Advances in Understanding Cadmium Stress and Breeding of Cadmium-Tolerant Crops

Cadmium(Cd) pollution has emerged as a critical global environmental concern, due to its significant toxicity, environmental persistence, and the pervasiveness of contamination. Significantly, the bioaccumulation of Cd in agricultural crops constitutes a primary vector for its entry into the human diet. This issue warrants urgent attention from both the scientific community and policymakers to develop and implement effective mitigation strategies. This review delves into the physiological impacts of Cd stress on plants, including the suppression of photosynthetic activity, amplification of oxidative stress, and disruptions in mineral nutrient homeostasis. Additionally, the resistance mechanisms deployed by plants in response to Cd stress have been explored, and the prospective contributions of molecular breeding strategies in augmenting crop tolerance to Cd and minimizing its bioaccumulation have been assessed. By integrating and analyzing these findings, we seek to inform future research trajectories and proffer strategic approaches to enhance agricultural sustainability, safeguard human health, and protect environmental integrity.

Effects of Different Kinds of Exogenous Auxin on the Growth of Rice Roots under Cadmium Stress

[Objective] The aim was to study the effect of different kinds of exogenous auxin on the growth of rice roots under cadmium stress.[Method] Oryza sativa L.cv Zhonghua No.11 was used as experimental materials to detect the effect of different kinds of exogenous auxin on the growth of rice roots.[Result] The results showed that 0.1 mmol/L Cd treatment could not only increase primary,adventitious and lateral root length but also lateral root number,whereas the shoot growth was inhibited.When supplemented with different concentrations of NAA,IAA,IBA and 2,4-D,the growth of root system varied and similar change trend had been found.At the auxin concentration of 10^-9-10^-7 mol/L in particular 10^-8 mol/L,all four kinds of auxin promoted the elongation growth of primary and adventitious roots,but inhibition was observed when auxin was higher than 10^-7 mol/L.The decreased shoot growth caused by Cd could not be counteracted by supplementing with the four kinds of auxin.However,at the auxin concentration of 10^-9-10^-8 mol/L,NAA could improve rice growth under Cd stress condition.The formation and development of lateral roots on primary and adventitious roots was not only similar but also different after applying the same concentration of four auxins.[Conclusion] The addition of suitable amount of auxin under cadmium stress （such as 10^-9-10^-8 mol/L of NAA and so on） could ease the damage of cadmium on plants to a certain extent.

Effects of Cadmium Stress on Key Enzymes Involved in Nitrogen Metabolism and Nitrogen,Phosphorus, Potassium Accumulation of Different Varieties of Rice

In this study, the pot experiment was used to explore the differences of activity of key enzymes involved in N metabolism and NPK accumulation under Cd stress during the til iering stage of differen varieties of rice. The results showed that：Cd stress could increase the NPK concentration of different rice type in the til ering stage, while Shen-Liangyou 5867,Yongyou 5550 and Wu-Yunjing 27 showed the highest amplification respectively. Morever, Cd stress can also contribute to the ac-tivity of NR,GS,GOGAT increasing.A s for NR,the Cd stress significantly contribute to NR activity increasing of Huang-Huazhan and Yongyou 538 but is not significant for Wu-Yunjing 27, Shen-Liangyou 5867 and Yongyou 5550, however, the difference among them is not obvious.However, for the activity of GS , Cd stress promote the GS activity. Huang-Huazhan and Wu-Yunjing 27 with low activity in Cd normal level are the most sensitive. Meanwhile the difference between two treatment is the most significant. To the contrary, restrain the GS activity of Shen-Liangyou 5867, Yongy-ou 5550 and the difference is not significant. And under Cd stress, either difference reached significant in GS activity. Cd stress also improve the activity of GOGAT, Wu-Yunjing 27 showed the highest inprovement which showed the lowest GOGAT activity under Cd normal level. Cd stress on rice growth and development of ad-verse, make its lower seed setting rate, 1 000 grain weight decreased, resulting in different degrees of reduction of output of rice.

