Advances in Biochemical Engineering/Biotechnology 172 Series Editor: T. Scheper Gashaw Mamo Bo Mattiasson Editors Alkaliphiles in Biotechnology 172 Advances in Biochemical Engineering/Biotechnology SeriesEditor T.Scheper,Hannover,Germany EditorialBoard S.Belkin,Jerusalem,Israel T.Bley,Dresden,Germany J.Bohlmann,Vancouver,Canada M.B.Gu,Seoul,Korea(Republicof) W.-S.Hu,Minneapolis,USA B.Mattiasson,Lund,Sweden H.Seitz,Potsdam,Germany R.Ulber,Kaiserslautern,Germany A.-P.Zeng,Hamburg,Germany J.-J.Zhong,Shanghai,China W.Zhou,Shanghai,China Aims and Scope This book series reviews current trends in modern biotechnology and biochemical engineering. Its aim is to cover all aspects of these interdisciplinary disciplines, whereknowledge,methodsandexpertisearerequiredfromchemistry,biochemistry, microbiology,molecularbiology,chemicalengineeringandcomputerscience. Volumes are organized topically and provide a comprehensive discussion of developments in the field over the past 3–5 years. The series also discusses new discoveries and applications. Special volumes are dedicated to selected topics whichfocusonnewbiotechnologicalproductsandnewprocessesfortheirsynthesis andpurification. In general, volumes are edited by well-known guest editors. The series editor and publisher will, however, always be pleased to receive suggestions and supplemen- taryinformation.ManuscriptsareacceptedinEnglish. In references, Advances in Biochemical Engineering/Biotechnology is abbreviated asAdv.Biochem.Engin./Biotechnol.andcitedasajournal. Moreinformationaboutthisseriesathttp://www.springer.com/series/10 (cid:129) Gashaw Mamo Bo Mattiasson Editors Alkaliphiles in Biotechnology With contributions by (cid:1) (cid:1) (cid:1) D. A. Cowan S. O. Hashim V. V. Kevbrin (cid:1) (cid:1) (cid:1) (cid:1) E. Khalikova T. Korpela P. H. Lebre G. Mamo (cid:1) (cid:1) (cid:1) B. Mattiasson F. J. Mulaa S. Somersalo W. C. Wanyonyi Editors GashawMamo BoMattiasson Billeberga DepartmentofBiotechnology IndienzAB LundUniversity Billeberga,Sweden Lund,Sweden ISSN0724-6145 ISSN1616-8542 (electronic) AdvancesinBiochemicalEngineering/Biotechnology ISBN978-3-030-49735-4 ISBN978-3-030-49736-1 (eBook) https://doi.org/10.1007/978-3-030-49736-1 ©SpringerNatureSwitzerlandAG2020 Thisworkissubjecttocopyright.AllrightsarereservedbythePublisher,whetherthewholeorpartofthe materialisconcerned,specificallytherightsoftranslation,reprinting,reuseofillustrations,recitation, broadcasting,reproductiononmicrofilmsorinanyotherphysicalway,andtransmissionorinformation storageandretrieval,electronicadaptation,computersoftware,orbysimilarordissimilarmethodology nowknownorhereafterdeveloped. Theuseofgeneraldescriptivenames,registerednames,trademarks,servicemarks,etc.inthispublication doesnotimply,evenintheabsenceofaspecificstatement,thatsuchnamesareexemptfromtherelevant protectivelawsandregulationsandthereforefreeforgeneraluse. Thepublisher,theauthors,andtheeditorsaresafetoassumethattheadviceandinformationinthisbook arebelievedtobetrueandaccurateatthedateofpublication.Neitherthepublishernortheauthorsorthe editorsgiveawarranty,expressedorimplied,withrespecttothematerialcontainedhereinorforany errorsoromissionsthatmayhavebeenmade.Thepublisherremainsneutralwithregardtojurisdictional claimsinpublishedmapsandinstitutionalaffiliations. ThisSpringerimprintispublishedbytheregisteredcompanySpringerNatureSwitzerlandAG. Theregisteredcompanyaddressis:Gewerbestrasse11,6330Cham,Switzerland Preface The notion that on earth there is life where there is liquid water seems holding true.Thebiospherehasmanyextremeenvironmentswhichwereoncethoughtfree of life. However, studies indicated that even these extreme sites that are bizarre to humans are found inhabited by richly diverse life forms known as extremophiles. Alkaliphiles are a class of extremophiles that thrive in alkaline habitats. To thrive in the extremely alkaline habitats, alkaliphiles evolved unique structural and functional adaptive features. These features are of great importance to biotechnol- ogy. High pH conditions are part of several industrial and environmental pro- cesses. Thus, alkaliphiles and their products have got several applications such as in detergent, paper and pulp, textile, and leather industries. These applications and