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Wood Structure and Environment

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Dendrochronology and wood anatomy developed for decades as two independent scientific fields. It was only in the last decade that it was made clear that the dimension of time is the fourth dimension for both sciences and that it was demonstrated that wood anatomy and dendrochronology are perfect partners.


The main aim of this book is to show the hidden ecological richness in stems and roots from trees, shrubs and herbs. It should encourage researchers to consider the anatomic microcosm of wood plants and use it as a retrospective source of information, solving problems related to ecophysiology, competition, site conditions, population biology, earth science, wood quality and even human history.

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Contents
1
2
3
Introductîon
Preparatîon of Wood and Herb Samples for Mîcroscopîc Analysîs 2.1 Labeîng, Transpor and Sorage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 2.2 Sampe Preparaîon or Mîcroome Secîonîng . . . . . . . . . . . . . . . . . . 3 2.3 Sampe Preparaîon or Epîscopîc Mîcroscopîc Examînaîon . . . . . . 3 2.4 Makîng hîn Secîons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 2.5 Saînîng o hîn Secîons or Permanen Sîdes . . . . . . . . . . . . . . . . . . . 5 2.6 Preparaîon o Impermanen Sîdes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 2.7 Mîcroscopîc Observaîon and Poograpy . . . . . . . . . . . . . . . . . . . . . . 5
Growth Zones and Stems în Relatîon to Genetîc Ecologîcal Factors he Xyem As an Indîcaor o Geneîc Idenîy . . . . . . . . . . . . . . . . . . . 7 Woody Pans Wîou Vesses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Woody Pans wî Vesses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Woody Pans wî Vesses and Incuded Poem . . . . . . . . . . . . . . . . 8 Inra-annua Anaomîca Varîaîons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Fa Ground Tîssue Ces wî hîck Ce Was . . . . . . . . . . . . . . . . . . 8 Recurren Cange o Dîferen Ce Types and Ce Sîzes în Decîduous Trees . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Perîodîc Sorage o Ce Compounds . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Ce-Type-Specîic Ce Wa hîckenîng and Lîgnîicaîon . . . . . . . 14 Inerannua Anaomîca Varîaîons (Tree-Rîng Boundarîes) . . . . . . 14 hîck-Waed, Fa Fîbers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Perîodîc Inra-annua Cange Beween Dîferen Ce Types and Sîzes în Decîduous Trees în Reaîon o Earywood and Laewood Boundarîes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 Perîodîc Deposîîon o Crysas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 Varîabe Ce Wa hîckenîng and Lîgnîicaîon . . . . . . . . . . . . . . . . . 19 Varîabe Geneîc Predîsposîîon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 Laera Varîaîons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 Dîsconînuous Grow and Varîabe Anaomîca Srucures . . . . . . 22 Wavy Tree-Rîng Paern, Dîspaced Rîngs and Irreguar Sem Cross Secîons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 Reaîonsîp Beween Tree-Rîng Paerns and Rays and Vesse Densîy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 Reaîonsîp Beween Tree-Rîng Paern and Canged Dîferenîaîon Processes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 Reaîonsîp Beween Tree-Rîng Paern and Pî Sape . . . . . . . . . 30 Reaîonsîp Beween Tree-Rîng Paern and Advenîîous and Laen Buds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30 Varîabe Axîa Fîber Arrangemen, Especîay Spîra Graîn . . . . . . . 30
3.1 3.1.1 3.1.2 3.1.3 3.2 3.2.1 3.2.2 3.2.3 3.2.4 3.3 3.3.1 3.3.2 3.3.3 3.3.4 3.3.5 3.5 3.5.1 3.5.2 3.5.2.1 3.5.2.2 3.5.2.3 3.5.2.4 3.5.3
VI
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Contents
3.6
3.6.1 3.6.1.1 3.6.1.2 3.6.1.3 3.6.1.4 3.6.1.5 3.6.1.6 3.6.1.7 3.6.2 3.6.3
