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Growth and Defence in Plants: Resource Allocation at Multiple Scales PDF

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Ecological Studies, Vol. 220 Analysis and Synthesis Edited by M.M. Caldwell, Washington, USA G. Heldmaier, Marburg, Germany R.B. Jackson, Durham, USA O.L. Lange, Wu¨rzburg, Germany H.A. Mooney, Stanford, USA E.-D. Schulze, Jena, Germany U. Sommer, Kiel, Germany Forfurthervolumesseelistattheendofthebookand: http://www.springer.com/series/86 . Rainer Matyssek (cid:129) Hans Schnyder (cid:129) Wolfgang Oßwald (cid:129) Dieter Ernst (cid:129) Jean Charles Munch (cid:129) Hans Pretzsch Editors Growth and Defence in Plants Resource Allocation at Multiple Scales Editors ProfessorDr.RainerMatyssek ProfessorDr.HansSchnyder TechnischeUniversit€atM€unchen TechnischeUniversit€atM€unchen ChairofEcophysiologyofPlants Lehrstuhlf€urGr€unlandlehre Freising,Germany Freising,Germany ProfessorDr.WolfgangOßwald Dr.Dieter Ernst TechnischeUniversit€atM€unchen HelmholtzZentrumM€unchen PhytopathologyofWoodyPlants InstituteofBiochemicalPlantBiology Freising,Germany Neuherberg,Germany ProfessorDr.JeanCharlesMunch ProfessorDr.HansPretzsch HelmholtzZentrumM€unchen TechnischeUniversit€atM€unchen InstituteofSoilEcology ChairofForestGrowthandYieldScience Neuherberg,Germany Freising,Germany ISSN0070-8356 ISBN978-3-642-30644-0 ISBN978-3-642-30645-7(eBook) DOI10.1007/978-3-642-30645-7 SpringerHeidelbergNewYorkDordrechtLondon LibraryofCongressControlNumber:2012948418 #Springer-VerlagBerlinHeidelberg2012 Thisworkissubjecttocopyright.AllrightsarereservedbythePublisher,whetherthewholeorpartof thematerialisconcerned,specificallytherightsoftranslation,reprinting,reuseofillustrations,recita- tion, broadcasting, reproduction on microfilms or in any other physical way, and transmission or 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. Theuseofgeneraldescriptivenames,registerednames,trademarks,servicemarks,etc.inthispublica- tiondoesnotimply,evenintheabsenceofaspecificstatement,thatsuchnamesareexemptfromthe relevantprotectivelawsandregulationsandthereforefreeforgeneraluse. 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) To our late colleague and friend, Prof. Dr. Heinrich Sandermann jr. ({ August 18, 2009), who was one of the initiators and members of the interdisciplinary research program SFB 607 “Growth and Parasite Defence in Plants – Competition for Resources in Economic Plants from Agronomy and Forestry”, giving rise to this book project, but to whom it was not granted to see its completion. . Prologue A New View on Systems Biology: Information, Knowledge, Understanding Cost/Benefit Scenarios and the Dilemma of Growth and Defence Cost/benefit relations determine the functions of plants in natural ecosystems as well as anthropogenically formed agro- and forestry ecosystems. Under the influ- ence of abiotic and biotic cues, they regulate the dynamics in the realm of the opposingforcesofgrowthanddefence(Chap.1).Inthecompetitionforresources thedilemmahasthecostsofproductivityontheonehandandofsurvivalofstress ontheotherhand. Talking of a dilemma we must ask the question if there are in fact options of ways out of it. Evidently, like all living organisms plants are also open systems subjecttonon-lineardynamicsinspaceandtime.Thisimpliesthatspatio-temporal developmentsunderaplethoraofexternalcuesandtheopposingforcesofgrowth and defence inevitably lead to branching points where alternatives or options are given,butwherechancemayeventuallyalsoleadthewayintodeterministicchaos. Thelatterisaninherentprobleminevaluationsofthedegreesofpossiblepredict- abilityinallsystemswithnon-lineardynamics(Schuster1995).Optionsinspatio- temporal responses of plants under the dilemma of growth and defence and potential predictability are the central themes underlying the hypotheses and conceptsevaluatedinthisbook. Scenariosaredominatedby (cid:129) differentlifeformsandtypesofhabitats,viz.grasslandandforest, (cid:129) competition (Chaps. 17 and 18), abiotic stress (ozone, elevated CO , nitrogen 2 nutrition),pathogens(Chaps.3and5),herbivores(Chap.4),symbionts(Chaps. 5and10), (cid:129) ontogeneticdevelopment(Chap.11), vii viii Prologue (cid:129) types of resources, e.g. originating from the soil (Chaps. 