human curiosity of understanding nature led to isolation and studies of a wide range of alkaliphiles from various alkaline environments. In recent years, these studies are contributing to expand the application horizon of alkaliphiles to new territories and technologies. This volume on “Alkaliphiles in Biotechnology” was thus intended to bring attention to these fascinating extremophiles and highlight their immense biotechnological potential. Alkaline habitats are widely distributed in the biosphere. However, the genesis andactualchemicalcompositionoftheseenvironmentsarequitedifferent,afactor which influences the diversity of the community inhabiting the alkaline environ- ments.Thisvariationalsoaffectsthesuccessofisolatingalkaliphilesfromdifferent habitats.Moreover,likeotherextremophiles,isolationofalkaliphilesisnotefficient and only a very tiny fraction of alkaliphiles community are culturable. However, there is some progress made on the isolation and cultivation of this group of organisms. These advances in isolation and cultivation of alkaliphiles are included inthisvolume. Theemergenceofmoleculartechniquessuchascloningandheterologousexpres- sion,metagenomics,systembiology,metabolicengineering,andsystemsmetabolic engineering which integrates genetic engineering, synthetic biology, and systems biology approaches has started to contribute to the advancement of biotechnology. v vi Preface To utilize alkaliphiles efficiently, it is necessary to implement these molecular techniques and hence, it is desirable to understand their genetic makeup. Thus, there have been studies on the genomes of alkaliphiles for which a chapter is dedicatedinthisvolume. It is not only the applications of alkaliphiles that are enthralling, but the mech- anism of how these organisms adapted to the extreme pH habitats which are often lethaltootherlifeformsisalsointriguing.Therearesomeprogressesthatdepictpart of the alkaline adaptations. In this volume, some of the most important adaptation mechanismsarehighlightedandnewinsightswhichmightstimulatefurtherstudies areincluded. Atleastsomeoftheuniqueproductsandphysiologiesofalkaliphileswhichareof greatimportancetobiotechnologyemanatefromthehighpHadaptationmechanism. Forinstance,alkaliphilesproduceorganicacidsthatalleviatethehighpHstress,an ability that can be tapped for commercial production of organic acids. Similarly, alkaliphilesadaptationresultedinnovelsiderophores,exopolysaccharides,caroten- oids,biosurfactants,andmanyothermetaboliteswithimmenseapplicationpotential. Oneofthemostimportantandwell-studiedalkaliphilesadaptations,whichattracted a great deal of interest, is their alkaline-active and -stable enzymes. These unique enzymes are suitable for high pH biocatalysis and are implemented in several processes. Among alkaline active enzymes, those enzymes which modify starch, hemicellulose,andproteinarerelativelythemoststudiedandaccountforthegreater portion of the current alkaline active enzymes’ applications. The real and potential applicationsoftheseextraordinaryextracellularenzymesofalkaliphilesarecovered inthevolume. Inadditiontotheirproducts,alkaliphileswholecellshavegotapplicationssuch as in food and feed supplementation, waste treatment, biofuel, and other chemical production. Another newly emerging application of alkaliphiles is in the construc- tion industry where alkaliphiles are winning growing attention to treat concrete. Alkaliphiles have been tried to make concrete surface coatings, repair concrete cracks, and engineer self-healing concretes. Moreover, it can also serve in many other related applications such as in bioclogging, bioaggregation, and as concrete admixture. This book highlights the major advances of alkaliphiles applications in theconstructionindustry. Althoughalkaliphilesareoneoftheoldestorganisms,itisamongthenewcomers into the circle of biotechnology. Hence, an active researchis ongoing and some of thesestudiestrickleinnewfascinatingapplicationsanddiscoveries.Theemergence of new techniques, the desire for new products, the growing demand for organic productsandbiotechnologicalprocesses,andtheprovenversatilityofalkaliphilesin serving different sectors of biotechnology may indicate the possible expansion of alkaliphilesbiotechnologicalutilizationinyearstocome. As