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The Plant Family Brassicaceae ENVIRONMENTAL POLLUTION VOLUME 21 Editors Brain J. Alloway, Department of Soil Science, The University of Reading, U.K. Jack T. Trevors, School of Environmental Sciences, University of Guelph, Ontario, Canada Editorial Board I. Colbeck, Interdisciplinary Centre for Environment and Society, Department of Biological Sciences, University of Essex, Colchester, U.K. R.L. Crawford, Food Research Center (FRC) 204, University of Idaho, Moscow, Idaho, U.S.A. W. Salomons, GKSS Research Center, Geesthacht, Germany For furthervolumes: http://www.springer.com/series/5929 Naser A. Anjum Iqbal Ahmad l M. Eduarda Pereira Armando C. Duarte l Shahid Umar Nafees A. Khan l Editors The Plant Family Brassicaceae Contribution Towards Phytoremediation Editors NaserA.Anjum IqbalAhmad CentreforEnvironmentaland CentreforEnvironmentalandMarine MarineStudies(CESAM)& Studies(CESAM)&Department DepartmentofChemistry ofChemistryandBiology UniversityofAveiro UniversityofAveiro 3810-193Aveiro 3810-193Aveiro Portugal Portugal M.EduardaPereira ArmandoC.Duarte CentreforEnvironmentaland CentreforEnvironmentaland MarineStudies(CESAM)& MarineStudies(CESAM)& DepartmentofChemistry DepartmentofChemistry UniversityofAveiro UniversityofAveiro 3810-193Aveiro 3810-193Aveiro Portugal Portugal ShahidUmar NafeesA.Khan DepartmentofBotany DepartmentofBotany FacultyofScience FacultyofLifeSciences HamdardUniversity AligarhMuslimUniversity NewDelhi,110062 Aligarh202002 India UttarPradesh,India ISSN1566-0745 ISBN978-94-007-3912-3 ISBN978-94-007-3913-0(eBook) DOI10.1007/978-94-007-3913-0 SpringerDordrechtHeidelbergNewYorkLondon LibraryofCongressControlNumber:2012936075 #SpringerScience+BusinessMediaB.V.2012 Thisworkissubjecttocopyright.AllrightsarereservedbythePublisher,whetherthewholeorpartof the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation,broadcasting,reproductiononmicrofilmsorinanyotherphysicalway,andtransmissionor informationstorageandretrieval,electronicadaptation,computersoftware,orbysimilarordissimilar methodologynowknownorhereafterdeveloped.Exemptedfromthislegalreservationarebriefexcerpts inconnectionwithreviewsorscholarlyanalysisormaterialsuppliedspecificallyforthepurposeofbeing enteredandexecutedonacomputersystem,forexclusiveusebythepurchaserofthework.Duplication ofthispublicationorpartsthereofispermittedonlyundertheprovisionsoftheCopyrightLawofthe Publisher’s location, in its current version, and permission for use must always be obtained from Springer.PermissionsforusemaybeobtainedthroughRightsLinkattheCopyrightClearanceCenter. ViolationsareliabletoprosecutionundertherespectiveCopyrightLaw. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publicationdoesnotimply,evenintheabsenceofaspecificstatement,thatsuchnamesareexempt fromtherelevantprotectivelawsandregulationsandthereforefreeforgeneraluse. While the advice and information in this book are believed to be true and accurate at the date of publication,neithertheauthorsnortheeditorsnorthepublishercanacceptanylegalresponsibilityfor anyerrorsoromissionsthatmaybemade.Thepublishermakesnowarranty,expressorimplied,with respecttothematerialcontainedherein. Printedonacid-freepaper SpringerispartofSpringerScience+BusinessMedia(www.springer.com) Foreword Plant Biology is reaching an extraordinary degree of conceptual complexity, especially since the crucial technological advances during the last two decades, halfway between the twentieth and twenty-first centuries. Integration from the molecular to the whole plant level of the mechanisms governing growth and developmentalprocessesprovidesnowabetterandanincreasinglydiversevision ofthefunctioningofplantsindifferenthabitatsandunderchangingenvironmental conditions. This is featuring a solid body of scientific knowledge which covers the various levels of plant research: cellular, biochemical, genetic, molecular, physiological,ecologicalandevolutionary.Thisplethoraofknowledgeandmeth- odological consolidation now allows to more precisely understand the physio- logical and