Effect of Cadmium Stress on the Growth, Antioxidative Enzymes and Lipid Peroxidation in Two Kenaf (Hibiscus cannabinus L.) Plant Seedlings

The effects of cadmium stress on the growth, antioxidative enzymes and lipid peroxidation in two kenaf plants, Fuhong 991 and ZM412, were analysed under control （0.5-strength Hoagland＇s nutrient solution） or five levels of cadmium stress （0.5- strength Hoagland＇s nutrient solution containing different concentrations of Cd2＋）. The leaves and roots of control and cadmium-stressed plants were harvested after 3 wk. At the same Cd concentration, the Cd tolerance index of Fuhong 991 was higher than that of ZM412, indicating that Fuhong 991 may be more tolerant to Cd than ZM412. The superoxide dismutase （SOD）, catalase activity （CAT） and peroxidase （POD） activities fluctuated in the leaves of the Cd-stressed plants compared to the control, whereas the glutathione reductase activity （GR） was much larger than the control for Fuhong 991, ensuring that sufficient quantities of GSH were available to respond to the cadmium stress. In comparison to the control, the dynamic tendency of the SOD, CAT and POD activities in roots of the Cd-stressed plants all increased and then declined, but the POD activity of Fuhong 991 remained nearly unchanged at all of the stress levels. The increase in the enzyme activities demonstrated that Fuhong 991 was more tolerant to cadmium than ZM 412. The lipid peroxidation was enhanced only in the leaves of Cd-stressed ZM 412. These findings indicated that antioxidative activities may play important roles in Cd-stressed Fuhong 991 and ZM 412 and that the leaf and root cell membranes of Fuhong 991 have a greater stability than those of ZM 412. For pollution monitoring purposes, the GR activity in the roots and leaves may serve as a biomarker of Cd for Fuhong 991, whereas lipid peroxidation may serve as biomarker for ZM 412.

Response of Glutathione and Glutathione S-transferase in Rice Seedlings Exposed to Cadmium Stress

A hydroponic culture experiment was done to investigate the effect of Cd stress on glutathione content （GSH） and glutathione S-transferase （GST, EC 2.5.1,18） activity in rice seedlings. The rice growth was severely inhibited when Cd level in the solution was higher than 10 mg/L. In rice shoots, GSH content and GST activity increased with the increasing Cd level, while in roots, GST was obviously inhibited by Cd treatments, Compared with shoots, the rice roots had higher GSH content and GST activity, indicating the ability of Cd detoxification was much higher in roots than in shoots. There was a significant correlation between Cd level and GSH content or GST activity, suggesting that both parameters may be used as biomarkers of Cd stress in rice.

Molecular cloning, sequence analysis, and cadmium stress-rated expression changes of BTG1 in freshwater pearl mussel(Hyriopsis schlegelii)

The B cells translocation gene 1 （BTG1） is a member of the BTG/TOB family of anti-proliferative genes, which have recently emerged as important regulators of cell growth and differentiation among vertebrates. Here, for the first time we cloned the full-length eDNA sequence of Hyriopsis schlegelii （Hs-BTG1）, an economically important freshwater shellfish and potential indicator of environmental heavy metal pollution, for the first time. Using rapid amplification of eDNA ends （RACE） together with splicing the EST sequence from a haemocyte eDNA library, we found that Hs-BTG1 contains a 525 bp open reading frame （ORF） encoding a 174 amino-acid polypeptide, a 306 bp 5＇ untranslated region （5＇ UTR）, and a 571 bp 3＇ UTR with a Poly（A） tail as well as a transcription termination signal （AATAAA）. Homologne searching against GenBank revealed that Hs-BTG1 was closest to Crassostrea gigas BTG1, sharing 50.57% of protein identities. Hs-BTG1 also shares some typical features of the BTG/TOB family, possessing two well-conserved A and B boxes. Clustering analysis of Hs-BTG1 and other known BTGs showed that Hs-BTG1 was also closely related to BTG1 of C. gigas from the invertebrate BTG1 clade. Function prediction via homology modeling showed that both Hs-BTG1 and C. gigas BTG1 share a similar three-dimensional structure with Homo sapiens BTG1. Tissue-specific expression analysis of the Hs-BTG1 via real-time PCR showed that the transcripts were constitutively expressed, with the highest levels in the hepatopancreas and gills, and the lowest in both haemocyte and muscle tissue. Expression levels of Hs-BTG1 in hepatopancreas （2.03-fold）, mantle （2.07-fold）, kidney （2.2-fold） and haemocyte （2.5-fold） were enhanced by cadmium （Cd2＋） stress, suggesting that Hs-BTG 1 may have played a significant role in H, schlegelii adaptation to adverse environmental conditions.

