ones. Spraying leaves with 8-hydroxyquinoline (an O ₂ ^- scavenger) and sodium benzoate (an .OH scavenger) reduced the oxidative damage and enhanced superoxide dismutase (SOD) activity. A short duration flooding treatment elevated the activities of SOD, catalase, ascorbate peroxidase (AP), and glutathione reductase (GR), while further flooding significantly reduced the enzyme activities but enhanced the concentrations of ascorbic acid and reduced form glutathione (GSH). It was noted that the decline in SOD activity was greater than that in H₂O₂ scavengers (AP and GR). The results suggested that O ₂ ^- induced lipid peroxidation and membrane damage, and that excessive accumulation of O ₂ ^- is due to the reduced activity of SOD under flooding stress.     

Increased Antioxidant Activity under Elevated Temperatures: A Mechanism of Heat Stress Tolerance in Wheat Genotypes

An experiment was conducted with three wheat (Triticum aestivum L.) genotypes C 306, HD 2285 and HD 2329 (differently susceptible to water and temperature stress) to study the extent of oxidative injury and activities of antioxidant enzymes in relation to heat stress induced by manipulating dates of sowing. Increase in temperature by late sowing significantly decreased leaf relative water content (RWC), ascorbic acid content, and increased H₂O₂ content and lipid peroxidation in all the genotypes at 8 and 23 d after anthesis. Temperature tolerant genotypes C 306, closely followed by HD 2285 were superior to HD 2329 in maintaining high RWC, ascorbic acid content, and lower H₂O₂ content and lipid peroxidation (malondialdehyde content) under high temperature (late sowing) at the two stages. Activities of superoxide dismutase and catalase were highest in HD 2285 followed by C 306 and minimum in HD 2329 while ascorbate peroxidase activity was highest in C 306.     

Effect of ultraviolet-B radiation on biomass production,lipid peroxidation,reactive oxygen species,and antioxidants in Withania somnifera

The present study was aimed at understanding the effects of long term supplemental UV-B (3.6 kJ m^?2 d^?1) on biomass production, accumulation of reactive oxygen species, lipid peroxidation, and enzymatic antioxidants in leaves and roots of Withania somnifera (an indigenous medicinal plant). Under the UV-B treatment, a reduction in biomass and an increased malondialdehyde content (a characteristic of lipid peroxidation) were observed in both the shoots and roots. Amongst ROS, H₂O₂ content increased under UV-B in the leaves, whereas it decreased in the roots, and superoxide radical production rate decreased in both the plant parts. The activities of all enzymatic antioxidants tested (ascorbate peroxidase, catalase, glutathione reductase, peroxidase, polyphenol oxidase, and superoxide dismutase) increased under the UV-B treatment, the increase being greater in the roots.     

Aluminum-inhibited root growth of rice seedlings is mediated through putrescine accumulation

The effects of AlCl₃ on growth and polyamine levels of rice roots were investigated. When rice roots were treated with AlCl₃, root growth was markedly inhibited. AlCl₃ treatment resulted in a higher putrescine content and lower spermidine and spermine contents in rice roots. d-Argnine and α-methylornithine, inhibitors of putrescine biosynthesis, caused a reduced content of putrescine in rice roots under Al stress. AlCl₃ treatment also resulted in a decrease in diamine oxidase activity in rice roots. The growth of rice roots in the presence of AlCl₃ was recovered after the addition of d-arginine or α-methylornithine. The protective effect of d-arginine or α-methylornithine in counteracting AlCl₃-inhibited growth of rice roots is unlikely caused by reduction of Al uptake. Furthermore, the effect of the growth recovery in AlCl₃-treated rice roots by d-arginine or α-methylornithine was reversed by the addition of putrescine. Our results strongly suggest that putrescine accumulation is a factor causing growth inhibition of rice roots under Al tress. Evidence is also presented to show that lignification is responsible for putrescine- and AlCl₃-inhibited growth of rice roots.     

Exogenous salicylic acid alleviates NaCl toxicity and increases antioxidative enzyme activity in <Emphasis Type="Italic">Lycopersicon esculentum</Emphasis>

Effects of exogenous salicylic acid (SA) on plant growth, contents of Na, K, Ca and Mg, activities of superoxide dismutase (SOD), guaiacol peroxidase (GPX), ascorbate peroxidase (APX), dehydroascorbate reductase (DHAR), glutathione reductase (GR) and catalase (CAT), and contents of ascorbate and glutathione were investigated in tomato (Lycopersicon esculentum L.) plants treated with 100 mM NaCl. NaCl treatment significantly increased H₂O₂ content and lipid peroxidation indicated by accumulation of thiobarbituric acid reactive substances (TBARS). A foliar spray of 1 mM SA significantly decreased lipid peroxidation caused by NaCl and improved the plant growth. This alleviation of NaCl toxicity by SA was related to decreases in Na contents, increases in K and Mg contents in shoots and roots, and increases in the activities of SOD, CAT, GPX and DHAR and the contents of ascorbate and glutathione.     

