Decay does not cause fungi
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- Dwight Heath
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1 Plant Health Care: Callus, Woundwood, Barrier Zones, Reaction Zones, and More Christopher J. Luley, Ph.D. Urban Forestry LLC Naples, NY Decay does not cause fungi Robert Hartig, 1874 Father of Forest Pathology 1
2 Wound Response, Decay Assessment and PHC Biology of tree response to wounding External Wound Response PHC and Pest resistance Decay assessment Internal Response CODIT Decay Progression, and Decay Fungi Decay Assessment ANSI Risk BMP Risk TRAQ 2
3 Wounds Depth, Size + Type Matter Most decay fungi infect through wounds *Depth-Expose heartwood* Width- Wide wounds more important than long wounds Size- Kills sapwood 11 inch (900 cm) Circle 2X more likely to become infected than smaller wounds *Type* Most Heart rot fungi infect through branch or root wounds Greater than 2 inches 3
4 Wound Type Schwarze and Heuser (2006) showed that shallow sapwood wounds are relatively unimportant in heartwood decay infection More important in wound parasite infection Sapwood type decay Wounds that expose heartwood are more likely to heart rot type decay 4
5 Berry,
6 6
7 Decay Incidence by DBH Random Survey of 1800 Street Trees Percent Diameter Class Decay Severity by Diameter SW/SR = 0.3 or less Number of Trees Diameter Class 7
8 Treatment of Wounds with Trichoderma sp. Extensive literature on effectiveness on wound treatment May not be long lasting enough F. Schwarze claims curing trees with decay with tailored Trichoderma treatment Two Responses to Wounding Closure/Sealing Callus Woundwood Compartmentalization Reaction zones Barrier zones 8
9 Bark or Wound Periderm G.W. Hudler Hudler, G.W Wound healing in bark of woody plants. Journal of Arboriculture 10: Wound Closure Callus and Woundwood 9
10 Callus or Woundwood? Callus or Woundwood? Everyone is WRONG! 10
11 University Extension Publication Biology of Callus and WW Urbanforestryllc.com 11
12 Archive.org Callus Initially forms adjacent to a wound Shigo
13 Callus K.T. Smith, USDA Forest Service Kuster (1913) homogenous, parenchymatic, very thin, walled undifferentiated cells Shigo (1984) initially lacking lignin Fink 1999 Undifferentiated parenchymatic proliferations frequently of mixed origin but having a homogeneous appearance Ikeheuchi et al 2013 Disorganized cell masses that can have varying levels of genetic and cellular differentiation Parenchyma Cells Thin walled living plant cells Parenchyma-tissue made up of parenchyma cells Ray parenchymacollection of cells 13
14 Callus Thin walled, undifferentiaed, mostly round cells that initially lack lignin Forms days/weeks after a wound Formed by Vascular cambium Sapwood ray parenchyma, and/or Bark (phloem) parenchyma Callus Totipotent Blank slate Can form vascular cambium, roots, shoots or whole plant 14
15 Callus 1. May differentiate to form vascular cambium 2. Covers VC while it regains function Short-lived Becomes lignified and loses mitotic ability Sloughed off as woundwood grows Shigo
16 Woundwood forms from 1. Vascular cambium (VC) 2. VC differentiated from Callus Initially shorter, denser cells Less vessels Resin canal increase in conifers Kuster
17 Scenario 1 Dujesiefken and Liese 2014 Callus Forms VC Directly 17
18 Callus Forms VC Directly DeBarked Tilia 1. Young callus forms as isodiametric parenchymatous cells Sapwood parenchyma 2. Cambial cells remaining at surface die 3. Callus forms wound periderm (Bark) 4. Cambial zone forms under new periderm From callus New cambium produces xylem and phloem Callus can form vascular cambium which then forms wound wood directly 18