editors of this volume, we have acquired a lot from the chapters written by scientists who are specialists in the field and hence, we would like to extend our heartfeltgratitudetoalltheauthorsfortheiroutstandingcontributionandeffortsfor Preface vii makingthisbookaninteresting,importantsourceofinformationonbiotechnology ofalkaliphiles.Inthemeantime,wewouldalsoliketotaketheopportunitytothank theserieseditorofAdvancesinBiochemicalEngineering/Biotechnologyforbeliev- inginthesignificanceofalkaliphilesinbiotechnologyandtheinspirationtoeditthis volume. Billeberga,Sweden GashawMamo Lund,Sweden BoMattiasson Contents Alkaliphiles:TheVersatileToolsinBiotechnology. . . . . . . . . . . . . . . . . 1 GashawMamoandBoMattiasson IsolationandCultivationofAlkaliphiles. . . . . . . . . . . . . . . . . . . . . . . . . 53 VadimV.Kevbrin ChallengesandAdaptationsofLifeinAlkalineHabitats. . . . . . . . . . . . 85 GashawMamo GenomicsofAlkaliphiles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135 PedroH.LebreandDonA.Cowan MetabolitesProducedbyAlkaliphileswithPotentialBiotechnological Applications. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157 ElviraKhalikova,SusanneSomersalo,andTimoKorpela AlkaliphilicEnzymesandTheirApplicationinNovelLeather ProcessingTechnologyforNext-GenerationTanneries. . . . . . . . . . . . . . 195 WycliffeC.WanyonyiandFrancisJ.Mulaa Starch-ModifyingEnzymes. . . . .. . . . . . . . . . . . . .. . . . . . . . . . . . . .. . 221 SuhailaOmarHashim AlkalineActiveHemicellulases. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 245 GashawMamo Alkaliphiles:TheEmergingBiologicalToolsEnhancingConcrete Durability. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 293 GashawMamoandBoMattiasson Index. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 343 ix AdvBiochemEngBiotechnol(2020)172:1–52 DOI:10.1007/10_2020_126 ©SpringerNatureSwitzerlandAG2020 Publishedonline:28April2020 Alkaliphiles: The Versatile Tools in Biotechnology GashawMamoandBoMattiasson Contents 1 Introduction................................................................................... 4 2 Extremophiles................................................................................. 5 3 Alkaliphiles................................................................................... 9 4 AlkaliphilesinBiotechnology................................................................ 10 4.1 AlkalineActiveEnzymes.............................................................. 10 4.2 PharmaceuticallyImportantMetabolites............................................... 21 4.3 CarotenoidsofAlkaliphiles............................................................ 23 4.4 SiderophoresofAlkaliphiles........................................................... 24 4.5 BiosurfactantProductionbyAlkaliphiles.............................................. 25 4.6 ExopolysaccharidesofAlkaliphiles.................................................... 25 4.7 AlkaliphilesinBiofuelProduction..................................................... 26 4.8 OrganicAcids.......................................................................... 28 4.9 AlkaliphileBiomassasFoodandFeedSupplement.................................. 29 4.10 AlkaliphilesinBioelectricity:MicrobialFuelCell.................................... 30 4.11 AlkaliphilesinBioconstruction........................................................ 32 4.12 AlkaliphilesinMiningandOilDrilling............................................... 32 4.13 AlkaliphilesinEnvironmentalBiotechnology......................................... 34 5 ConcludingRemarks.......................................................................... 36 References......................................................................................... 37 Abstract The extreme environments within the biosphere are inhabited by organ- ismsknownasextremophiles.Lately,theseorganismsareattractingagreatdealof interest from researchers and industrialists. The motive behind this attraction is mainly related to the desire for new and efficient products of biotechnological G.Mamo(*) IndienzAB,Billeberga,Sweden e-mail:[email protected] B.Mattiasson DepartmentofBiotechnology,LundUniversity,Lund,Sweden e-mail:[email protected]