Anatomical Adaptations of Halophytes: A Review of Classic Literature and Recent Findings PDF
Preview Anatomical Adaptations of Halophytes: A Review of Classic Literature and Recent Findings
Marius-Nicușor Grigore Constantin Toma Anatomical Adaptations of Halophytes A Review of Classic Literature and Recent Findings Anatomical Adaptations of Halophytes Marius-Nicușor Grigore (cid:129) Constantin Toma Anatomical Adaptations of Halophytes A Review of Classic Literature and Recent Findings Marius-NicușorGrigore ConstantinToma FacultyofBiology FacultyofBiology AlexandruIoanCuzaUniversity PlantAnatomy&EcologyLaboratory Iasi,Romania AlexandruIoanCuzaUniversity Iasi,Romania ISBN978-3-319-66479-8 ISBN978-3-319-66480-4 (eBook) DOI10.1007/978-3-319-66480-4 LibraryofCongressControlNumber:2017951218 ©SpringerInternationalPublishingAG2017 Thisworkissubjecttocopyright.AllrightsarereservedbythePublisher,whetherthewholeorpartof the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilarmethodologynowknownorhereafterdeveloped. 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Printedonacid-freepaper ThisSpringerimprintispublishedbySpringerNature TheregisteredcompanyisSpringerInternationalPublishingAG Theregisteredcompanyaddressis:Gewerbestrasse11,6330Cham,Switzerland Foreword When my colleague and good friend, Marius-Nicus¸or Grigore, asked me to write the Foreword to this book, “Anatomical adaptations of halophytes,” I was flattered… but I thought I was not the most appropriate person to do so. I have worked for over 30 years on various aspects of plant development and plant responses tostress and the last 15 more specifically on salt tolerance mechanisms in various plant species, including several halophytes. However, I am a plant biochemist and molecular biologist and I do not consider myself an expert in plant anatomy … well, let’s be honest: myknowledge of plant anatomy is practi- callyzero.So,IfeltIhadnothingtosaytothepotentialreadersofthebook,whoI assumedshouldbemostlybotanistsandplantanatomists.Whilelookingforapolite waytodeclinetheinvitation,Iglancedatthesepagesthatarenowinyourhands, dear reader—or, nowadays, more probably on your computer screen—and I completely changed my mind. The book fascinated me, and I realized that it could be just as interesting for any scientist working with this amazing group of salt-tolerantplants,regardlessoftheirparticularfieldofresearch.Whatfollowsare, therefore, some brief comments from a nonspecialist, addressed to those re- searchers with various backgrounds and expertise, about a much specialized— and very special—book, which I believe should be present on the shelves (or harddisks)ofanylaboratoryinvolvedinthestudyofhalophytes. Iamnotgoingindetailintothecontentsofthebook,itsscope,ortheauthors’ reasons to write it; I will not even mention the (multiple) definitions of “halo- phytes,”apartfromthefactthattheyareplantsabletosurviveandreproduceunder salinityconditionsthatwillkillthevastmajorityofplantspecies.Thereaderscan lookthemselvesatthepreface,thetableofcontents,ortheintroductionofthebook. I would like, instead, to frame the study of halophytes in a wider context: as the mostappropriatemodelstoinvestigatethemechanismsofsalttoleranceinplants— which should be obvious, despite the opinion of some hardcore plant molecular biologists (no, this book does not deal with Arabidopsis thaliana). The study of thesemechanismsisatpresentaveryactivefieldinplantbiologyresearch,because ofitsunquestionableacademicinterestbutalsoduetoitspracticalimplicationsin v vi Foreword agriculture. Soil salinity is—together with drought—the most important environ- mental stress factor responsiblefor the reductionofcrop yields worldwide.In the context of global climate change, increasing scarcity of good-quality water for irrigation, and population growth, the biotechnological improvement of the salt toleranceofourcropshasbecomeanurgentneedforthefutureofagricultureand food production, especially in arid and semiarid regions; this, in