19 Scenario 2. Callus Covers VC and VC Regains Function Shigo, 1984 Callus can form vascular cambium which then forms wound wood directly 19
20 Callus may Seal small wounds Sloughed off as wound ages and VC regains function Shigo, 1984 Correct Answer? Both WW and Callus Early in the wound sealing process 20
21 With Time Normal Wood Develops Woundwood Normal wood eventually forms 21
22 Conifers Form traumatic resin canals in woundwood Do NOT form surface callus and woundwood Wound and Callus Why do I Care? Plant Health Care, Pest Resistance, Decay Assessment, Forensics and More Indicator of response Predictor of internal decay resistance 22
23 Woundwood Seals a Wound The Seal not the Heal Callus/WW seal small wounds quickly Injection treatments may impact callus and woundwood formation G. W. Hudler Woundwood Seals a Wound Sets the Stage for Frost Cracks 23
24 Ribs and Frost Cracks Related to old wound and WW Indicate an internal defect, possibly decay Rib Seam 24
25 Rams Horns The wound will never seal Woundwood may be stronger than normal wood (Kane and Ryan 2003) REALLY needs more study Shigo 1984 Woundwood has fungicidal properties Phenolic extractives of wound-associated wood of beech and their fungicidal effect (Vek, Oven, and Humar, 2012) 25
26 Callus and Woundwood Forms quickest when trees are actively growing Marshall, R. P The relationship of season of wounding and shellacking to callus formation in tree wounds. USDA Technical Bulletin p. Impact of Time of year on Wound Closure Wounds made in the fall seal slowest and may have dieback associated with the injury Dujesiefken and Liese
27 Toole and Gammage, Forestry Wound Sealing High Humidity Protection Promote Callus and Woundwood Formation POLYETHYLENE PLASTIC WRAP FOR TREE WOUNDS: A PROMOTER OF WOUND CLOSURE ON FRESH WOUNDS -McDougall and Blanchette, 1996 Effect goes away after 1 week Species effects present 27
28 Callus and Borer Resistance Fierke and Stephen Callus formation and bark moisture as potential physical defenses of northern red oak, Quercus rubra, against red oak borer, Enaphalodes rufulus (Coleoptera: Cermabycidae). Canadian Entomolgist 140: Callus, Woundwood and Borer Resistance 28
29 Assessment of Woundwood in Tree Health and Stability Rate of Formation Age of Wound/Decay Forensics Presence or Absence Forensics-Aging Wounds Shigo
30 Aging EAB Infestation 30
31 Forensics WW is a Type of Response Growth Presence or Absence is a Key PHC Diagnostic 1. Lack of Vitality 2. Pathogen or Insect Attack 3. Location-supply shadow 31
32 Decay Assessment-Presence/Absence of Woundwood Response Decay Assessment- Presence/Absence of Woundwood 32
33 Diagnostics Absence of WW 33
34 Presence/Absence of Response Growth including Woundwood Scenario 1 Dujesiefken and Liese
35 Basic Wood Anatomy Rays- Living parenchyma cells React to decay fungi invasion Attached to phloem and sugar source! Presence or Absence of WW Proxy for tree defense against decay spread 35
36 -Low Failure Rate? (Kane and Ryan 2003) The Lag 36
37 WW is a Type of Response Growth Presence or Absence is a Key PHC Diagnostic 1. Lack of Vitality 2. Pathogen or Insect Attack 3. Location-supply shadow Potential Indicators Old Cankers 37
38 Eutypella canker Killing Woundwood Cerrena unicolor killing woundwood 38
39 Schizophyllum commune attacking woundwood and decaying wood Schizophyllum commune 39
40 Sap Rot Importance Stem Strength Loss Sap Rot Assessment Safest=Assume the stem is completely decayed 40
41 Urbanforestryllc.com Sap Rot-Living Trees 41
42 Sap Rot Assessment Sounding Not Useful for Quantitative Assessment Case Hardening 42
43 Kretzschmaria deusta killing woundwood and decaying wood 43
44 WW is a Type of Response Growth Presence or Absence is a Key PHC Diagnostic 1. Lack of Vitality 2. Pathogen or Insect Attack 3. Locationsupply shadow Supply Shadow Absence of WW 44