9 and 10), from aboveground components, from the atmosphere, of the solar radiation (Chap.8),ofspaceitselfasaresource(Chap.12,GramsandL€uttge2010), (cid:129) allocation, partitioning and consumption of resources in allometry relations (Chaps.6,7,9,11–18). These scenarios are elements of the formation of integrative networks within various scalar levels which are also connected between each other over both ascending and descending scales and thus can form supra-networks. This determines a new view on systems biology in an ecological context (Chap. 19), which must advance from information via knowledge towards understanding (Liessmann2008), i.e. frompurely descriptivecompilation ofdata via correlative interpretation of processes towards reflection and causal assessment. Combining theapproachesofexperiment,theoryandmodellingdevelopspredictivepoweron thebasisofunderstandingobservedpastevents. An Architectural Metaphor In a metaphoric way we may visualise the architecture of the Sonderforschungsbereich (SFB) “Wachstum und Parsitenabwehr – Wettbewerb um Ressourcen in Nutzpflanzen aus Land- und Forstwirtschaft”, from whose work the present volume is emerging, like the knots and edges of networks. We mayconsidertheexperimentalapproachesontwobasicallydifferentlevelsastwo columnsortowers(knots)whichbylongwings(edges)areconnectedtotheoryand modelling(knot). I cannot imagine a better way of depicting this than choosing the Castle of Hirschberg above the city of Beilngries in the valley of the Altm€uhl in Central Bavaria. For more than a decade the SFB had regular internal meetings there for brainstormingandstrategyplanning.Whenwearriveatthecastlewepassthetwo towersandweseethetwowingsofthebuildingguidingourviewinaremarkably suggestivewaytowardstheentrance(Fig.1).Thearchitectmanagedthisbyhaving thetwowingsinanangleandmuchwiderapartfromeachotheratthestartthanat theendwiththeentrance.Letusconsiderthetwotowersasthetwosectionsofthe SFB. One of the towers (one knot) of the SFB or its section A (Chaps. 2–5) is linkingmolecularbiologywithbiochemistryandphysiology(Chap.2)and,thus,is developingmolecularecologyof (cid:129) geneinductionandtranscription, (cid:129) proteinsandenzymeactivities, (cid:129) primaryandsecondarymetabolism, (cid:129) signallingbyphytohormonesandmetabolites. The other tower (knot) of the SFB, its section B (Chaps. 6–14), is developing physiologicalecologyattheorganismic,standandecosystemlevels,with Prologue ix Fig.1 HirschbergCastleofthebishopricofEichst€attabovethecityofBeilngriesinthevalleyof theAltm€uhl(photographs:K.-H.H€aberle) (cid:129) primaryproduction,allocation,partitioningandallometry, (cid:129) occupationofspaceaboveandbelowground, (cid:129) theuseofreservesunderstressofresourceavailability,ozonepollution,elevated atmosphericCO ,pathogenattack. 2 x Prologue With the two wings the two towers are connected to the entrance, section C (Chaps. 15–19) of the SFB. What is this an entrance to? Transduction of the informationandknowledgesensuLiessmann(2008)fromthetwosections,towers orknotsAandBviathewingsoredgesintosectionorknotCentersthewealthof observations (information) and interpretations (knowledge) into understanding sensuLiessmann(2008)asprobedandadvancedby (cid:129) statisticalmodelling(Chap.16), (cid:129) mechanisticmodelling(Chaps.14,15,17and18), (cid:129) integrationviaabstraction. Thus, metaphorically we can see that, as in the most compelling way in which the two wings of the castle guide our anticipation to arrive towards the entrance, sectionsAandBoftheSFBfunnelourattentiontowardstheintegrationbytheory andmodellingofsectionC. What Is a “System” in Biology? TheextremeviewisGaiaconsideringthewholebiosphereasonesinglesystemor onemega-organism(Lovelock1979,2009). Inthe life sciences theterm “systemsbiology” originated fromdescribingvast accumulations of data in “genomics”, and in a noteworthy inflationary fashion it now covers all kinds of “-omics”, such as transcriptomics, proteomics, metabolomics and channelomics. The present book presents examples of such databases (Chaps. 2–5). Remarkably comprehensive sets of data on mineral macro- and micro-nutrients in plants have also been obtained (Chap. 9). In the SFB this has been called “mineralomics”. The term “ionomics“ is somewhat of a misnomerintheliterature(Saltetal.2008)aslongasitrefersonlytoinorganicions anddoesnotcompriseontheonehandthewealthoforganicionsimportantinthe functioning of organisms and realise on the other hand that many minerals in organismsoccurandfunctioninnon-ionicboundorchelatedstates.Intheveinof a certain omics-inflation we may also allude to the necessity of considering structuralorfunctionalcomplements,i.e.“anatomics”or“functionomics”. However, are such data bases “systems”? They are really nothing more than primary information. They constitute a necessary basis but themselves they have nothingtodowithknowledgewhichmustbeadvancedfromtheinformation. Anotherwayoflookingfromamolecularpointofviewtosystemsistoconsider specifickeyfunctionswhichareoftenexpressedbymanydifferentisogenescoding differentisoenzymes.Interestinglyabioticstressduetoozoneandbioticstressdue to pathogen attack may elicit very similar reactions at the molecular level, viz. transcription (Chap. 2). The translational and post-translational regulation and modulationofisoenzymeactivitiesaswellastheirlocalisationandcompartmenta- tioncancharacterisethespatio-temporalfunctioningofthesystemsofwholeplants or even higher integrated consortia, such as host/parasite or host/symbiont

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