turn, requires a deep knowledge of the mechanisms used by plants to counteract the deleterious effectsofhighsoilsalinity.Halophytescannotonlycontributebasicknowledgebut also provide biotechnological tools—salt tolerance genes and salt-induced pro- moters—which could be eventually used for the genetic improvement of salt tolerance of our crops. Some of them could even be domesticated to generate new crops for food, feed, fiber, biofuel production, or other industrial uses. They wouldbethebasisofa“salineagriculture,”couldbegrowninsalinisedsoils and irrigated with brackish or sea water, avoiding in this way competition with our conventional crops for scarce resources: fertile farmland and freshwater for irrigation. Coming back to the book itself, there are several aspects that, in my opinion, makeitspecialandshouldbehighlighted.First,thebookshouldappealtoawide readership;duetotheirinterestinbasicresearchandpossiblepracticalapplications —asmentionedabove—halophytesarebeingstudiedfrommanydifferentpointsof view by researchers with quite different scientific backgrounds: ecologists, bota- nists, agronomists, plant physiologists, biochemists, and molecular biologists, manyofthem(especiallythelatter)withlittleknowledge,ifany,ofplantanatomy. The information contained in this book is organized basically around specific anatomic adaptations, which represent structural strategies of halophytes to with- standhighsoilsalinity(succulence,saltglands,Kranzanatomy,etc.),andnotbased ontaxonomicclassifications,asonecouldexpectforabookofthiskind.Ifindthis approach very convenient and accessible, particularly for nonspecialists. Those scientistsinvestigatingthephysiological,biochemical,andmolecularmechanisms activated in plants, in general, and in halophytes, in particular, in response to salt stress—Iknowitfrommyownexperience—oftenneglectconstitutivemechanisms ofdefensebasedonstructuraladaptationsoftheirinvestigatedspecies.Thisbook canhelpallofustolookatourownresearchwithawiderperspective.Ialsofind veryinterestingandintellectuallyrewarding(thisisobviouslyapersonalopinion) thehistoricalapproachunderlyingtheentirebook’scontentsandorganization,with numerous references to the work of classical botanists—which in other scientific fieldswouldbeconsidered“ancient”butinplantanatomywillneverbeoutdated.I would like to point out, especially, the high quality of the (also “classical”) ink drawings in many of the figures; probably no “modern” micrograph can show anatomicstructureswithsuchclarityanddetail. Inatimeof“omics”technologies,fullgenomesequences,DNA“barcodes,”and Appsthatcansupposedlyidentifyanyplantspeciesfromapicturetakenwithyour mobilephone,itisgoodtolookbackandacknowledgetheenormouscontribution Foreword vii ofclassicalbotaniststoourpresentknowledgeinthisfield.Itisaswellrefreshingto look at the beauty of nature, also reflected in the anatomic adaptations to salinity shownhere.Thisis,basically,whatthisbookisabout. InstituteofPlantMolecularandCellular OscarVicente Biology(IBMCP,UPV-CSIC) UniversitatPolite`cnicadeVale`ncia Valencia,Spain Preface Succulence,tracheoidioblasts(spiralcells),saltsecretion,Kranzanatomy,succes- sive cambia, and bulliform cells represent major anatomical adaptations found in halophytes;theyaretreatedinthismonographwithanemphasisonliteraturefrom the end of the nineteenth century and the beginning of the twentieth century. Of course, the list of adaptations is not exhaustive; other anatomical features in halophytes may be associated with saline environments. Naturally, the reviewed literaturehasnopretensiontobecomplete.Actually,partofrecentliteraturemight be regarded—at first glance—as being repetitive, at least when it refers to purely anatomicaldescriptiveapproachesinspeciesthathavealsobeeninvestigatedinthe past. This is an inherent risk in plant anatomy research, and the real challenge wouldbetofindtherightframeinwhichtointegratethiskindofapproach. Anotherimportantobservationinrelationtothisbookisthatwetried—asmuch as possible—to