Effect of Biochar on Relieving Cadmium Stress and Reducing Accumulation in Super japonica Rice

It is of great importance to solve the threats induced by cadmium pollution on crops. This paper examined the effect of biochar on cadmium accumulation in japonica rice and revealed the mechanism underlying the response of protective enzyme system to cadmium stress. Biochar derived from rice straw was applied at two application rates under three cadmium concentrations. Shennong 265, super japonica rice variety, was selected as the test crop. The results indicated that cadmium content in above-ground biomass of rice increased with increasing soil cadmium concentrations, but the biochar application could suppress the accumulation of cadmium to some extent. Under high concentrations of cadmium, content of free proline and MDA （malondialdehyde） were high, so did the SOD （superoxide dismutase）, POD （peroxidase） and CAT （catalase） activity in the flag leaf of rice. However, the protective enzyme activities remained at low level when biochar was added.

Effects of Melatonin on Growth,Physiology and Gene Expression in Rice Seedlings Under Cadmium Stress

Melatonin(MLT)is a hormonal substance found in many organisms and can improve plant stress resistance.In this study,the japonica rice variety Y32 and indica rice variety NJ6 were cultivated in hydroponics under different concentrations of CdCl2 at the two-leaf stage.The growth,physiological and biochemical responses of the seedlings and the expression of cadmium(Cd)-related genes under exogenous melatonin(MLT)treatment were assessed.The results indicated that Cd stress destroyed the dynamic balance between reactive oxygen species(ROS)production and removal,resulting in ROS accumulation,membrane lipid peroxidation,and impaired growth and development.Following the application of exogenous MLT to rice seedlings,increases in plant biomass including both underground and above-ground areas were observed.MLT also scavenged the inhibition of superoxide dismutase(SOD)and peroxidase(POD)in a concentration dependent manner in response to Cd stress.Catalase(CAT)activity and malondialdehyde(MDA)expression also decreased following MLT treatment.Amongst the six Cd-related genes assessed,five genes were downregulated and one was up-regulated in response to MLT treatment.Taken together,these data demonstrate that MLT improves the resilience of rice seedlings at the biochemical,physiological,and molecular levels,and diminishes the damage caused by Cd stress.

Effects of Waterlogging Time on Rice Cd Absorption and Accumulation under Cadmium Stress

A pot experiment with exogenous cadmium was utilized to study the effects of waterlogging time on rice yield and Cd accumulationin different growth stages including top tillering stage and filling stage.The results showed that the rice yields of all flooding treatments were lower than the CK(CI).The WI,T1,T2,T3 and T4 decreased significantly by 23.7%,16.0%,15.5%,20.2%and 18.6%respectively.The Cd content of brown rice decreased with the extension of waterlogging time.And WF was the lowest,at only 3.4%of the wet irrigation of the whole growth period(WI).Under the same waterlogging condition,the Cd content in brown rice with 1 to 4 weeks of flooding treatment at the top tillering stage decreased by 27.1%(P﹤0.05),46.6%(P﹤0.05),56.0%(P﹤0.05)and 35.2%(P﹥0.05)respectively,compared with the treatment at the filling stage.And the average decrease was 41.2%.The variation tendency of Cd content in stems and leaves was similar to brown rice.The translocation efficiency of Cd from stems and leaves to rice seeds decreased with the extension of waterlogging time.The Cd enrichment factor of stems and leaves,as well as brown rice,varied greatly with different treatments.Specifically,the Cd enrichment factors in brown rice and in stems and leaves under WI were 28.0 and 17.8 times higher respectively than those under WF.The findings of this study demonstrated that flooding could inhibit the uptake and accumulation of Cd in rice,with significant positive correlation between them.The inhibition effect of flooding treatment on Cd accumulation in rice at the top tillering stage was superior to that at the filling stage.