Grow Zones în Perennîa Pans wî Very Sor or Long Sem Axes: Perennîa Herbaceous Pans, Dwar Srubs, Cîmbers and Lîanas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30 Pans wî Sor Sem Axes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 Exîsence o Grow Rîngs în Dwar Srubs and Herbaceous Pans în Cenra Europe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 he Anaomy o Grow Rîngs în Herbaceous Pans . . . . . . . . . . . . 32 Age Deermînaîon o Herbaceous Pans . . . . . . . . . . . . . . . . . . . . . . . 40 Reaîonsîp Beween Grow Rîngs în Herbaceous Pans and heîr Grow Form and Taxonomy . . . . . . . . . . . . . . . . . . . . . . . . 41 Reaîonsîp Beween Grow Rîngs în Herbaceous Pans and Sîe Condîîons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 Common Caracerîsîcs în e Grow Rîng Srucure o Herbaceous Pans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 Ouook . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 Pans wî Exremey Eongaed Sem Axes, în Parîcuar rom Temperae and Semîarîd Regîons . . . . . . . . . . . . . 42 Comparîson o Wood Anaomîca Srucures o Dwar Srubs and Lîana-Lîke Woody Pans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
Modîicatîon of the Tree-Rîng Anatomy Due to Ageîng and the Posîtîon Wîthîn the Tree 4.1 Onogenesîs o e Xyem în Roos and Soos . . . . . . . . . . . . . . . . . . 45 4.1.1 he Cener o e Roo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45 4.1.2 he Cener o e Sprou . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48 4.2 Onogeneîc and Pysîoogîca Ageîng o Wood . . . . . . . . . . . . . . . . . 49 4.2.1 Deinîîon o e Term Ageîng . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49 4.2.2 he Efecs o Envîronmena Facors on Ageîng . . . . . . . . . . . . . . . . 49 4.2.2.1 Varîabe Speed o Ageîng Wîîn e Tree Crown and e Roo Sysem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49 4.2.2.2 Speed o Ageîng Under Varîous Sîe Condîîons . . . . . . . . . . . . . . . . 52 4.2.2.3 Genoypîc Ageîng . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52 4.2.2.4 Ce-Specîic Ageîng în e Xyem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52 4.2.3 Anaomîca Dîferences Beween Juvenîe and Adu Wood . . . . . . 52 4.3 Brances and Twîgs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56 4.3.1 he Dendrocronoogîca Imporance o Brances and Twîgs . . . . 56 4.3.2 Bud-Scae Scars and Tree Rîngs în Long and Sor Soos Expressîng Crown Dynamîcs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56 4.3.2.1 Bud-Scae Scars . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56 4.3.2.2 Tree Rîngs în Long and Sor Soos în Four Dîferen Tree Specîes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57 4.3.3 he Eîmînaîon o Brances and Twîgs . . . . . . . . . . . . . . . . . . . . . . . . 60 4.3.3.1 Grow Pase Beore Dea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60 4.3.3.2 Comparmenaîzaîon o Dead Brances and Twîgs . . . . . . . . . . . . . 63 4.3.3.3 Acîve Seddîng o Sor Soos (Cadoposîs) . . . . . . . . . . . . . . . . . . 63 4.3.4 Soo Meamorpîsm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66 4.3.4.1 Sor Soos în Conîers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66 4.3.4.2 horns on Decîduous Woody Pans . . . . . . . . . . . . . . . . . . . . . . . . . . . 66 4.3.4.3 Sperobass . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66 4.3.4.4 Neede Traces . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66 4.3.4.5 Traces o Mae Inlorescences în Conîers . . . . . . . . . . . . . . . . . . . . . . . 69 4.3.4.6 Traces o Femae Cones în Conîers . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70 4.3.4.7 Traces o Fruî Saks în Decîduous Woody Pans . . . . . . . . . . . . . . . 70
5
4.4 4.4.1 4.4.2 4.4.3 4.4.4 4.4.5 4.4.6 4.4.6.1 4.4.6.2 4.4.6.3 4.4.6.4 4.4.7
5.1 5.2 5.2.1 5.2.2 5.2.3 5.2.4 5.3 5.3.1 5.3.2 5.3.3 5.3.4
Contents
VII
Roos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72 he Dendrocronoogîca Imporance o Roos . . . . . . . . . . . . . . . . . 72 Srucura Caracerîsîcs o Laen Roos . . . . . . . . . . . . . . . . . . . . . . 72 Roo Grow Subjec o Lîe Mecanîca Sress . . . . . . . . . . . . . . . . 72 Roo Grow Subjec o Mecanîca Sress (Excep Buress Roos) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72 Srucura Canges Caused by e Exposure o Roos . . . . . . . . . . . . 75 Srucura Canges Caused by e Burîa o Soos . . . . . . . . . . . . . . 79 Burîa o Sow-Growîng Seedîngs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79 Burîa o e Soo Base and o Creepîng Soos . . . . . . . . . . . . . . . . 79 Burîa o Conîer Brances Cose o e Ground . . . . . . . . . . . . . . . . 79 he Esabîsmen o Spruce Seedîngs în Spagnum Cusîons . . . 81 Sprou and Roo Formaîon în Sems and Roos (Laen and Advenîîous Sprous/Roos) . . . . . . . . . . . . . . . . . . . . . . . 81