molecular mechanisms, to see the richness of phenotypic responses of plants to the diverse environmental situations, to deepen consistently in the evolutionary lines, to discern the molecular basis of variation for including all within the system’s biology, and to consider the diversity and specificities of plantfunctionalism. Spectacularprogressisbeingmadeinthefundamentalknowledgeofreception of external signals and internal signaling pathways, the vital mechanisms of regulationofhomeostasis,theintegrationofmetabolicfunctions,andthedifferen- tiation and development of plants exposed to biotic or abiotic stress factors. As expected, this advanced insight into basic processes governing plants under adverse environmental conditions has substantially improved our expertise in practical applications, not only in the field of agronomy, but also in that of environmental restoration. Phytoremediation as a clean, sustainable technology is becominganattractivewaytorecoverdegradedland.Nowonderthatresearchand publications on this topic have increased exponentially during the last years and thatwefacearapidlyexpandingareawherebasicknowledgeisbeingturnedinto effectiveenvironmentaltechnology. Itisnotaccidentalthatthisnewbookentitled‘ThePlantFamilyBrassicaceae: ContributionTowardsPhytoremediation’focusestheissueofphytoremediationon theBrassicaceae.Thislargebotanicalfamilywith360genusand3,700speciesnot v vi Foreword only includes important food and industrial plants, but also Arabidopsis thaliana theplantmodelspeciesparexcellence.Smallgenomesize,self-compatibility,short life cycle, small biomass, and large production of seeds make this Arabidopsis thalianaanidealresearchobjectforstudiesintofunctionalgenomics.Identification of genes involved in metal uptake, transport and compartmentation and knowl- edgeofkeyenzymesinmetabolicpathwaysabletotransformanddetoxifyorganic pollutantsprovidethefundamentaltoolsforthedevelopmentofphytoremediation technologies. This extensive basic research in Arabidopsis thaliana allowed fast advancesintheknowledgeofthefunctioningofBrassicaceaespeciesofhighinterest in phytoremediation processes, such as the close relative Arabidopsis halleri and Thlaspi caerulescens (Noccaea caerulescens), both species with ability to hyper- accumulate cadmium and zinc. Also the nickel hyperaccumulators of the genus Alyssum are within the Brassicaceae. Furthermore, the non-hyperaccumulating species of the genus Brassica, especially Brassica juncea and Brassica napus, the seed-oil canola which combines both industrial applications and phytoremediation potential, are being explored for phytoremediation technologies. These Brassica species have considerably lower shoot metal concentrations than metal hyperaccu- mulators.Thisiscompensated,however,bythemuchhigherbiomassinBrassicasp. Thus, the focus on Brassicaceae in this new book is fully awarded by this predominant presence of representatives of this family in current research in phytoremediation. This leads to the question about the characteristics that makes Brassicaceaespeciessooutstandinglyattractiveforrecoveringcontaminatedland. Toouropinion,twofeaturescommontoBrassicaceaemaycontributetothis:special pathwaysofsecondarysulphurmetabolismforglucosinolatesynthesisand,probably inpartrelatedtothis,thefactthatmostBrassicaceae,arenonmycorrhizal.Ahigh tissuelevelofantioxidantsisafurthercharacteristicofimportanceforperformance understressfulconditions. Brassicaceaehaveahighsulphurrequirement.Sulphuravailabilityinminespoils usestorangefromnormaltoextremelyhighandmostcontaminatedsoilscanfully satisfy the higher sulphur demand of the Brassicaceae. Moreover, in Brassicaceae the levels of sulphur containing metabolites that are important for basic metal tolerance, such as glutathion and phytochelatins seem to be affected only under extremesulphurdeficiency. Many Brassicaceae species are pioneers on waste areas and seem well- adaptedtorecentlyformed,badlystructuredsubstrates.Noneedformycorrhization undertheseconditionsimplieshighnutrientefficiency,especiallyforphosphorous. InBrassicacropsdifferencesinphosphorusacquisitionarecloselyrelatedtoboth