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Hindawi Publishing CorporationThe Scientific World JournalVolume 2013 Article ID 714639 9 pageshttpdxdoiorg1011552013714639

Review ArticlePeroxidase(s) in Environment Protection

Neelam Bansal and Shamsher S Kanwar

Department of Biotechnology Himachal Pradesh University Shimla 171 005 India

Correspondence should be addressed to Shamsher S Kanwar kanwarss2000yahoocom

Received 30 August 2013 Accepted 5 October 2013

Academic Editors S Menoret and P Poltronieri

Copyright copy 2013 N Bansal and S S Kanwar This is an open access article distributed under the Creative Commons AttributionLicense which permits unrestricted use distribution and reproduction in any medium provided the original work is properlycited

Industrial discharges of untreated effluents into water bodies and emissions into air have deteriorated the quality of water andair respectively The huge amount of pollutants derived from industrial activities represents a threat for the environment andecologic equilibrium Phenols and halogenated phenols polycyclic aromatic hydrocarbons (PAH) endocrine disruptive chemicals(EDC) pesticides dioxins polychlorinated biphenyls (PCB) industrial dyes and other xenobiotics are among the most importantpollutants Peroxidases are enzymes that are able to transform a variety of compounds following a free radical mechanism therebyyielding oxidized or polymerized productsTheperoxidase transformation of these pollutants is accompanied by a reduction in theirtoxicity due to loss of biological activity reduction in the bioavailability or the removal from aqueous phase especially when thepollutant is found in waterThe review describes the sources of peroxidases the reactions catalyzed by them and their applicationsin the management of pollutants in the environment

1 Introduction

Two unpredicted challenges for human being are energy andenvironment Functioning of society as a whole and itsfuture progress are dependent on the availability of new andrenewable sources of energy and on the capacity to changepolluting productive processes for new environment friendlyprocesses Together these developments have led to a grow-ing awareness of the central importance for the environmen-tal sciences as humankind attempts to transition to a moresustainable relationship with the Earth and its naturalresources [1] Peroxidases have potential to decrease environ-mental pollution by bioremediation ofwastewater containingphenols cresols and chlorinated phenols for biopulpingand decolourization of synthetic textile azo-dyes Peroxidases(EC 11117) are oxidoreductases that catalyse the reductionof peroxides such as hydrogen peroxide (H

2O2) and the

oxidation of a variety of organic and inorganic compounds[2 3] Specifically peroxidase activity involves donating elec-trons that bind to other substrates such as ferricyanides andascorbate in order to break them into harmless components

Peroxidases have potential for bioremediation of wastewater contaminated with phenols cresols and chlorinatedphenols for biopulping [4] biobleaching in paper industry

textile-dye degradation and removal of peroxide frommate-rials such as food stuffs and industrial wastes Process waterfrom textile mills often features a strong coloration due topresence of rhodamine dyes which are resistant to conven-tional bleaching treatment and can be degraded by perox-idase [5] The unique ability of white rot fungi to degradelignin is largely attributable to the nonspecific free radicalmediated oxidizing reactions carried out by their extracellu-lar peroxidases [6] Peroxidase oxidizes dimethoxybenzenelignin dimers phenols amines dyes and aromatic alcoholsin the absence of Mn(II) peroxidase oxidizes phenolic andnonphenolic substrates Yet another peroxidase designateddye-decolorizing peroxidase from Agaricus type fungi wasreported to catalyze the oxidation of dyes and phenolic com-pounds [7] Peroxidases from different sources are relativelynonspecific and provide white rot fungi the unique abilityto degrade abroad array of environmental pollutants such asdioxins polychlorinated biphenyls petroleumhydrocarbonsmunitions wastes (such as trinitrotoluene) industrial dyeeffluents herbicides and pesticides [8]

2 Sources of Peroxidase

Peroxidases (EC 11117) are widely distributed in natureThese enzymes are produced by a variety of sources including

2 The Scientific World Journal

Substratereduced + H2O2

HRP(EC 11117)

Substrateoxidized + 2H2O

Figure 1 A general reaction catalyzed by HRP

plants animals and microbes Peroxidases produced frommicrobial sources such as bacteria (Bacillus sphaericus Bacil-lus subtilis Pseudomonas sp Citrobacter sp) Cyanobacteria(Anabaena sp) fungi (Candida krusei Coprinopsis cinereaPhanerochaete chrysosporium) actinomycetes (StreptomycesspThermobifida fusca) and yeast are used in decompositionof pollutants production of animal feedstock and raw mate-rials for the chemical agricultural paper industries textiledye degradation paper-pulp industry for lignin degradationdye decolorization sewage treatment and also as biosen-sors Many plant sources for peroxidases production havebeen reported such as horseradish papaya (Carica papaya)banana (Musa paradisiacal) and bare (Acorus calamus)Peroxidase obtained from horseradish (HRP) is extensivelyused in diagnostic kits in ELISA for labeling an antibodysynthesis of various aromatic chemicals and removal ofperoxides from materials such as foodstuff and industrialwastes (Figure 1)

3 Characteristics of Peroxidase(s)

Peroxidases are oxidoreductases that catalyse a variety ofreactions such as reduction of peroxides such as hydrogenperoxide and oxidation of a variety of organic and inorganiccompounds These are heme proteins and contain iron (III)protoporphyrin IX as the prosthetic group They have amolecular weight ranging from 30 to 150 kDa The termperoxidase represents a group of specific enzymes such asNADH peroxidase (EC 11111) glutathione peroxidase (EC11119) and iodine peroxidase (EC 11118) as well as a varietyof nonspecific enzymes that are simply known as peroxidases

4 Applications and Peroxidase Biocatalysis inManagement of Environmental Pollutants

41 Decolorization of Synthetic Dyes Dye wastes representone of the most problematic groups of pollutants consideredas xenobiotics that are not easily biodegradable [9] Thesedyes are mostly used in textile dyeing paper printing colourphotography and as additive in petroleum products Whenthese synthetic dyes are discharged into industrial effluentsthey cause environmental pollution Textile industries playa vital role in the economic increases in India Water isone of the major products of nature used enormously byhuman beings and it is not unnatural that any growingcommunity generates enormous waste water or sewage [10]To achieve the biodegradation of environmentally hazardouscompounds white rot fungi appear as a valuable alternative

The capability of oxidation is based on the ability of white rotfungi to produce oxidative enzymes such as laccase mang-anese peroxidase and lignin peroxidase [11] These oxidasesand peroxidases have been reported as excellent oxidantagents to degrade dyes [12]

