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Physical and Physiological Forest Ecology Pertti Hari • Kari Helio¨vaara (cid:129) Liisa Kulmala Editors Physical and Physiological Forest Ecology 123 http://avaxhome.ws/blogs/ChrisRedfield Editors PerttiHari KariHelio¨vaara DepartmentofForestSciences DepartmentofForestSciences UniversityofHelsinki UniversityofHelsinki Helsinki Helsinki Finland Finland LiisaKulmala DepartmentofForestSciences UniversityofHelsinki Helsinki Finland ISBN978-94-007-5602-1 ISBN978-94-007-5603-8(eBook) DOI10.1007/978-94-007-5603-8 SpringerDordrechtHeidelbergNewYorkLondon LibraryofCongressControlNumber:2012951122 ©SpringerScience+BusinessMediaDordrecht2013 Thisworkissubjecttocopyright.AllrightsarereservedbythePublisher,whetherthewholeorpartof thematerialisconcerned,specificallytherightsoftranslation,reprinting,reuseofillustrations,recitation, broadcasting,reproductiononmicrofilmsorinanyotherphysicalway,andtransmissionorinformation storageandretrieval,electronicadaptation,computersoftware,orbysimilarordissimilarmethodology nowknownorhereafterdeveloped.Exemptedfromthislegalreservationarebriefexcerptsinconnection with reviews or scholarly analysis or material supplied specifically for the purpose of being entered and executed on a computer system, for exclusive use by the purchaser of the work. Duplication of this publication or parts thereof is permitted only under the provisions of the Copyright Law of the Publisher’slocation,initscurrentversion,andpermissionforusemustalwaysbeobtainedfromSpringer. PermissionsforusemaybeobtainedthroughRightsLinkattheCopyrightClearanceCenter.Violations areliabletoprosecutionundertherespectiveCopyrightLaw. Theuseofgeneraldescriptivenames,registerednames,trademarks,servicemarks,etc.inthispublication doesnotimply,evenintheabsenceofaspecificstatement,thatsuchnamesareexemptfromtherelevant protectivelawsandregulationsandthereforefreeforgeneraluse. 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) Preface Photosynthesis is the source of energy for nearly all life on Earth and the source of the carbon in all the organic compounds within the bodies of organisms. Photosynthesisisthebiochemicalreactionthroughwhichallgreenplants,including trees,liveandgrow.We startedfield measurementsofphotosynthesisin Hyytia¨la¨, the Forestry Field Station at the Universityof Helsinki, in the year 1972,and this wasthebeginningofastillcontinuingworkintheforestsaroundthestation.Wehad thevisionthatphysicalandphysiologicalknowledgeinadditiontothequantitative methods, especially field measurements and dynamic modelling, should have an importantrole in forest ecology.We introducedinto our vision the importantrole ofmassandenergyfluxeswhenwestudiedtheChernobylfalloutinthesummerof 1986.TheconstructionoftheSMEARIImeasuringstationclarifiedourvision,and weincludedthefluxesinthesoilandbetweenthesoilandtreesintoourapproach. Some members in our team have had good knowledge in tree physiology, and they introducedthe physiologicalaspect into the research.We have been lucky to workwith physicistswho are deeplyinterestedin the role of physicalphenomena inforests.Dr.TaistoRaunemaa(1939–2006)startedcommonprojectswithus,and thephysicistsbecomeanaturalcomponentinourresearchteam.Theco-operation betweentheAcademyofFinlandandSovietAcademyofSciencesenabledcommon research in Finland, Soviet Karelia and Estonia. Physicists, Academician Juhan Ross(1925–2002),Dr.LeoKaipiainen(1932–2004)andDr.TiitNilsonhavebeen thekeypersonsintheteam. The co-operation with physicists was intensified when Dr. Markku Kulmala began to lead the team of young aerosol physics at the Department of Physics, Helsinki University. He was the other principalplanner of the measuring systems SMEAR I and II. In addition, he is interested in the role of boreal forest in the formation and growth of aerosols. Dr. Timo Vesala introduced the important role ofmeasurementsofecosystemfluxeswiththeEddyCovariancemethod.Heisalso interestedintheroleofnitrogeninforestecosystems. Thefeedbackfromourmeasurementshasbeenclearandpositive.Thedeveloped modelstructureshavebeenabletoexplainwellthebehaviouroftreesandground vegetation in field conditions. We have published our results in international v vi Preface literature, although some of the most relevant papers have remained unpublished in scientific journals. During the last 20 years, we have actively participated into Europeanresearchco-operation. The Academyof FinlandandHelsinkiUniversityhave beenthe main financial supportersduringthewholelongperiod. Ourfirstreader,Dr.JohnGracefromEdinburgh,hasmadevaluablecommentson ourmanuscriptandweexpressourgratitudetohim.WeacknowledgeDr.Kourosh KabiriKoupaei,forhiscontributiontotheeditingofthebook. Ourbookisthesummaryofourworkduring40yearsatthepresentDepartment of Forest Sciences, previously Department of Forest Ecology and Department of Silvicultureat HelsinkiUniversity.Our visionstressing the importanceof physics andphysiologicalknowledge,quantitativemethods,materialandenergyfluxesand fieldworkingintheforestshashadthecentralroleintheplanningoftheresearch. Helsinki Pertti(Pepe)Hari,D.Sc.