Purification and Identification of Glutathione S-transferase in Rice Root under Cadmium Stress

Cadmium （Cd） contamination in paddy soils poses a serious threat to the production and quality of rice. Among various biochemical processes related to Cd detoxification in rice, glutathione S-transferase （GST） plays an important role, catalyzing Cd complexation with glutathione （GSH） and scavenging reactive oxygen species （ROS） in cells. In this study, a hydroponic experiment was conducted to investigate the response of GST isozymes in rice roots upon Cd exposure. Results showed that the GST activity in rice roots was clearly enhanced by 50 pmol/L Cd treatment for 7 d. The GST isozymes were purified by ammonium sulphate precipitation, gel filtration chromatography and affinity chromatography. After being separated by SDS-PAGE and visualized by silver staining, GSTU6 was identified by in-gel digestion, MALDI-TOF-MS analysis and peptide mass fingerprint. The results confirm the vital function of tau class rice GST in Cd detoxification.

Effects of Cadmium Stress on the Growth of Rice Seedlings

The seedlings of 6 rice varieties were treated with Cd concentrations of 25,50 and 100μmol/L for 5 d.The results showed that the longest root length,seedling height,fresh weight per plant and dry weight per plant of 6 rice varieties were significantly inhibited.The inhibiting effects showed a trend as the following:the longest root length>fresh weight per plant>dry weight per plant>seedling height,and the inhibiting effect increased with the increase of Cd stress concentration.It is more economical and reasonable to use 50μmol/L Cd stress concentration to identify Cd tolerance of rice seedlings.And it is more scienti fic to evaluate the Cd tolerance of rice seedling comprehensively by using the 4 indexes of longest root length,fresh weight per plant,dry weight per plant and seedling height.

Transcriptome Sequencing and Analysis of Agaricus blazei Mycelia Under Cadmium Stress

To explore the mechanism of Agaricus blazei Murrill enrichment of the heavy metal cadmium,we employed Illumina high-throughput sequencing to analyze the transcriptomes of A.blazei mycelia treated with and without exogenous cadmium addition,and then the differentially expressed genes(DEGs)of the strains with high and low cadmium enrichment between the control and cadmium treatment were screened out.The results showed that the DEGs were mainly involved in steroid biosynthesis,antibiotic biosynthesis,protein processing in endoplasmic reticulum,glutathione metabolism and other pathways.Carbon metabolism and glutathione metabolism may play an important role in the response of A.blazei mycelium to cadmium stress.

Alterations of Antioxidative Enzymes Activities and Induction of Lipid Peroxidation in Germinating Wheat Seeds Subjected to Cadmium Stress

Germinating wheat （Triticum aestivum L.） seeds were exposed to CdCI2 （50, 100 and 200 μM） for 48 h and some aspects of oxidative metabolism was assessed in the embryonic tissues. The germination percentage and the soluble protein content of the embryonic tissues were found to decrease with increasing of Cd concentration. There was elevation in superoxide dismutase （SOD） and decline in catalase （CAT） and peroxidase （POX） activities. The increasing of lipid peroxidation levels indicated the prevalence of oxidative stress in the tissues which was probably due to the alteration of antioxidative enzymes activities. The adding of ascorbate, along with CdCl2, has resulted in restoration of the Cd induced decline in CAT activity. Weakening in H2O2 detoxification system seems to be the principal reason behind Cd induced oxidative stress in germinating seeds. Thus, imposition of oxidative stress might be the consequence of cadmium stress and this finding may help in elucidating the mechanisms underlying cadmium mediated toxicity in germinating seeds.

Physiological Response of Hydrilla verticillata （I.f.） Royle Exposed to Cadmium Stress

Hydrilla verticillata （I.f.） Royle twigs were exposed to CdCI2 （50, 100, 200 and 500 BM） under continuous light for 48 hrs and the physiological parameters like photosynthetic pigment （chlorophylls a, b and carotenoids） contents, activities of antioxidative enzymes like superoxide dismutase （SOD）, catalase （CAT） and peroxidase （POX） along with lipid peroxidation level were determined. With respect to increase in Cd concentration in the medium and exposure duration, decrease in pigment contents, and decrease in the activities of SOD, CAT and POX were found. The increased levels of lipid peroxidation indicated the prevalence of oxidative stress situation in the tissues which might be one of the reasons behind Cd induced toxicity in Hydrilla verticillata. Since there was significant decrease in the activities of key antioxidative ezymes, the study suggests that Hydrilla verticillata may not be effective for phytoremediation of cadmium in these concentration ranges.