Several bacterial peroxidases have been used for decol-orization of synthetic textile dyes Removal of chromate Cr(VI) and azo dye Acid Orange 7 (AO7) using Brevibacteriumcasei under nutrient-limiting conditions has been studiedAO7 was used as an electron donor by the reduction enzymeof Brevibacterium casei for the reduction of Cr (VI) Thereduced chromate Cr (III) complexed with the oxidizedAO7 formed a purple intermediate [13] Decolorization ofdifferent azo dyes by Phanerochaete chrysosporium RP 78under optimized conditions was studied [14] by reactionmechanism via azo dye Peroxidase was produced underaerobic conditions as a secondarymetabolite in the stationaryphase Bacillus sp VUS isolated from textile effluent con-taminated soil showed capability for degrading a variety ofdyes [15] The production of ligninolytic peroxidases directlyoxidizing aromatic compounds has been described in fungi[16] Other peroxidases were detected in microorganismsresponsible for the biodegradation of industrial dyes togetherwith lignin peroxidase [17] An edible macroscopic fungiPleurotus ostreatus produced an extracellular peroxidase thatcan decolorize remazol brilliant blue and other structurallydifferent groups including triarylmethane heterocyclic azoand polymeric dyes Bromophenol blue was decolorized best(98) while methylene blue and toluidine blue O were leastdecolorized 10 [18] HRP was found to degrade industriallyimportant azo dyes such as remazol blue This dye containsat least one aromatic group in its structure making it apossible substrate of HRP [19] The dyeing and bleachingunitrsquos contaminants seeping into the ground have pollutedthe ground water rendering it unsuitable for consumption(Table 1)

42 Bioremediation of Waste Water Removal of PhenolicContaminants and Related Compounds Industrial pollutionhas been a major factor causing the degradation of the envi-ronment around us affecting the water we use its quality andhuman health is directly related issues Improved quality andincreased quantity of water would bring forth health benefitsSafe water eliminates the infective agents associated withwater borne diseases availability of greater quantity of watercan improve health by allowing improved personal hygieneWater pollution caused industrial waste products to releaseinto lakes rivers and other water bodies that make marinelife no longer hospitable Peroxidases have been applied to

The Scientific World Journal 3

Table 1 Decolorization and detoxification of synthetic textile dyes and other industry effluent by microbial peroxidase(s)

SNo Type of peroxidase Type of

microorganism Microorganism Application Reference

(1) Peroxidase Bacteria E coli Dye degradation [20](2) Peroxidase Bacteria Bacillus sp F31 Dye degradation [21]

(3)

Manganese-dependentperoxidase (MnP) lignin

peroxidase (LiP)Fungi

Fourbasidiomycetous fungi(Pleurotus ostreatussensu CookeCoriolus versicolor (L) Quel

Tyromyces albidus (Schaeff) Donkand Trametes gallica

Biodelignification [22]

(4) Lignin peroxidase Bacteria Citrobacterfreundii (FJ581026) andCitrobacter sp (FJ581023)

Black liquor (pulping byproduct cause serious

environmental problem)[4]

(5) Lignin peroxidase Yeast Candida krusei

Basic Violet 3 (BV)extensively used in humanand veterinary medicine asa biological stain and in

various commercial textileprocesses

[23]

(6) Lignin peroxidase Bacterium Pseudomonas desmolyticum Diazo dye Direct Blue-6 [24]

(7) Mn-peroxidase Bacterium Pseudomonas sp

Malachite green awidely-used recalcitrantdye has been confirmed to

be carcinogenic andmutagenic against many

organisms

[25]

(8) Lignin peroxidase White rotfungi Pleurotusostreatus Remazol Brilliant Blue R

(Artificial dye) [26]

(9) Peroxidase Bacterium Pseudomonas sp Congo red decolorization [27]

(10)

Lignin peroxidaseisoenzymes (LiP 465 LiP

415 and LiP 385)Fungus Phanerochaete chrysosporium

Azo triphenyl methaneheterocyclic andpolymeric dyes

[28]

(11) Peroxidase bacterium Clostridium bifermentans Reactive azo dyes [29]

(12) Versatile peroxidase Fungus Thanatephorus cucumeris Anthraquinone dyeReactiveblue 5 [30]

(13) DyP-type peroxidases Fungi Auricularia auricula-judae High-redox potential dyes [31]

(14)Extracellular

LiP Bacteria Bacillus sp Navy blue 2GL-azo dye [15]

(15)Dye-decolorizingperoxidases (DyP) Fungi Pleurotusostreatus Azo dyes [32]

the bioremediation of waste waters contaminated with phe-nols cresols and chlorinated phenols [2 9] Aromatic com-pounds including phenols and aromatic amines constituteone of the major classes of pollutants They are found inthe waste waters of a wide variety of industries includingcoal conversion petroleum refining resins and plasticswood preservation metal coating dyes and other chemicalstextiles mining and dressing and pulp and paper industries[34] Phenols and halogenated phenols present in textileindustries processed water are known to be toxic and alsosome of them are hazardous carcinogens that can accumulatein the food chain [5]

Peroxidases comprise an important class of enzymesable to catalyze the oxidative coupling reactions of a broadrange of phenolic compounds [35] Lignin peroxidasefrom Phanerochaete chrysosporium HRP myeloperoxidase

lactoperoxidase microperoxidase-8 a versatile peroxidasefrom Bjerkandera adusta and chloroperoxidase fromCaldar-iomyces fumago [36] were able to transform pentachlorophe-nol totetrachloro-14-benzoquinone by an oxidative dehalo-genation in the presence of H

2O2 An extracellular man-

ganese peroxidase produced by P chrysosporium P sordidaC subvermispora P radiata D squalens and P rivulosuTwo electron oxidation of that extracellular peroxidase byH2O2yields Compound I which undergoes two consecutive

one-electron reduction steps by oxidizing Mn2+ into Mn3+that in turn oxidizes phenolic compounds [37] Many toxicaromatic and aliphatic compounds occur in waste water ofa number of industries Among these phenol is the mostcommon aromatic pollutant and is also found in contami-nated drinking water Phenol can be toxic when present at anelevated level and is known to be carcinogenic It has an effect


OH

OH O

O

BQ

QRMycelium

Ominus

Fe+3

Fe+3

Fe+2Fe+2

BQminus

H2O2

BQH2

O2

O2

minuslowast

OHlowast

OlowastPero

xidase

Figure 2 Reaction scheme involved in the production of hydroxylradical by white rot fungi via quinone redox cycling [33] 14-benzoquinone (BQ) is reduced by quinone reductase (QR) pro-ducing hydroquinone (BQH

2) which is oxidized by any of the

lignin modifying enzymes to semiquinones (BQminus) The productionof superoxide anion radicals (O