Modîicatîon of the Tree-Rîng Structure Due to Extreme Sîte Condîtîons he Inluence o e Regîona Cîmae on Ecopyogeneîc Deveopmen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85 Modîicaîons Caused by Exreme Weaer Condîîons . . . . . . . . . . 85 Cambîum Injury Caused by Haîsorms . . . . . . . . . . . . . . . . . . . . . . . . 85 Cambîum Injury Caused by Ice Sorms . . . . . . . . . . . . . . . . . . . . . . . . . 87 Cambîum Injury Caused by Lîgnîng . . . . . . . . . . . . . . . . . . . . . . . . . 87 Cambîum Injury Caused by Fros (Fros Rîngs) . . . . . . . . . . . . . . . . . 87 Modîicaîons Caused by Exreme Hydroogîca Canges în e Soî . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92 Arîicîa Raîsîng o e Groundwaer Leve în a Draîned Bog . . . . 93 Rîsîng Coasîne Waer Leve Due o Subsîdence o e Land . . . . . 94 Lowerîng o e Groundwaer Leve în Waerogged Cay Soîs . . . 94 Reduced Soî Waer Conen Ater e Abandonmen o Od Agrîcuura Irrîgaîon Sysems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96 he Perîodîca Dryîng-ou o Rîverbeds and Coasînes . . . . . . . . . . 97 Rîng-Porous Wood Specîes în a Rîverbed wî a Reguaed Waer Leve . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99 Dîfuse-Porous Wood Specîes în e Bed o a Torren wî a Varîabe Waer Leve . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101 Perîodîca Summer Droug în Medîerranean Cîmaes . . . . . . . . 101 Sress Reacîons în Poed Pans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101 Insoaîon As a Facor o Grow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102 he Anaomy o Pans Growîng în Lîg or Sade . . . . . . . . . . . . . 102 Genera Wood-Anaomîca Caracerîsîcs o Densey Saded Pans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103 Reduced Lîg Inensîy Modîies e Sand and Afecs Moraîy107 Reduced Lîg Inensîy în Eac Indîvîdua’s Crown . . . . . . . . . . . 110 Reduced Lîg Inensîy în e Canopy o Even-Aged Sands . . . . 110 Reduced Lîg Inensîy în Open, Man-Made Sands . . . . . . . . . . . 110 Reduced Lîg Inensîy în Tree Sands wî Severa Even-Aged Tree Generaîons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112 Reduced Lîg Inensîy în Coppîce Woods . . . . . . . . . . . . . . . . . . . . 112 Reduced Lîg Inensîy on Abandoned Agrîcuura Land . . . . . . 112 Improved Lîg Condîîons Favor Grow and e Formaîon o New Sands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112 Modîicaîons Due o Permaneny Unavorabe Envîronmena Condîîons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
5.3.5 5.3.5.1 5.3.5.2 5.3.6 5.3.7 5.4 5.4.1 5.4.2 5.4.3 5.4.3.1 5.4.3.2 5.4.3.3 5.4.3.4 5.4.3.5 5.4.3.6 5.4.4 5.5
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5.5.1 5.5.2 5.5.3 5.5.4 5.5.4.1 5.5.4.2 5.5.4.3 5.5.4.4 5.5.5 5.5.6
Dwar Conîers rom Permaros Sîes . . . . . . . . . . . . . . . . . . . . . . . . . 118 Dwar Conîers în Rock Crevîces . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 118 Dwar Conîers on Raîsed Bogs and Fens . . . . . . . . . . . . . . . . . . . . . . 120 Dwar Srubs a e Lîmî o heîr Dîsrîbuîon în e Apîne Be . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120 Dwar Srubs on a Sou-Facîng Sope Above e Tîmberîne . . . 121 Traîîng Azaea on Wînd-Exposed Rîdges în e Apîne Be . . . . . 123 Pans Coonîzîng Rocky Oucrops în Snow-Covered, Exposed Locaîons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124 Dwar Wîows în Apîne Snow Pockes . . . . . . . . . . . . . . . . . . . . . . . 124 Young Conîers a a Rîsîng Upper Tîmberîne . . . . . . . . . . . . . . . . . . 124 Srubs and Dwar Srubs on Saow Soîs în Arîd Desers . . . . . 124
Modîicatîon of the Tree-Rîng Structure Caused by Compressîon and Tensîon 6.1 Opîmîzaîon o Vîa Funcîons Durîng Exreme Mecanîca Sress . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127 6.2 Caracerîsîcs o Reacîon Wood . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127 6.2.1 Compressîon Wood în Gymnosperms . . . . . . . . . . . . . . . . . . . . . . . . . 128 6.2.2 Tensîon Wood în Angîosperms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129 6.2.3 he Dendrocronoogîca Inormaîon Obaîned în Reacîon Wood . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132 6.3 Eccenrîc and Irreguary Saped Sems Wîou Reacîon Wood 132