rootarchitecturetraitsandinternalphosphorususeefficiency.Metalhyperaccumu- latorslikeThlaspicaerulescensseemwelladaptedtothelowphosphorusavailabil- ity of mine waste areas and surplus phosphorus supply may not enhance their biomassproductionand/ormetalextractionpotential. Many species that perform well on mine spoils are mycorrhizal. Considerable amounts of heavy metals can be retained inthe cellwalls, especially of ecto- and ericoidmycorrhizalfungi,thuslimitingmetaluptakebytheplants.Metallophytes thatarenon-mycorrhizal,suchasmostBrassicaceaeandCaryophyllaceaespecies Foreword vii (e.g.Silenevulgaris),havetheirownmechanismstoexcludemetalsfromeitheror both roots and shoots. Nonetheless, on polluted soils they tend to take up higher amounts of heavy metals than mycorrhizal plants. Thus tolerance to higher tissue concentrations of metals is a prerequisite for good performance of these species on metal contaminated sites. The evolution of metal hyperaccumulation behavior with an especially high number ofrepresentativesinthe Brassicaceae is afurther argument for meriting special attention. These plants do not efficiently exclude metalsneitherfromtherootsnortheshoots,butevenforageandaccumulatemetals from soils with low concentrations. Binding of the metals in non-toxic form and efficientcompartmentationintocellwallsandvacuolesarefundamentalmechanisms inthehyperaccumulationlifestyle. Sothegreatachievement andnoveltyofthisbookeditedbyeminentscientists Naser A. Anjum, Iqbal Ahmad, M. Eduarda Pereira, Armando C. Duarte, Shahid UmarandNafeesA.Khanresidesinthefocusonphytoremediationreferredtothe Brassicaceaefamilycontainingspeciesthatarewelladaptedtoharshconditionson wastelandandwithefficientmetaluptakeandtolerancemechanisms.Thedifferent chapters written by experienced specialists provide a unique compilation of the dispersedliteratureonthistopic.Bysure,thereadersofthebookwillbenefitfrom thisjointvisionoftheactualandthepotentialcontributionofthisbotanicalfamily totheemergingdisciplineofphytoremediation. Therefore, we believe that this book targets a potentially broad spectrum of audience ranging from the researchers in this field of increasing importance to undergraduateandgraduatestudentswhowantanoverviewofcurrentknowledge referredtotheBrassicaceaeanditsutilityinphytoremediation.Nolessimportantis thistextforallthoseinterestedintheagriculturaluseoftheBrassicaceaeandtheir contributiontothesustainableproductionofhealthyfood. CharlottePoschenriederandJuanBarcelo´ Lab.Fisiolog´ıaVegetal,FacultaddeBiociencias UniversidadAuto´nomadeBarcelona,E-08193Bellaterra Spain. e-mail:[email protected](C.P.), [email protected](J.B.) Preface The sustainability of agriculture (the only source of food) and the civilization ofmankindrelyontwopreciousnaturalresourcesnamelylandandwater.Unfor- tunately, both land and water have been subjected to maximum exploitation and severely degraded or polluted by kinds of contaminants originating mainly from anthropogenic activities. Majority of contaminants get accumulated in plants and either directly or indirectly, find their way into food web thus cause severe secondaryconsequences.Abovefactstogethercanproduceseveredeteriorationof naturalresources,disturbanceofecosystemsthus,creatingenormoushumanhealth and welfare challenges globally. The remediation of varied environmental con- taminants using plants (phytoremediation; “phyto” meaning plant, and the Latin suffix “remedium” meaningtoclean orrestore)have been acceptedadvantageous over commonly used physical remediation methods in costs, practice and the scale at which the processes operate. This approach is emerging as an innovative tool with greater potential for achieving sustainable development and also to decontaminateair,soils/sedimentsandwater,andforotherenvironmentalrestora- tionapplications. The angiosperm family Brassicaceae – commonly termed as the mustard family (or Cruciferae, because of their characteristic flowers consisting of four petalsintheformofaGreekcross),isalargedicotplantfamilycomprisingnearly 