2

minus) by BQminus autoxidation is mainlycatalyzed by Fe3+ that is reduced to Fe2+ Fentonrsquos reagent formationis accomplished by O

2-dismutation to H

2O2

on health even at low concentration A laboratory phenolwas treated with turnip root enzyme (peroxidase) extract inthe presence of H

2O2as an oxidant to form corresponding

free radicals Free radicals polymerize to form substances thatare less soluble in water The precipitates were removed bycentrifugation and residual phenol was estimated [38] Theresults showed that turnip root enzyme extract degraded phe-nol more efficiently Another versatile peroxidase producedby P eryngii and P ostreatus oxidizedMn2+ intoMn3+ similarto the action of MnP and also high redox potential aromaticcompounds as LiP does had broad specificity and oxidizednonphenolic compounds [39]

421 Mechanism of HRP-H2O2-Phenol Reaction Horsera-

dish peroxidase undergoes a cyclic reaction when reactingwith phenolic substratesThis sequence is summarized in thefollowing reactions

E +H2O2997888rarr Ei +H

2O (1)

Ei + PhOH1015840 997888rarr Eii + PhO (2)

Eii + PhOH10158401015840 997888rarr E + PhO +H2O (3)

The enzyme starts in its native form (E) and is oxidizedby H2O2to form an active intermediate compound known

as compound 1 (Ei) Compound 1 oxidizes one molecule ofphenol (PhOH) to form a phenol free radical (PhO) andbecome compound II (Eii) Compound II oxidizes a secondphenol molecule to produce another phenol free radical andcomplete the cycle by returning to its native form E The freeradicals polymerize and form insoluble compounds which

precipitate from solution [40] The polymerization reactionis illustrated in

PhO + PhO 997888rarr Polymer of aromatic compounds (4)

Yet another peroxidase designated dye-decolorizing peroxi-dase (EC 1 1 1 119909) from Agaricus type fungi was reported tocatalyze the oxidation of dyes and phenolic compounds [7](Figure 2)

43 Removal of Endocrine Disruptive Chemicals (EDCs) Sev-eral classes of oxidative enzymes have shown promise forefficient removal of EDCs that are resistant to conventionalwaste water treatments Although the kinetics of reactionsbetween individual EDCs and selected oxidative enzymessuch as HRP are well documented in the literature there hasbeen little investigation of reactions with EDC mixtures [41]EDCs are a group of compounds that due to their chemicalstructure are able to act as agonists or antagonists of hor-mones They can disturb the synthesis secretion transportbinding action and elimination of the endogenous hor-mones which are responsible for maintaining homeostasisreproduction development and integrity in living organismsand their progeny [42] They are widely dispersed in theenvironment but are mainly found in waste water effluentsSeveral works reported the EDC oxidation by manganeseperoxidase Using 10UmL of manganese peroxidase fromPleurotus ostreatus 04mM bisphenol was eliminated in 1 h[43] Peroxidases are also helpful in removal or degradation ofother potent environmental pollutants such as chloroanilinesand polycyclic aromatic hydrocarbons [44]

44 Degradation of Polychlorinated Biphenyls (PAHs) Pesti-cides Pesticides include a broad range of substances mostcommonly used to control insects weeds and fungi Pesticideexposure in human is associated with chronic health prob-lems or health symptoms such as respiratory problemsmem-ory disorders dermatologic conditions cancer depressionneurologic deficits miscarriages and birth defects [45] Bio-logical decomposition of pesticides is themost important andeffective way to remove these compounds from the envi-ronment Microorganisms have the ability to interact bothchemically and physically with substances leading to struc-tural changes or complete degradation of the target molecule[46]

Peroxidases extracted from some fungal species havegreat potential to transform several pesticides into harmlessform(s) Transformation of organophosphorus pesticides bywhite rot fungi has been studied [47] and transformation ofseveral organophosphorus pesticides by the chloroperoxidasefrom Caldariomyces fumago has been reported PAHs arecomposed of two or more fused aromatic rings and arecomponents of crude oil creosote and coal [48] Most of thecontamination by PAHs had originated from the extensiveuse of fossil fuels as energy sources Peroxidases and phenoloxidases can act on specific PAHrsquos by transforming them toless toxic or products easier to degrade PAHs are oxidizedby peroxidases such as lignin peroxidase [49] andmanganeseperoxidase [50] In spite of their versatility and potential use


in environmental processes peroxidases are not applied atlarge scale yet Diverse challenges such as stability redoxpotential and the production of large amounts should beaddressed in order to apply peroxidases in the pollutanttransformation [51] Peroxidases extracted from some fungalspecies have great potential to transform several pesticidesinto harmless form(s) In spite of their versatility and poten-tial use in environmental processes peroxidases are notapplied at large scale yet Diverse challenges such as stabilityredox potential and the production of large amounts shouldbe addressed in order to apply peroxidases in the pollutanttransformation

45 Degradation of Chlorinated Alkanes and Alkenes Con-tamination of soils and aquifers by the aliphatic halocar-bons trichloroethylene (TCE) and perchloroethylene (PCE)widely used as degreasing solvents is a serious environmentalpollution problem TCE is subject to in vitro reductivedehalogenation catalyzed by LiP of P chrysosporium in thepresence of tertiary alcohol H

2O2 and EDTA (or oxalate)

leading to the production of the corresponding reducedchlorinated radicals [52] One strain of bacterium IM-4capable of degrading imazethapyr (IMZT) was isolated fromthe IMZT-contaminated soil This strain also showed thecapability to degrade other imidazolinone herbicides suchas imazapyr imazapic and imazamox [53] Extracellularhydroxyl radicals produced byT versicolor via quinone redoxcycling were also shown to catalyze degradation of PCE andTCE [1] TCE is mineralized by P chrysosporium culturesgrown aerobically These investigators proposed that TCEis subject to in vitro reductive dehalogenation catalyzed byLiP of P chrysosporium in the presence of tertiary alcoholH2O2 and EDTA (or oxalate) leading to the production of

the corresponding reduced chlorinated radicals [8]

46 Degradation of Phenoxy Alkanoic and TriazineherbicidesThe most commonly used broad leaf herbicides aroundthe world are 24-dichlorophenoxyacetic acid (24-D) and245-trichlorophenoxyaceticacid (245-T) 24-D and per-haps 245-T are a component of Agent Orange that waswidely used as a defoliant 24-D is quite susceptible tobacterial degradation and generally does not persist for longin the environment On other hand 245-T is relatively moreresistant to microbial degradation and tends to persist in theenvironment It has been blamed for serious illnesses inmanyveterans of Vietnam War where they got exposed to AgentOrange that was used as a defoliantThese were also reportedto be mutagenic agents and thus very toxic to humansLigninolytic peroxidases of P chrysosporium and Dichomitusqualens were involved in the degradation of chlorinated phe-nolic intermediates of 24-D and 245-T These results werebased on the increased degradation of ring-labeled and sidechain-labeled 245-T and 24-D by D Squalens on additionof Mn2+ (a known inducer of MnP) to the medium and onincreased degradation by P chrysosporium in nitrogen-limitedmedium (inwhich production of both LiP andMnP isinduced) Atrazine is a commonly used triazine herbicide andis degraded by a number of white rot fungi produced laccasesand peroxidase [54]