remain close to anatomy and not to go into details of associated issues emerging from many of the discussed adaptations. For instance, Kranz anatomy is related to C photosynthetic pathway, and this requires a very sophis- 4 ticatedapproachintermsofphysiology,biochemistry,andpaleo-ecologyofhalo- phytes.Inthisrespect,aluxuriantliteraturehasbeenproducedinthepast20years, includingeditedbooksandpublishedpapers—originalandreviews.Ofcourse,our intention didnotinterfere with these research directions.Thesameis truefor salt glands, where many results derived from ultramicroscopic observations or are focused on the ecophysiology of salt secretion. Again, we tried to remain at the borderofthisapproach. However,thesetwoexamplesclearlysuggestthattheanatomyofhalophytesisa refreshing field that opened many advanced research directions, such as plant ecophysiology,saltstressphysiology,andmolecularapproaches. Thechapterdedicatedtosalineenvironmentsisintendedonlytofamiliarizethe reader with several operational terms related to salt areas. This should not be regarded a soil science chapter, since we are aware that the terminology and taxonomy of salt-affected soils is very problematic and the strongest evidence is the multitude of classification systems existing worldwide. Therefore, in this ix x Preface chapter several concepts are discussed, since the chosen terms are largely used in thebotanicalliterature,especiallyinthatfromthepastdecades. Because of the specificity of this book, mentioned species were kept in the genuine nomenclatural state, with no intention to find actual synonyms. This would have been tricky, especially for species from Chenopodiaceae and Plumbaginaceae; in addition, consistent with the vision of this book, readers interested in the history and evolution of plant taxonomy should be invited to discover by themselves the exciting journey toward—likely—the most romantic andfruitfulperiodinthehistoryofbotany. Iasi,Romania Marius-NicușorGrigore ConstantinToma Acknowledgements Wehavetothankseveralpersons: Oscar Vicente (Instituto de Biolog´ıa Molecular y Celular de Plantas, UPV-CSIC) was willing to write a Foreword for our book. In the past, he helped me very much to integrate in his research group from UPV, giving me the possi- bility to work and extend my research interests in halophytes. Our collaboration was and still is very fruitful, and I’m deeply indebted to him for all the support during the last 7 years. Together with Monica Boscaiu (Instituto Agroforestal Mediterra´neo, UPV), we managed to create a strong research interest network betweenRomaniaandSpain. UniversidadPolite´cnicadeValencia(UPV)hostedmethreetimesatInstitutode Biolog´ıaMolecularyCelulardePlantas,fordevelopingscientificmissionsrelated to the study of halophytes. Two stays were funded by COST Action FA0901: “Putting Halophytes to Work—From Genes to Ecosystems,” in a frame of Short- Term Scientific Missions. One stay was funded by Universidad Polite´cnica de ValenciathatsupportedmeintheanatomicalstudyofSpanishhalophytes.During thesestays,Ihadtheexcellentopportunitytocollectalargenumberofhalophytes, which were anatomically investigated. Part of the obtained results is included in thisbook. COSTActionFA0901:“PuttingHalophytestoWork—FromGenestoEcosys- tems”alsofundedmeforattendingameetinginTorun(Poland),whenIcollected twohalophytespeciesfromsalt-affectedareas,duringafieldtrip.Onthisoccasion, IthankAgnieszkaPiernik(NicolausCopernicusUniversity,Torun)andAlexander Fehe´r(SlovakUniversityofAgriculture,Nitra,Slovakia)forhelpingmeidentify- ingseveraltaxainthefield. Surveyandanalysisofoldbotanicalliteraturerepresentapivotalaspectforthis book. Not all the bibliographic references were available to us. A part of old papers—thatwasnotonourdisposal—hasbeenprovidedbyfewpersons.Roberta Gasparrifromthe DepartmentofAgricultural, FoodandEnvironmentalSciences, MarchePolytechnicUniversity(Ancona,Italy),sentusLicopoli’spapers. xi
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