338 genera (assigned to 10–19 tribes) and 3,700 species with worldwide distribu- tion, of major agro-economic and scientific importance. The family Brassicaceae represents hundreds of plant families reported so far for their potential use in the remediation of varied environmental contaminants including toxic metals- metalloids.ThemostofthemembersoftheBrassicaceaeplantfamily wellrepre- sent the metal hyperaccumulation among 0.2% of all angiosperms and thus, have key role in phytoremediation technology. Many of the plant species within Brassicaceae family such as Alyssum, Arabidopsis, Berkheya, Bornmuellera, Cardamine, Cochlearia, Crambe, Peltaria, Pseudosempervivum, Stanleya, Strep- tanthus,Thlaspi including oilseed Brassicas grow fast, yield high biomass and are well adapted to a range of environmental conditions. Some species are tolerant ix x Preface tohighlevelsoftracemetals,andthereisthepotentialtoselectsuperiorgenotypes for phytoremediation. Of 87 different metals/metalloids-hyperaccumulator plant species in the family Brassicaceae, in particular model metals/metalloids hyper- accumulators Alyssum sp., Thlaspi sp. and Arabidopsis sp. have been studied extensivelyfortheirabilitytohyperaccumulatevariedenvironmentalcontaminants including metals and metalloids. In addition, they are well suited to genetic manipulation and in vitro culture techniques and are attractive candidates for the introductionofgenesaimedatphytoremediation. Although, the number of publications focused on plant-based remediation of varied environmental contaminants has been growing exponentially in the last decade but the significant role/contribution of the members of the plant family Brassicaceae for decontaminating varied environmental compartments either remaineduntouchedorlittledebated. Written by an international team of authors and as a significant addition to theSpringers’EnvironmentalPollutionbookseriesthecurrentvolume‘ThePlant Family Brassicaceae: Contribution Towards Phytoremediation’ aims mainly to update our understanding on the recent breakthroughs in the area of differential adaptationoftheplantspecieswithinBrassicaceaefamilytocontaminantsthrough mechanisms involving physiological, biochemical and molecular processes and theircumulativeroleincontaminantsaccumulation,toleranceandremediation. Besides highlighting the taxonomy of the family Brassicaceae, the current status of contaminants (metals/metalloids)-addition to varied environment and its consequences, the information included in this book will bring very stimulating insights into the mechanism involved in the physiological, biochemical and molecular processes responsible for environmental contaminants accumulation, toleranceandremediationinBrassicaceaeans. Additionally,the book alsointends toexplorethevariouspotentialstrategies(suchasthephysiological,biochemical, plant breeding and genetic engineering approaches) for the enhancement of contaminants-tolerance,-accumulationand-remediationpotentialsofthemembers of the plant family Brassicaceae. This volume promises to be a useful asset for researchers,students,otheracademiciansandpolicymakersinvolvedinsustainable remediationofvariedenvironmentalcompartments. Wearethankfultocontributorsfortheirinterests,significantcontributionsand cooperation that eventually made the present volume possible. Thanks are also duetoallthewell-wishers,teachers,seniors,researchstudentsandfamily.Without their unending support, motivation and encouragement the present grueling task wouldhaveneverbeenaccomplished. We would like to offer our sincere thanks to Professors Brian J Alloway and JackTTrevors,EnvironmentalPollutionbookserieseditorsfortheirkindconsent to include the current volume to the series. Exceptional kind support provided by TamaraWelschot,JudithTerposandtheirteamatSpringerdeservespraiseswhich madeoureffortssuccessful. Last, but not least, the financial supports to our research from Foundation for Science & Technology (FCT), Portugal, the Aveiro University Research

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