47 Degradation of Chlorinated Dioxins Polychlorinateddibenzodioxins (PCDDs) are a group of highly toxic envi-ronmental pollutants that are confirmed human carcinogensand tend to bioaccumulate in humans and animals due totheir lipophilic properties Polychlorinated dibenzodioxins(PCDD) and polychlorinated dibenzofurans (PCDF) havebeen shown to be degraded by several species of white rotfungi [55] suggesting the possible involvement of LiP andMnP A fungus P sordida produced MnP but no LiP andcrude MnP showed degradation of the dioxins

48 Degradation of Chlorinated Insecticides Lindane (c iso-mer of hexachlorocyclohexane) was a widely used pesticidein the past and an estimated 600000 tons of lindane wereproduced globally between the year 1950 and 2000 Thereis a global ban on the use of lindane now because of itsenvironmental persistence as a pollutant P chrysosporiumcultured under ligninolytic conditions was reported to par-tially mineralize lindane in liquid cultures and in a corncob-amended soils inoculated with P chrysosporium [56] but lin-dane degradationwas not observed in vitro using purified LiPand MnP from P chrysosporium [57] DDT (111-trichloro-22-bis [4-chlorophenyl] ethane) the first of the chlorinatedorganic insecticides was used quite heavily after World WarII High levels of DDT found in agricultural soils are of deepconcern because they present serious threats to food securityand human health The white rot fungi P chrysosporium Postreatus T versicolor and Phellinus weirii have been shownto mineralize DDT [58]

49 Peroxidase as Biosensors Biosensors have been definedas analytical devices which tightly combine biorecognitionelements with physical transducers for detection of the targetcompound Several examples of biosensors developed for rel-evant environmental pollutants Biosensors can be useful forexample for the continuous monitoring of a contaminatedarea [59] They may also present advantageous analytical fea-tures such as high specificity and sensitivity (inherent in theparticular biological recognition bioassay H

2O2is consid-

ered the mediator of the biochemistry of cellular pathologyand maybe involved in the etiology of aging and progressiveneurodegenerative diseases such as Parkinsonrsquos disease Dueto its crucial role in neurochemistry determination of theconcentration of H

2O2has been a considerable interesting

research field Electrochemical methods based on peroxidasebiosensors have proved to be significantly advantageous tothe biosciences due to their direct real-time measurementsand capability for practical applications [60] A novel thirdgeneration biosensor for hydrogen peroxide was constructedby cross-linking HRP onto an electrode modified with multi-wall carbonnanotubes [61] At the same time biosensors offerthe possibility of determining not only specific chemicalsbut also their biological effects such as toxicity cytotoxicitygenotoxicity or endocrine disrupting effects that is relevantinformation that in some occasions is more meaningful thanthe chemical composition Enzymatic biosensors are basedon the selective inhibition of specific enzymes by differ-ent classes of compounds with the decrease in activity of


the immobilized enzyme in the presence of the target analyteas the parameter that is frequently used for quantification

A novel myoglobin-based electrochemical biosensorbased on a nanocomposite prepared frommultiwalled carbonnanotubes that were coated with ceria nanoparticles has beendeveloped [62] Another application of whole-cell biosen-sors is the determination of the biological oxygen demand(BOD) Pesticides (herbicides fungicides and insecticides)are widely used in the agriculture and industry around theworld due to their high insecticidal activity Biosensors arepotentially useful as they detected pesticides quickly and havebeen active in the research area for some years Another valu-able HRP-based biosensor was developed in which polyvinylpyrrolidone (PVP) nanofibers were spun with incorporationof the enzyme HRP Scanning electron microscopy (SEM)of the spun nanofibers was used to confirm the non wovenstructure which had an average diameter of 155 plusmn 34 nmThe HRP containing fibers were tested for their changein activity following electrospinning and during storage Acolorimetric assay was used to characterize the activity ofHRP by reaction with the nanofiber mats in a microtiterplate and monitoring the change in absorption over time[63] Rapid and sensitive detection methods are of utmostimportance for the identification of pathogens related tohealth and safety Peroxidase used in development of a nucleicacid sequence-based lateral flow assay which achieves a lowlimit of detection using chemiluminescence and enzymaticsignal amplification [64]

410 Use in Pulp-Paper Industries The pulping byproducts(black liquor) and pulp mill waste water cause seriousenvironmental problemdue to its high pollution load Solvingthe pulp and paper industriesrsquo environmental problems isessential to maintaining the forest industry and accommo-dating the changing economic needs of forest communities[65] Pulp manufacturing in pulp paper industries involvetwo main processes that is wood digestion and bleachingIn the process of wood digestion wood chips are cooked inthe solution of sodium hydroxide and sodium sulphate atelevated temperature and pressure to break chips into fibermass The chemical reaction with wood fibers dissolves allthe depository materials which are hard to degrade andthese derivatives are washed away from the fiber duringwashing and dewatering process Various extracts duringwashing include mainly lignins cellulose phenolics resinsfatty acids and tannins mixed together to make a dark blackviscous alkaline waste known as black liquor The alkalineeffluent consists only in 10ndash15 of total waste water butcontributes in almost above 90ndash95 of the total pollutionload in terms of high pH BOD COD and color whichmakesit significantly toxic to the environment [66] Hence theadequate treatment of black liquor prior to its discharge intoenvironment is warranted The biological methods for thetreatment of black liquor involving the use of fungi bacteriaalgae and enzymes as single step treatment or in combinationwith other physical and chemical methods seem to be moreeconomical and ecofriendly [4] Among biological methodstried so far most of the literature is confined to a few generaof white rot fungi because of their nonspecific extracellular

enzymatic system (LiP MnP and Laccase) involved in ligninbiodegradation [67]