(Agr.&For.),Prof. 6thDecember2012 KariHelio¨vaara,D.Sc.(Agr.&For.),Prof. LiisaKulmala,D.Sc.(Agr.&For.) Contents 1 IntroductiontoPhysical,PhysiologicalandCausalForestEcology .. 1 PerttiHari,LiisaKulmala,andMikkoHavimo 2 TheApproachtoConstructandTesttheTheoryofForestEcology.. 7 PerttiHari 3 EnvironmentalFactors..................................................... 27 U¨llarRannik, SamuliLauniainen, JukkaPumpanen, LiisaKulmala, PasiKolari, TimoVesala, JanneF.J.Korhonen,andPerttiHari 4 ProcessesinLivingStructures ............................................ 43 JaanaBa¨ck,EeroNikinmaa,LiisaKulmala,AskoSimojoki, TuomoKalliokoski, PerttiHari, RistoHa¨kkinen, TapioLinkosalo, HeikkiHa¨nninen, EijaJuurola, PasiKolari,KouroshKabiriKoupaei,AlbertPorcar-Castell, BenãtOlascoagaGracia,JussiHeinonsalo,SariTimonen, KariHelio¨vaara, MaaritRaivonen, JohannaJoensuu, MariPihlatie, JukkaPumpanen, JukkaKurola, MirjaSalkinoja-Salonen,andMikaKa¨hko¨nen 5 FluxesofCarbon,WaterandNutrients.................................. 225 TeemuHo¨ltta¨,PerttiHari,KariHelio¨vaara,EeroNikinmaa, JukkaPumpanen, TimoVesala, PasiKolari, SamuliLauniainen, U¨llarRannik, LiisaKulmala, KouroshKabiriKoupaei,MinnaPulkkinen,MariPihlatie, JanneF.J.Korhonen,AskoSimojoki,Antti-JussiKieloaho, JaanaBa¨ck,andMarkkuKulmala 6 StructuralRegularitiesinTrees........................................... 329 PerttiHari, MikkoHavimo, JuhoAalto, PauliinaSchiestl-Aalto, EeroNikinmaa, AnnaLintunen, TuomoKalliokoski,Helja¨-SiskoHelmisaari,andIngeStupak vii viii Contents 7 Dynamicsof CarbonandNitrogenFluxesandPools inForestEcosystem ........................................................ 349 PerttiHari, MikkoHavimo, KouroshKabiriKoupaei, KalevJo¨giste, AhtoKangur, MirjaSalkinoja-Salonen, TuomasAakala, JuhoAalto, PauliinaSchiestl-Aalto, JariLiski,andEeroNikinmaa 8 HowtoUtilisetheKnowledgeofCausalResponses?................... 397 PerttiHari, MikkoHavimo, Helja¨-SiskoHelmisaari, LiisaKulmala,EeroNikinmaa,TimoVesala,JouniRaïsa¨nen, TuukkaPeta¨ja¨, ErkkiSiivola, HeikkiTuomenvirta, JaanaBa¨ck,JohnGrace,FedericoMagnani,TwanvanNoije, JukkaPumpanen, DavidStevenson, MarkkuKulmala, SampoSmolander,IlonaRiipinen,andMiikkadalMaso 9 StationforMeasuringEcosystem-AtmosphereRelations:SMEAR.. 471 PerttiHari, EeroNikinmaa, ToivoPohja, ErkkiSiivola, JaanaBa¨ck,TimoVesala,andMarkkuKulmala 10 ThePhysicalandPhysiologicalTheoryofForestEcology andItsEvaluation.......................................................... 489 PerttiHari Appendices........................................................................ 505 Glossary........................................................................... 521 Index............................................................................... 531 Contributors TuomasAakala DepartmentofForestSciences,UniversityofHelsinki,Helsinki, Finland Juho Aalto Hyytia¨la¨ Forestry Field Station, University of Helsinki, Helsinki, Finland Jaana Ba¨ck Department of Forest Sciences, University of Helsinki, Helsinki, Finland John Grace Institute for Atmospheric and EnvironmentalScience, University of Edinburgh,Edinburgh,UK BenãtOlascoagaGracia DepartmentofForestSciences,UniversityofHelsinki, Helsinki,Finland RistoHa¨kkinen FinnishForestResearchInstitute,Vantaa,Finland Heikki Ha¨nninen Department of Biosciences, University of Helsinki, Helsinki, Finland Pertti Hari Department of Forest Sciences, University of Helsinki, Helsinki, Finland Mikko Havimo Departmentof ForestSciences,UniversityofHelsinki,Helsinki, Finland Jussi Heinonsalo Department of Forest Sciences, Department of Physics and DepartmentofFoodandEnvironmentalSciences,UniversityofHelsinki,Helsinki, Finland KariHelio¨vaara DepartmentofForestSciences,UniversityofHelsinki,Helsinki, Finland Helja¨-Sisko Helmisaari Department of Forest Sciences, University of Helsinki, Helsinki,Finland ix