Mannose-Induced Modulations in Antioxidants,Protease Activity,Lipid Peroxidation,and Total Phenolics in Etiolated Wheat Leaves

Modulation of different antioxidants, total phenolics, lipid peroxidation, and protease activity as a result of mannose treatment (1%) was studied in leaves of etiolated wheat seedlings. Changes in these biochemicals were monitored up to 96 h after treatment at 24-h intervals. Mannose treatment induced a significant increase in protease activity throughout the scanning period, coupled with a gradual decrease in leaf protein content. Membrane lipid peroxidation (MDA content) was higher at 24 and 72 h after treatment. MDA content remained higher for a longer period due to mannose treatment. During the initial 24 h of mannose treatment, only catalase and total phenolic contents were increased. Catalase activity was down regulated with increasing duration of treatment. On the other hand, peroxidase (POD, APX) activities were initially unaffected but increased with increasing treatment duration. The decreased level of lipid peroxidation at 96 h may be due to detoxification of H₂O₂ by peroxidases. Superoxide dismutase activity was not affected by mannose treatment. In conclusion, evidence is provided that mannose can modulate the expression of the enzymatic antioxidant defense system in wheat leaves.     

Response of Barley Seedlings to UV-B Radiation as Affected by Proline and NaCl

Barley (Hordeum vulgare L. cv. Alfa) seedlings were treated for 4 d before UV-B irradiation with 0.05 mM proline or 150 mM NaCl. UV-B exposure induced synthesis of yellow coloured compounds with maximum absorbance at 438 nm. The content of these compounds was increased in proline-treated and decreased in NaCl-treated plants. UV-B radiation reduced chlorophyll/carotenoids ratio, oxygen evolution rate and photochemical efficiency of PS 2 as estimated by chlorophyll fluorescence and increased proline accumulation, H₂O₂ generation and lipid peroxidation. Exogenous proline had no effect on the parameters studied and did not change the response of plants to UV-B radiation. NaCl inhibited photochemical efficiency of PS 2, reduced oxygen evolution and increased H₂O₂ concentration and lipid peroxidation. The combination of NaCl and proline treatment led to lowering the inhibitory effect of NaCl in non UV-B irradiated seedlings. There was not relationship between the level of UV-B-induced compounds and UV-B tolerance of barley seedlings.     

Antioxidative defense system in pigeonpea roots under waterlogging stress

Pigeonpea [Cajanus cajan (L.) Millsp.] is a waterlogging-sensitive legume crop. We studied the effect of waterlogging stress on hydrogen peroxide (H₂O₂) content, lipid peroxidation and antioxidant enzyme activities in two pigeonpea genotypes viz., ICPL-84023 (waterlogging resistant) and MAL-18 (waterlogging susceptible). In a pot experiment, waterlogging stress was imposed for 6 days at early vegetative stage (20 days after sowing). Waterlogging treatment significantly increased hydrogen peroxide accumulation and lipid peroxidation, which indicated the extent of oxidative injury posed by stress conditions. Enzyme activities of peroxidase (POX), catalase (CAT), ascorbate peroxidase (APX), superoxide dismutase (SOD) and polyphenol oxidase (PPO) increased in pigeonpea roots as a consequence of waterlogged conditions, and all the enzyme activities were significantly higher in waterlogged ICPL-84023 than in MAL-18. POX activity was the maximum immediately after imposing stress, therefore, it was suggested to be involved in early scavenging of H₂O₂, while rest of the enzymes (CAT, APX, SOD and PPO) were more important in late responses to waterlogging. Present study revealed that H₂O₂ content is directly related to lipid peroxidation leading to oxidative damage during waterlogging in pigeonpea. Higher antioxidant potential in ICPL-84023 as evidenced by enhanced POX, CAT, APX, SOD and PPO activities increased capacity for reactive oxygen species (ROS) scavenging and indicated relationship between waterlogging resistance and antioxidant defense system in pigeonpea.     

Evalution of toxicity of abcisic acid and gibberellic acid in rats: 50 days drinking water study

In the present study, the influence of subchronic effects of two plant growth regulators (PGRs) [Abcisic acid (ABA) and Gibberellic acid (GA₃)] on antioxidant defense systems [reduced glutathione (GSH), glutathione reductase (GR), superoxide dismutase (SOD), glutathione-S-transferase (GST) and catalase (CAT)] and lipid peroxidation level (malondialdehyde = MDA) in various tissues of the rat were investigated during treatment as a drinking water model. 75 ppm of ABA and GA₃ in drinking water were continuously administered orally to rats (Sprague-Dawley albino) ad libitum for 50 days. The PGRs treatments caused different effects on the antioxidant defense systems and MDA content of dosed rats compared to controls. The lipid peroxidation end product MDA significantly increased in the lungs, heart and kidney of rats treated with GA₃ without significant change in the spleen. ABA caused also a significant increase in MDA content in the spleen, lungs, heart and kidney. The GSH levels were significantly depleted in the spleen, lungs and stomach of rats treated with ABA without any change in the tissues of rats treated with GA₃ except the kidney where it increased. Antioxidant enzyme activities such as SOD significantly increased in the lungs and stomach and decreased in the spleen and heart tissues of rats treated with GA₃. Meanwhile, SOD significantly decreased in the spleen, heart and kidney and increased in the lungs of rats treated with ABA. While CAT activity significantly decreased in the lungs of rats treated with GA₃, a significant increase occurred in the heart of rats treated with both PGRs. On the other hand, the ancillary enzyme GR activity in the tissues were either significantly depleted or not changed with PGRs treatment. The drug metabolizing enzyme GST activity significantly decreased in the lungs of rats treated with ABA but increased in the stomach of rats treated with both PGRs.

As a conclusion, the rats resisted oxidative stress via the antioxidant mechanism. But the antioxidant mechanism could not prevent the increases in lipid peroxidation in rat's tissues. This data, along with changes, suggests that PGRs produced substantial systemic organ toxicity in the spleen, lungs, stomach, heart and kidney during a 50-day period of subchronic exposure.