5 Conclusion

Importance of peroxidase in the detoxification of pollutedenvironments relies on their capability to catalyse the reduc-tion of peroxides such as hydrogen peroxide and the oxi-dation of a variety of organic and inorganic compoundsand the polymerization of toxic compounds or by crossreaction with other phenolics or with cosubstrates withtoxic and harmless characteristics and generates polymericproducts (dimmers trimmers hybrid oligomers) which willvery likely accumulate in soil andor in water systemsPeroxidases have potential for bioremediation of waste watercontaminated with phenols cresols and other industrialeffluents for decolourization of textile dyes removal ofendocrine disruptive chemicals degradation of pesticidespolychlorinated biphenyls chlorinated alkanes and alkenes ofsoil phenoxy alkanoic herbicides triazine herbicides chlo-rinated dioxins and chlorinated insecticides Peroxidasesare also used as biosensors Rapid progresses in the use ofperoxidase for the degradation of pollutants have thrownmore light on sustainable bioremediation strategies for pol-luting compounds and environmental protection by usingdifferent enzymes Environmental protection is influenced bythree interwoven factors environmental legislation ethicsand education Each of these factors plays an importantrole in influencing national-level environmental decisionsand personal-level environmental values and behaviors Forenvironmental protection to become a reality it is importantfor societies and the nations to develop each of these areasthat together will inform and drive environmental decisions

Acknowledgments

This work has been funded by Council for Scientific andIndustrial Research New Delhi under CSIR-NET JuniorResearch Fellowship awarded to one of the authors (NB)Theauthors are thankful to CSIR New Delhi and Department ofBiotechnology Himachal Pradesh University Shimla

References

[1] E Marco-Urrea E Aranda G Caminal and F Guillen ldquoInduc-tion of hydroxyl radical production in Trametes versicolorto degrade recalcitrant chlorinated hydrocarbonsrdquo BioresourceTechnology vol 100 no 23 pp 5757ndash5762 2009

[2] H Hamid and K U Rehman ldquoPotential applications of perox-idasesrdquo Food Chemistry vol 115 no 4 pp 1177ndash1186 2009

[3] T Chanwun N Muhamad N Chirapongsatonkul and NChurngchow ldquoHevea brasiliensis cell suspension peroxidasepurification characterization and application for dye decol-orizationrdquo AMB Express vol 3 p 14 2013

[4] R Chandra A Abhishek and M Sankhwar ldquoBacterial decol-orization and detoxification of black liquor from rayon gradepulp manufacturing paper industry and detection of theirmetabolic productsrdquo Bioresource Technology vol 102 no 11 pp6429ndash6436 2011


[5] P Huber and B Carre ldquoDecolorization of process waters indeinking mills and similar applications a reviewrdquo BioResourcesvol 7 no 1 pp 1366ndash1382 2012

[6] T K Lundell M R Makela and K Hilden ldquoLignin-modifyingenzymes in filamentous basidiomycetes-ecological functionaland phylogenetic reviewrdquo Journal of Basic Microbiology vol 50no 1 pp 5ndash20 2010

[7] M Hofrichter R Ullrich M J Pecyna C Liers and T LundellldquoNew and classic families of secreted fungal heme peroxidasesrdquoApplied Microbiology and Biotechnology vol 87 no 3 pp 871ndash897 2010

[8] E Marco-Urrea and C A Reddy ldquoDegradation of chloro-organic pollutants by white rot fungi microbial degradation ofxenobioticsrdquo Environmental Science and Engineering vol 2 pp31ndash66 2012

[9] S T Ong P S Keng W N Lee S T Ha andW T Hung ldquoDyewaste treatmentrdquoWater vol 3 no 1 pp 157ndash176 2011

[10] V Gopi A Upgade and N Soundararajan ldquoBioremediationpotential of individual and consortium non-adaptedfungalstrains on Azo dye containing textile effluentrdquo Advances inApplied Science Research vol 3 no 1 pp 303ndash311 2012

[11] H T Tien Z Salamon J Kutnik et al ldquoBilayer Lipid Mem-branes (BLM) an experimental system for biomolecular elec-tronic device developmentrdquo Journal of Molecular Electronicsvol 4 supplement 4 pp S1ndashS30 1988

[12] N Kirby G McMullan and R Marchant ldquoDecolourisation ofan artificial textile effluent by Phanerochaete chrysosporiumrdquoBiotechnology Letters vol 17 no 7 pp 761ndash764 1995

[13] T W Ng Q Cai C Wong A T Chow and P Wong ldquoSimul-taneous chromate reduction and azo dye decolourization byBrevibacterium casei Azo dye as electron donor for chromatereductionrdquo Journal of Hazardous Materials vol 182 no 1ndash3 pp792ndash800 2010

[14] F Ghasemi F Tabandeh B Bambai and K R S SambasivaRao ldquoDecolorization of different azo dyes by Phanerochaetechrysosporium RP78 under optimal conditionrdquo InternationalJournal of Environmental Science and Technology vol 7 no 3pp 457ndash464 2010

[15] V V Dawkar U U Jadhav S U Jadhav and S P GovindwarldquoBiodegradation of disperse textile dye Brown 3REL by newlyisolated Bacillus sp VUSrdquo Journal of Applied Microbiology vol105 no 1 pp 14ndash24 2008

[16] M Krishnaveni and R Kowsalya ldquoCharacterization and decol-orization of dye and textile effluent by laccase from Pleurotusflorida- A white-rot fungirdquo International Journal of Pharma andBio Sciences vol 2 no 1 pp 117ndash123 2011

[17] F PomarNCaballeroMA Pedreo andARos Barcelo ldquoH2O2

generation during the auto-oxidation of coniferyl alcohol drivesthe oxidase activity of a highly conserved class III peroxidaseinvolved in lignin biosynthesisrdquo FEBS Letters vol 529 no 2-3pp 198ndash202 2002

[18] J S Shin and B G Kim ldquoKinetic resolution of 120572-methy-lbenzylamine with120596-transaminase screened from soil microor-ganisms application of a biphasic system to overcome productinhibitionrdquo Biotechnology and Bioengineering vol 55 no 2 pp348ndash335 1997

[19] A Bhunia S Durani and PWangikar ldquoHorseradish peroxidasecatalyzed degradation of industrially important dyesrdquo Journal ofBiotechnology and Bioengineering vol 72 pp 562ndash567 2002

[20] P D Gennaro A Bargna F Bruno and G Sello ldquoPurificationof recombinant catalase-peroxidase HPI from Ecoli and its

application in enzymatic polymerization reactionsrdquo Joural ofApplied Microbiology and Biotechnology 2013

[21] N Bansal and S S Kanwar ldquoPurification and characterizationof an extracellular peroxidase of a bacterial isolate Bacillus spF31rdquo Current Biotechnology vol 2 no 2 pp 155ndash161 2013

[22] Y Hong M Dashtban S Chen R Song andW Qin ldquoEnzymeproduction and lignin degradation by four basidiomycetous-fungi in submerged fermentation of peatrdquo International Journalof Biology Medium vol 4 pp 172ndash180 2012