Modîicatîon of the Tree-Rîng Structure Due to Defolîatîon and Pollardîng 7.1 he Reacîon o Conîers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140 7.1.1 he Arîicîa Remova o Needes rom e Conîer Pinus pinasterBeore Bud-Break . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140 7.1.2 Removîng Woe rees în Managed oress (poardîng). Lîvîng sumps o Anasomoîc Trees . . . . . . . . . . . . . . . . . . . . . . . . . . 142 7.1.3 he Remova o Brances: Cuîng of Lower Brances and Branc Prunîng . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144 7.1.4 Tree Neede Loss Due o Sadîng . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144 7.1.5 Deoîaîon by Insecs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144 7.1.5.1 Larc Budmo Oubreaks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146 7.1.5.2Spruce Budworm Outbreaks (Choristoneura occidentalis and Choristoneura fumiferana). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146 . 7.1.6 Deoîaîon Caused by Fungî . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149 7.1.7 Deoîaîon o Fîr Trees Caused by Unknown Paogens (Fores Decîne în Fîrs) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152 7.1.8 Deoîaîon Caused by Anropogenîc Aîr Pouîon . . . . . . . . . . . . 153 7.1.8.1 Suur Dîoxîde: Dead and Dyîng Larces, Spruces and Bîrces în e Fores Decîne Area o Norîsk, Russîa . . . . . . . . . . . . . . . . . . . 153 7.1.8.2 Suur Dîoxîde: Dead Fîr Trees a Lake Baîka . . . . . . . . . . . . . . . . . . 154 7.1.8.3 Radîaîon Damage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 155 7.2 he Reacîon o Decîduous Wood . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157 7.2.1 Tree Crown Remova . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157 7.2.1.1 he Managemen o Young, Dense Sands . . . . . . . . . . . . . . . . . . . . . 157 7.2.1.2 Decapîaîon Beeces . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 160 7.2.2 Branc Loppîng în Cuura Landscapes . . . . . . . . . . . . . . . . . . . . . . . 162 7.2.2.1 Branc Loppîng o Provîde Lea Fodder . . . . . . . . . . . . . . . . . . . . . . . 162 7.2.2.2 Branc Loppîng o Fruî and Parkand Trees . . . . . . . . . . . . . . . . . . . 165
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7.2.2.3 7.3 7.4 7.5 7.6 7.6.1 7.6.2 7.7
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Hedge Cuîng . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 167 Deoîaîon Caused by Insecs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 170 Deoîaîon Caused by Mammas: Browsîng . . . . . . . . . . . . . . . . . . . 170 Deoîaîon Caused by Fîre . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 170 Adverse Lîg Condîîons Eîmînae e Assîmîaîon Area . . . . . . 176 Lealess Beeces wî a Lîve Cambîum în Coppîce Woods . . . . . . 176 Dyîng Long Soos on Swee Cesnus . . . . . . . . . . . . . . . . . . . . . . . 176 Deoîaîon and Dea Due o Unknown Paogens . . . . . . . . . . . . 176
Modîicatîon of the Tree-Rîng Structure Due to Destructîve Agents în Wood and Bark Reaîonsîp Beween Causes, Pysîoogîca Processes and Tree-Rîng Caracerîsîcs – an Overvîew . . . . . . . . . . . . . . . . . . 179 he Reacîon o e Cambîum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181 Secreory Ducs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181 Resîn Ducs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181 Gum Ducs în Decîduous Wood . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 185 Resîn Pockes (Pîc Pockes) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 187 Caus Tîssue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 188 Wound Cosure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 188 Hîdden Cambîum Injurîes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 194 Open Cambîum Injurîes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 194 he Reacîon o Lîve, Dîferenîaîng Ces . . . . . . . . . . . . . . . . . . . . . 195 he Reacîon o Fuy Dîferenîaed Ces . . . . . . . . . . . . . . . . . . . . . 197 Tyosîs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197 Poypenos în Hearwood and Wound-Margîn Ces . . . . . . . . . . . 199 Cemîca Boundarîes and Demarcaîon Lînes . . . . . . . . . . . . . . . . . 199 Cemîca Boundarîes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 199 Demarcaîon Lînes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 202 he Reacîon Wîîn Dead Wood . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 202 Tangenîa Cracks Wîou Caus Tîssue în Lîve Tree Sems . . . . 