Proteomic Study for Responses to Cadmium Stress in Rice Seedlings

A proteomic approach including two-dimensional electrophoresis and mass spectrometric （MALDI-TOF MS） analyses was used to investigate the responses to cadmium （Cd） stress in seedlings of rice （Oryza sativa L.） varieties Shanyou 63 and Aizaizhan. Cd stress significantly inhibited root and shoot growth, and affected the global proteome in rice roots and leaves, which induced or upregulated the expression of corresponding proteins in rice roots and leaves when rice seedlings were exposed to 0.1 or 1.0 mmol/L Cd. The Cd-induced proteins are involved in chelation and compartmentation of Cd, elimination of active oxygen free radicals, detoxification of toxic substances, degradation of denatured proteins or inactivated enzymes, regulation of physiologic metabolism and induction of pathogenesis-related proteins. Comparing the Cd-induced proteins between the two vadeties, the β-glucosidase and pathogenesis-related protein family 10 proteins were more drastically induced by Cd stress in roots and leaves of Aizaizhan, and the UDP-glucose protein transglucosylase and translational elongation factor Tu were induced by 0.1 mmol/L Cd stress in roots of Shanyou 63. This may be one of the important mechanisms for higher tolerance to Cd stress in Shanyou 63 than in Aizaizhan.

Dynamic changes of root proteome reveal diverse responsive proteins in maize subjected to cadmium stress

Toxic symptoms and tolerance mechanisms of heavy metal in maize are well documented. However, limited information is available regarding the changes in the proteome of maize seedling roots in response to cadmium(Cd) stress. Here, we employed an i TRAQ-based quantitative proteomic approach to characterize the dynamic alterations in the root proteome during early developmental in maize seedling. We conducted our proteomic experiments in three-day seedling subjected to Cd stress, using roots in four time points. We identified a total of 733, 307, 499, and 576 differentially abundant proteins after 12, 24, 48, or 72 h of treatment, respectively. These proteins displayed different functions, such as ribosomal synthesis, reactive oxygen species homeostasis, cell wall organization, cellular metabolism, and carbohydrate and energy metabolism. Of the 166 and 177 proteins with higher and lower abundance identified in at least two time points, 14 were common for three time points. We selected nine proteins to verify their expression using quantitative real-time PCR. Proteins involved in the ribosome pathway were especially responsive to Cd stress. Functional characterization of the proteins and the pathways identified in this study could help our understanding of the complicated molecular mechanism involved in Cd stress responses and create a list of candidate gene responsible for Cd tolerance in maize seeding roots.

Hydrogen Peroxide-Mediated Growth of the Root System Occurs via Auxin Signaling Modification and Variations in the Expression of Cell-Cycle Genes in Rice Seedlings Exposed to Cadmium Stress

The link between root growth, H2O2, auxin signaling, and the ceil cycle in cadmium （Cd）-stressed rice （Oryza sativa L. cv. Zhonghua No. 11） was analyzed in this study. Exposure to Cd induced a significant accumulation of Cd, but caused a decrease in zinc （Zn） content which resulted from the decreased expression of OsHMA9 and OsZIP. Analysis using a Cd-specific probe showed that Cd was mainly localized in the meristematic zone and vascular tissues. Formation and elongation of the root system were significantly promoted by 3-amino-l,2,4-triazole （AT）, but were markedly inhibited by N,N＇. dimethylthiourea （DMTU） under Cd stress. The effect of H2O2 on Cd-stressed root growth was further confirmed by examining a gain-of-function rice mutant （carrying catalasel and glutathione-S-transferase） in the presence or absence of diphenylene iodonium. DR5-GUS staining revealed close associations between H2O2 and the concentration and distribution of auxin. H2O2 affected the expression of key genes, including OsYUCCA, OsPIN, OsARF, and OslAA, in the auxin signaling pathway in Cd-treated plants. These results suggest that H2O2 functions upstream of the auxin signaling pathway. Furthermore, H2O2 modified the expression of cell-cycle genes in Cd-treated roots. The effects of H2O2 on root system growth are therefore linked to auxin signal modification and to variations in the expression of cell-cycle genes in Cd-stressed rice. A working model for the effects of H2O2 on Cd-stressed root system growth is thus proposed and discussed in this paper.