[23] C Deivasigamani and N Das ldquoBiodegradation of Basic Violet3 by Candida krusei isolated from textile wastewaterrdquo Biodegra-dation vol 22 no 6 pp 1169ndash1180 2011

[24] S D Kalme G K Parshetti S U Jadhav and S P GovindwarldquoBiodegradation of benzidine based dye Direct Blue-6 by Pseu-domonas desmolyticum NCIM 2112rdquo Bioresource Technologyvol 98 no 7 pp 1405ndash1410 2007

[25] L N Du S Wang G Li et al ldquoBiodegradation of malachitegreen by Pseudomonas sp strain DY1 under aerobic conditioncharacteristics degradation products enzyme analysis andphytotoxicityrdquo Ecotoxicology vol 20 no 2 pp 438ndash446 2011

[26] K S Shin H K Young and J S Lim ldquoPurification andcharacterization of manganese peroxidase of the white-rotfungus Irpex lacteusrdquo Journal of Microbiology vol 43 no 6 pp503ndash509 2005

[27] A A Telke S M Joshi S U Jadhav D P Tamboli andS P Govindwar ldquoDecolorization and detoxification of Congored and textile industry effluent by an isolated bacteriumPseudomonas sp SU-EBTrdquo Biodegradation vol 21 no 2 pp283ndash296 2010

[28] P Ollikka K Alhonmaki V-M Leppanen T Glumoff TRaijola and I Suominen ldquoDecolorization of azo triphenylmethane heterocyclic and polymeric dyes by lignin peroxidaseisoenzymes from Phanerochaete chrysosporiumrdquo Applied andEnvironmentalMicrobiology vol 59 no 12 pp 4010ndash4016 1993

[29] M H Joe S Y Lim D H Kim and I N Lee ldquoDecolorizationof reactive dyes by Clostridium bifermentans SL186 isolatedfrom contaminated soilrdquo World Journal of Microbiology andBiotechnology vol 24 no 10 pp 2221ndash2226 2008

[30] Y Sugano Y Matsushima and M Shoda ldquoComplete decol-orization of the anthraquinone dye Reactive blue 5 by theconcerted action of two peroxidases from Thanatephorus cuc-umerisrdquo Applied Microbiology and Biotechnology vol 73 no 4pp 862ndash871 2006

[31] C Liers C Bobeth M Pecyna R Ullrich and M HofrichterldquoDyP-like peroxidases of the jelly fungus Auricularia auricula-judae oxidize nonphenolic lignin model compounds and high-redox potential dyesrdquo Applied Microbiology and Biotechnologyvol 85 no 6 pp 1869ndash1879 2010

[32] V Faraco A Piscitelli G Sannia and P Giardina ldquoIdentifica-tion of a newmember of the dye-decolorizing peroxidase familyfrom Pleurotus ostreatusrdquo World Journal of Microbiology andBiotechnology vol 23 no 6 pp 889ndash893 2007

[33] V Gomez-Toribio A B Garcıa-Martın M J Martınez AT Martınez and F Guillen ldquoEnhancing the production ofhydroxyl radicals by Pleurotus eryngii via quinone redoxcycling for pollutant removalrdquo Applied and EnvironmentalMicrobiology vol 75 no 12 pp 3954ndash3962 2009

[34] D Kauspediene E Kazlauskiene A Gefeniene and RBinkiene ldquoComparison of the efficiency of activated carbon andneutral polymeric adsorbent in removal of chromium complexdye from aqueous solutionsrdquo Journal of Hazardous Materialsvol 179 no 1ndash3 pp 933ndash939 2010


[35] E L K Mui W H Cheung M Valix and G McKay ldquoDyeadsorption onto activated carbons from tyre rubber wasteusing surface coverage analysisrdquo Journal of Colloid and InterfaceScience vol 347 no 2 pp 290ndash300 2010

[36] A Longoria R Tinoco and R Vazquez-Duhalt ldquoChloroper-oxidase-mediated transformation of highly halogenated mono-aromatic compoundsrdquoChemosphere vol 72 no 3 pp 485ndash4902008

[37] T K Hakala K Hilden P Maijala C Olsson and A HatakkaldquoDifferential regulation of manganese peroxidases and charac-terization of two variable MnP encoding genes in the white-rot fungus Physisporinus rivulosusrdquo Applied Microbiology andBiotechnology vol 73 no 4 pp 839ndash849 2006

[38] J Mamatha A B Vedamurthy and S D Shruthi ldquoDegradationof phenol by turnip root enzyme extractrdquo Journal of Microbiol-ogy and Biotechnology Research vol 2 no 3 pp 426ndash430 2012

[39] F J Ruiz-DuenasMMorales E Garcıa YMikiM JMartınezand A T Martınez ldquoSubstrate oxidation sites in versatileperoxidase and other basidiomycete peroxidasesrdquo Journal ofExperimental Botany vol 60 no 2 pp 441ndash452 2009

[40] K F Mossallam F M Sultanova and N A Salemova ldquoPeroxi-dase catalysed the removal of phenol of phenol) from syntheticwaste waterrdquo in Proceedings of the 13th International WaterTechnology Conference (IWTC rsquo13) pp 1009ndash1020 HurghadaEgypt 2009

[41] W Zheng and L M Colosi ldquoPeroxidase-mediated removal ofendocrine disrupting compoundmixtures fromwaterrdquo Chemo-sphere vol 85 no 4 pp 553ndash557 2011

[42] H Cabana J P Jones and S N Agathos ldquoElimination ofendocrine disrupting chemicals using white rot fungi andtheir lignin modifying enzymes a reviewrdquo Engineering in LifeSciences vol 7 no 5 pp 429ndash456 2007

[43] Q Huang and W J Weber Jr ldquoTransformation and removalof bisphenol A from aqueous phase via peroxidase-mediatedoxidative coupling reactions efficacy products and pathwaysrdquoEnvironmental Science and Technology vol 39 no 16 pp 6029ndash6036 2005

[44] G Renner ldquoMetabolic studies on pentachloronitrobenzene(PCNB) in ratsrdquo Xenobiotica vol 10 no 7-8 pp 537ndash550 1980

[45] L A McCauley W K Anger M Keifer R Langley MG Robson and D Rohlman ldquoStudying health outcomes infarmworker populations exposed to pesticidesrdquo EnvironmentalHealth Perspectives vol 114 no 6 pp 953ndash960 2006

[46] J Raymond T Rogers D Shonnard and A Kline ldquoA review ofstructure-based biodegradation estimationmethodsrdquo Journal ofHazardous Materials vol 84 no 2-3 pp 189ndash215 2001