202 Radîa Cracks în Lîve Sems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203 Efecs o Geopysîca Evens on Trees . . . . . . . . . . . . . . . . . . . . . . . . 205 Avaances, Snow Movemen, Rock Fa, Landsîdes, Foods, Movîng Ice Paes, Wînd . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 205 Vocanoes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 205 Fores Fîres . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207 Marks în e Cambîum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207 he Reacîon o Bîoogîca-Mecanîca Inluences . . . . . . . . . . . . . . 208 Damage o Trampîng by Red Deer . . . . . . . . . . . . . . . . . . . . . . . . . . . 208 Traces o Bark-Eaîng Rodens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 210 Cambîum Injurîes Caused by Woodpeckers . . . . . . . . . . . . . . . . . . . 210 Pî Fecks, Caus Spos and Gas: Traces o Cambîum-Damagîng Insecs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 214 Tumorous Grow Caused by Fungî . . . . . . . . . . . . . . . . . . . . . . . . . . 215 Sem Rus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215 Wîces’-Broom on Sîver Fîr . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215 Beec Bark Canker . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218 Bacerîa Canker în As Trees . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218 Tumors on Conîers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218 Sem Hyperpasîa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218 Swee Cesnu Bark Canker . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220 Duc Em Dîsease . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220
8.1 8.2 8.2.1 8.2.1.1 8.2.1.2 8.2.2 8.2.3 8.2.4 8.2.4.1 8.2.4.2 8.3 8.4 8.4.1 8.4.2 8.4.3 8.4.3.1 8.4.3.2 8.5 8.5.1 8.5.2 8.6 8.6.1 8.6.2 8.6.3 8.6.4 8.7 8.7.1 8.7.2 8.7.3 8.7.4 8.7.5 8.7.5.1 8.7.5.2 8.7.5.3 8.7.5.4 8.7.5.5 8.7.5.6 8.7.6 8.7.7
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Contents
8.7.8 8.7.9 8.7.10
Honey Fungus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222 he Dea o Twîgs on Fruî Trees . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222 Mîseoe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222
Modîicatîon of the Tree-Rîng Structure Due to Deformed Stems and Anastomosîs 9.1 Naura Anasomosîs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229 9.1.1 Pase One: e Efecs o Pressure . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229 9.1.2 Pase Two: Fusîon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 231 9.2 he Ecoogîca Imporance o Naura Anasomosîs . . . . . . . . . . . . 231 9.3 Gratîng . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 232
Modîicatîon of the Tree-Rîng Structure Due to Wood Decay 10.1 Probems or Dendrocronoogy Because o Wood Decay . . . . . . 237 10.2 Wood Decay Wîou Mîcroorganîsms (Abîoîc Decomposîîon) 237 10.3 Wood Decay Due o Mîcroorganîsms . . . . . . . . . . . . . . . . . . . . . . . . . 237 10.3.1 he Bîoogy o Wood Decay Due o Mîcroorganîsms . . . . . . . . . . . 237 10.3.2 Organîsms ha Lîve în Wood Seen Under e Mîcroscope . . . . . 239 10.3.3 Funga Inesaîon and Decomposîîon o Wood . . . . . . . . . . . . . . . 239 10.3.3.1 Bue Saîn . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 239 10.3.3.2 Red Saîn . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 239 10.3.3.3 Wîe Ro . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 239 10.3.3. Brown Ro . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 239 10.3.3.5 Sot Ro . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 244 10.4 Wood Decay în e Absence o Oxygen (Anaerobîc Condîîons) . 244 10.4.1 Decay în e Lîvîng Sem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 244 10.4.2 Decay Under Waer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 245 10.4.3 he Conservaîon o We Wood . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 245 10.5 Wood Decay Due o Insecs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 245 10.6 Mecanîca Wood Deormaîon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 245 10.7 Fossîîzaîon and Mîneraîzaîon o Wood . . . . . . . . . . . . . . . . . . . . . 245 10.8 Carbonîzaîon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 246
11 Conclusîons
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .257
Subject Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .271
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