bHLH104 confers tolerance to cadmium stress in Arabidopsis thaliana

Cd is a non-essential heavy metal that is toxic to both plants and animals. Here, we reveal that the transcription factor bHLH104 positively regulates Cd tolerance in Arabidopsis thaliana. We show that Fe deficiency-responsive genes were induced by Cd treat- ment, and that their upregulation was suppressed in bhlh104 loss-of-function mutants, but enhanced upon overexpression of bHLH104. Correspondingly, the bhlh104 mutants displayed sensitivity to Cd stress, whereas plants overexpressing bHLH104 exhibited enhanced Cd tolerance. Further analysis suggested that bHLH104 positivelyregulates four heavy metal detoxification-associated genes, IREG2, A4TP3, HA4A3 and NAS4, which play roles in Cd sequestration and tolerance. The bHLH104 overexpres- sion plants accumulated high levels of Cd in the root but low levels of Cd in the shoot, which might contribute to the Cd tolerance in those lines. The present study thus points to bHLH104 as a potentially useful tool for genetic engineering of plants with enhanced Cd tolerance.

An Arabidopsis mutant atcsr-2 exhibits high cadmium stress sensitivity involved in the restriction of H_2S emission

The gene AtCSR encodes peptidyl-prolyl cis/trans isomerases (PPIases) that accelerate energetically unfavorable cis/trans isomerization of the peptide bond preceding proline production.In our studies,we found that AtCSR was associated with cadmium (Cd)-sensitive response in Arabidopsis.Our results show that AtCSR expression was triggered by Cd-stress in wild type Arabidopsis.The expression of some genes responsible for Cd2+ transportation into vacuoles was induced,and the expression of the iron-regulated transporter 1 (IRT1) related to Cd2+ absorption from the environment was not induced in wild type with Cd2+ treatment.The expression of Cd-transportation related genes was not in response to Cd-stress,whereas IRT expression increased dramatically in atcsr-2 with Cd2+ treatment.The expression of glutathione 1 (GSH1) was consistent with GSH being much lower in atcsr-2 in comparison with the wild type with Cd2+ treatment.Additionally,malondialdehyde (MDA),hydrogen peroxide,and Cd2+ contents,and activities of some antioxidative enzymes,differed between the wild type and atcsr-2.Hydrogen sulfide (H2S) has been confirmed as the third gas-transmitter over recent years.The findings revealed that the expression pattern of H2 S-releasing related genes and that of Cd-induced chelation and transportation genes matched well in the wild type and atcsr-2,and H2S could regulate the expression of the Cd-induced genes and alleviate Cd-triggered toxicity.Finally,one possible suggestion was given:down-regulation of atcsr-2,depending on H2S gas-transmitter not only weakened Cd2+ chelation,but also reduced Cd2+ transportation into vacuoles,as well as enhancing the Cd2+ assimilation,thus rendering atcsr-2 mutant sensitive to Cd-stress.

A novel aldo-keto reductase gene, IbAKR, from sweet potato confers higher tolerance to cadmium stress in tobacco

High concentrations of Cd can inhibit growth and reduce the activity of the photosynthetic apparatus in plants. In several plant species, aldo-keto reductases(AKRs) have been shown to enhance tolerance to various abiotic stresses by scavenging cytotoxic aldehydes;however, few AKRs have been reported to enhance Cd stress tolerance. In this study, the gene Ib AKR was isolated from sweet potato. The relative expression levels of Ib AKR increased significantly(approximately 3-fold) after exposure to 200 mmol$L–1 Cd Cl2 or 10 mmol$L–1 H2 O2. A subcellular localization assay showed that Ib AKR is predominantly located in the nucleus and cytoplasm.Ib AKR-overexpressing tobacco plants showed higher tolerance to Cd stress than wild-type(WT). Transgenic lines showed a significant ability to scavenge malondialdehyde(MDA) and methylglyoxal(MG). In addition,proline content and superoxide dismutase activity were significantly higher and H2 O2 levels were significantly lower in the transgenic plants than in the WT. Quantitative real-time PCR analysis showed that the reactive oxygen species(ROS) scavenging genes encoding guaiacol peroxidase(GPX), ascorbate peroxidase(APX), monodehydroascorbate reductase(MDHAR) and peroxidase(POD) were significantly upregulated in transgenic plants compared to WT under Cd stress. These findings suggest that overexpressing Ib AKR enhances tolerance to Cd stress via the scavenging of cytotoxic aldehydes and the activation of the ROS scavenging system.