[47] J Jauregui B Valderrama A Albores and R Vazquez-DuhaltldquoMicrosomal transformation of organophosphorus pesticidesby white rot fungirdquo Biodegradation vol 14 no 6 pp 397ndash4062003

[48] SHarayama ldquoPolycyclic aromatic hydrocarbon bioremediationdesignrdquoCurrent Opinion in Biotechnology vol 8 no 3 pp 268ndash273 1997

[49] R Weber C Gaus M Tysklind et al ldquoDioxin- and POP-contaminated sites-contemporary and future relevance andchallenges overview on background aims and scope of theseriesrdquoEnvironmental Science andPollutionResearch vol 15 no5 pp 363ndash393 2008

[50] C FHarford-Cross A B Carmichael F KAllan P A EnglandD A Rouch and LWong ldquoProtein engineering of cytochromeP458(cam) (CYP101) for the oxidation of polycyclic aromatic

hydrocarbonsrdquo Protein Engineering vol 13 no 2 pp 121ndash1282000

[51] K Kordon A Mikolasch and F Schauer ldquoOxidative dehalo-genation of chlorinated hydroxybiphenyls by laccases of white-rot fungirdquo International Biodeterioration and Biodegradationvol 64 no 3 pp 203ndash209 2010

[52] J S Yadav C Bethea and C A Reddy ldquoMineralization oftrichloroethylene (TCE) by the white rot fungus Phanerochaetechrysosporiumrdquo Bulletin of Environmental Contamination andToxicology vol 65 no 1 pp 28ndash34 2000

[53] X Huang J Pan B Liang J Sun Y Zhao and S Li ldquoIsolationcharacterization of a strain capable of degrading imazethapyrand its use in degradation of the herbicide in soilrdquo CurrentMicrobiology vol 59 no 4 pp 363ndash367 2009

[54] G D Bending M Friloux and A Walker ldquoDegradation ofcontrasting pesticides by white rot fungi and its relationshipwith ligninolytic potentialrdquo FEMSMicrobiology Letters vol 212no 1 pp 59ndash63 2002

[55] N Kasai S Ikushiro R Shinkyo et al ldquoMetabolism of mono-and dichloro-dibenzo-p-dioxins by Phanerochaete chrysospo-rium cytochromes P450rdquo Applied Microbiology and Biotechnol-ogy vol 86 no 2 pp 773ndash780 2010

[56] J C Quintero M T Moreira G Feijoo and J M LemaldquoScreening of white rot fungal species for their capacity todegrade lindane and other isomers of hexachlorocyclohexane(HCH)rdquo Ciencia e Investigacion Agraria vol 35 no 2 pp 123ndash132 2008

[57] C Mougin C Pericaud C Malosse C Laugero and C AstherldquoBiotransformation of the insecticide lindane by the white rotsbasidiomycete Phanerochaete chrysosporiumrdquo Journal of PestScience vol 47 no 1 pp 51ndash59 1996

[58] A S Purnomo T Mori I Kamei T Nishii and R KondoldquoApplication of mushroom waste medium from Pleurotusostreatus for bioremediation of DDT-contaminated soilrdquo Inter-national Biodeterioration and Biodegradation vol 64 no 5 pp397ndash402 2010

[59] S Ahammad ldquoHydrogen peroxide biosensors based onhorseradish peroxidase and haemoglobinrdquo Jounal of Biosensorsamp Bioelectronics 2013

[60] J Song J Xu P Zhao L Lu and J Bao ldquoA hydrogen peroxidebiosensor based on direct electron transfer from hemoglobin toan electrode modified with Nafion and activated nanocarbonrdquoMicrochimica Acta vol 172 no 1 pp 117ndash123 2011

[61] S Xu X Zhang T Wan and C Zhang ldquoA third-generationhydrogen peroxide biosensor based on horseradish peroxidasecross-linked to multi-wall carbon nanotubesrdquo MicrochimicaActa vol 172 no 1 pp 199ndash205 2011

[62] J Qiu S Cui and R Liang ldquoHydrogen peroxide biosensorbased on the direct electrochemistry of myoglobin immobi-lized on ceria nanoparticles coated with multiwalled carbonnanotubesby a hydrothermal synthetic methodrdquo MicrochimicaActa vol 171 no 3 pp 333ndash339 2010

[63] M Dai S Jin and S R Nugen ldquoWater-soluble electrospunnanofibers as a method for on-chip reagent storagerdquo Biosensors vol2 no 4 pp 388ndash395 2012

[64] Y Wang C Fill and S R Nugen ldquoDevelopment of chemilu-miniscent lateral flow assay for the detection of nucleic acidsrdquoBiosensors vol 2 no 1 pp 32ndash42 2012

[65] A Khalid M Arshad and D E Crowley ldquoBiodegradationpotential of pure and mixed bacterial cultures for removal of 4-nitroaniline from textile dye wastewaterrdquo Water Research vol43 no 4 pp 1110ndash1116 2009


[66] R Grover S S Marwaha and J F Kennedy ldquoStudies onthe use of an anaerobic baffled reactor for the continuousanaerobic digestion of pulp and paper mill black liquorsrdquoProcess Biochemistry vol 34 no 6-7 pp 653ndash657 1999

[67] K E Hammel and D Cullen ldquoRole of fungal peroxidases inbiological ligninolysisrdquo Current Opinion in Plant Biology vol11 no 3 pp 349ndash355 2008

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Anatomy Research International

PeptidesInternational Journal of


Hindawi Publishing Corporation httpwwwhindawicom

International Journal of

Volume 2014

Zoology


Molecular Biology International

GenomicsInternational Journal of


The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014


BioinformaticsAdvances in

Marine BiologyJournal of



Signal TransductionJournal of


BioMed Research International

Evolutionary BiologyInternational Journal of



Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014


Genetics Research International


Advances in

Virolog y

Hindawi Publishing Corporationhttpwwwhindawicom

Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational



Enzyme Research



Microbiology


Substratereduced + H2O2

HRP(EC 11117)

Substrateoxidized + 2H2O

Figure 1 A general reaction catalyzed by HRP

plants animals and microbes Peroxidases produced frommicrobial sources such as bacteria (Bacillus sphaericus Bacil-lus subtilis Pseudomonas sp Citrobacter sp) Cyanobacteria(Anabaena sp) fungi (Candida krusei Coprinopsis cinereaPhanerochaete chrysosporium) actinomycetes (StreptomycesspThermobifida fusca) and yeast are used in decompositionof pollutants production of animal feedstock and raw mate-rials for the chemical agricultural paper industries textiledye degradation paper-pulp industry for lignin degradationdye decolorization sewage treatment and also as biosen-sors Many plant sources for peroxidases production havebeen reported such as horseradish papaya (Carica papaya)banana (Musa paradisiacal) and bare (Acorus calamus)Peroxidase obtained from horseradish (HRP) is extensivelyused in diagnostic kits in ELISA for labeling an antibodysynthesis of various aromatic chemicals and removal ofperoxides from materials such as foodstuff and industrialwastes (Figure 1)

3 Characteristics of Peroxidase(s)

Peroxidases are oxidoreductases that catalyse a variety ofreactions such as reduction of peroxides such as hydrogenperoxide and oxidation of a variety of organic and inorganiccompounds These are heme proteins and contain iron (III)protoporphyrin IX as the prosthetic group They have amolecular weight ranging from 30 to 150 kDa The termperoxidase represents a group of specific enzymes such asNADH peroxidase (EC 11111) glutathione peroxidase (EC11119) and iodine peroxidase (EC 11118) as well as a varietyof nonspecific enzymes that are simply known as peroxidases

4 Applications and Peroxidase Biocatalysis inManagement of Environmental Pollutants

41 Decolorization of Synthetic Dyes Dye wastes representone of the most problematic groups of pollutants consideredas xenobiotics that are not easily biodegradable [9] Thesedyes are mostly used in textile dyeing paper printing colourphotography and as additive in petroleum products Whenthese synthetic dyes are discharged into industrial effluentsthey cause environmental pollution Textile industries playa vital role in the economic increases in India Water isone of the major products of nature used enormously byhuman beings and it is not unnatural that any growingcommunity generates enormous waste water or sewage [10]To achieve the biodegradation of environmentally hazardouscompounds white rot fungi appear as a valuable alternative

The capability of oxidation is based on the ability of white rotfungi to produce oxidative enzymes such as laccase mang-anese peroxidase and lignin peroxidase [11] These oxidasesand peroxidases have been reported as excellent oxidantagents to degrade dyes [12]

Several bacterial peroxidases have been used for decol-orization of synthetic textile dyes Removal of chromate Cr(VI) and azo dye Acid Orange 7 (AO7) using Brevibacteriumcasei under nutrient-limiting conditions has been studiedAO7 was used as an electron donor by the reduction enzymeof Brevibacterium casei for the reduction of Cr (VI) Thereduced chromate Cr (III) complexed with the oxidizedAO7 formed a purple intermediate [13] Decolorization ofdifferent azo dyes by Phanerochaete chrysosporium RP 78under optimized conditions was studied [14] by reactionmechanism via azo dye Peroxidase was produced underaerobic conditions as a secondarymetabolite in the stationaryphase Bacillus sp VUS isolated from textile effluent con-taminated soil showed capability for degrading a variety ofdyes [15] The production of ligninolytic peroxidases directlyoxidizing aromatic compounds has been described in fungi[16] Other peroxidases were detected in microorganismsresponsible for the biodegradation of industrial dyes togetherwith lignin peroxidase [17] An edible macroscopic fungiPleurotus ostreatus produced an extracellular peroxidase thatcan decolorize remazol brilliant blue and other structurallydifferent groups including triarylmethane heterocyclic azoand polymeric dyes Bromophenol blue was decolorized best(98) while methylene blue and toluidine blue O were leastdecolorized 10 [18] HRP was found to degrade industriallyimportant azo dyes such as remazol blue This dye containsat least one aromatic group in its structure making it apossible substrate of HRP [19] The dyeing and bleachingunitrsquos contaminants seeping into the ground have pollutedthe ground water rendering it unsuitable for consumption(Table 1)

42 Bioremediation of Waste Water Removal of PhenolicContaminants and Related Compounds Industrial pollutionhas been a major factor causing the degradation of the envi-ronment around us affecting the water we use its quality andhuman health is directly related issues Improved quality andincreased quantity of water would bring forth health benefitsSafe water eliminates the infective agents associated withwater borne diseases availability of greater quantity of watercan improve health by allowing improved personal hygieneWater pollution caused industrial waste products to releaseinto lakes rivers and other water bodies that make marinelife no longer hospitable Peroxidases have been applied to






(3)












[23]





organisms

[25]




(10)




[28]




(14)Extracellular










OH

OH O

O

BQ

QRMycelium

Ominus

Fe+3

Fe+3

Fe+2Fe+2

BQminus

H2O2

BQH2

O2

O2

minuslowast

OHlowast

OlowastPero

xidase




2


2-dismutation to H

2O2







2O (1)





















2O2is consid-









5 Conclusion


Acknowledgments


References

















































































Volume 2014

Zoology






















Advances in

Virolog y



Volume 2014




Enzyme Research



Microbiology






(3)












[23]





organisms

[25]




(10)




[28]




(14)Extracellular










OH

OH O

O

BQ

QRMycelium

Ominus

Fe+3

Fe+3

Fe+2Fe+2

BQminus

H2O2

BQH2

O2

O2

minuslowast

OHlowast

OlowastPero

xidase




2


2-dismutation to H

2O2







2O (1)





















2O2is consid-









5 Conclusion


Acknowledgments


References

















































































Volume 2014

Zoology






















Advances in

Virolog y



Volume 2014




Enzyme Research



Microbiology


OH

OH O

O

BQ

QRMycelium

Ominus

Fe+3

Fe+3

Fe+2Fe+2

BQminus

H2O2

BQH2

O2

O2

minuslowast

OHlowast

OlowastPero

xidase




2


2-dismutation to H

2O2







2O (1)





















2O2is consid-









5 Conclusion


Acknowledgments


References

















































































Volume 2014

Zoology






















Advances in

Virolog y



Volume 2014




Enzyme Research



Microbiology











2O2is consid-









5 Conclusion


Acknowledgments


References

















































































Volume 2014

Zoology






















Advances in

Virolog y



Volume 2014




Enzyme Research



Microbiology






5 Conclusion


Acknowledgments


References

















































































Volume 2014

Zoology






















Advances in

Virolog y



Volume 2014




Enzyme Research



Microbiology













































































Volume 2014

Zoology






















Advances in

Virolog y



Volume 2014




Enzyme Research



Microbiology












































Volume 2014

Zoology






















Advances in

Virolog y



Volume 2014




Enzyme Research



Microbiology











Volume 2014

Zoology






















Advances in

Virolog y



Volume 2014




Enzyme Research



Microbiology








Volume 2014

Zoology






















Advances in

Virolog y



Volume 2014




Enzyme Research



Microbiology

Documents

Review Article Peroxidase(s) in Environment Protectiondownloads.hindawi.com/journals/tswj/2013/714639.pdf · Review Article Peroxidase(s) in Environment Protection ... agricultural,