45 Woundwood Formation Basipetal Flow (Leaves to Roots) Wound Location Supply Shadow Supply Shadow 45
46 Woundwood Formation Basipetal Flow (Leaves to Roots) Wound Location Supply Shadow Large wounds More WW on sides of wound Pruning wounds and branch unions EAB Assessment Supply Shadow? 46
47 Two Responses to Wounding 1. Closure WW and callus 1. Compartmentalization Barrier zone- new tissue layer Reaction zones-existing tissues Aka Boundary zones, Boundary layers Natural boundaries Reaction to Wounding Tom Smiley Bartlett Tree Experts 47
48 Barrier Zones, Reactions Zones and CODIT Increasing Decay and Pest Resistance Wounds that expose heartwood are more likely to heart rot type decay 48
49 Basic Wood Anatomy Heartwood No living cells! Some heartwood is more resistant decay Heartwood still reacts to wounding Heartwood Resistance to Decay USDA Comparative decay resistance of heartwood of native species. FPL-0513 Greatest resistance of heartwood to decay is in the butt of a tree 49
50 Ripewood Heartwood that is not more resistant to decay False Heartwood Wound Induced Basic Wood Anatomy Sapwood or Xylem Not inherently resistance to decay Resistance due to living cells 50
51 Sapwood We believe that aeration and moisture effects are primarily responsible for the observed patterns of establishment, but we are still unable to exclude the more widely held view that they are dictated by the host responses of living sapwood (Boddy & Rayner, 1983) Vessels-conducting water Tyloses Vessels 51
52 Tyloses Formed by Axial Parenchyma Wood Structure Angiosperms (Hardwoods, Deciduous) Vessels-conduct water Fibers-support 52
53 Conifers-Tracheids Basic Wood Anatomy Rays- Living parenchyma cells React to decay fungi invasion Attached to phloem and sugar source! 53
54 Wood Structure Rays-living parenchyma cells in wood Radial arrangement (Medullary rays- in and out to center of the tree) Axial arrangement (up and down) Wood Structure Cambium Lateral Meristem Phloem to the outside Xylem to the inside Cambial Zone Shigo
55 Basic Wood Anatomy Cambial Zone Basic Wood Anatomy Phloem Inner bark No structural importance Note rays Note lighter color 55
56 Sieve Tube Elements Movement of sugars in the phloem requires energy Basic Wood Anatomy Bark Cambium Trees have 2 cambial layers Phellogen 56
57 Basic Wood Anatomy Bark Inner and Outer bark 57
58 Identification and Interpretation? Dendrothele species Typically Host Specific Widely distributed Host specific Able to degrade suberin Most trees species have one Ostrya Hickory White oak Ash Basidiomycetes/ Agaricales Dendrothele species Pignut hickory White oak 58
59 Dendrothele species Cause of Smooth Patch Require Microscopic ID to Species Dendrothele macrodens White ash Compartmentalization of Decay in Trees or CODIT 59
60 CODIT In Order of Increasing Strength! Walls 1-3 Reactions zones or Boundary layers Wall 1 = Vessels and tracheid plugging Wall 2 = Heartwood and inner growth rings Wall 3 = Rays Strongest Formed by Cambium after Wounding Wall 4 = Barrier zone WALL 4 Formed de novo by cambium Strongest protection layer Designed to keep new wood free of decay Physical boundary Chemical boundary Larger up and down from the wound Rather than around the wound Barrier Zone 60
61 Barrier Zone-formed by cambium Only forms once at a wound site High density of parenchyma cells High suberin content Dujesiefken and Liese 2014 Suberin-Waxy Substance Seals the Wound Chemically variable Cork Polyphenolic and polyaliphatic Water proof Fungitoxic Physical and Chemical barrier 61
62 Barrier Zone Formed in annual ring present at the time of wounding If dormant forms in next ring 62
63 Physical Barrier Barrier Zone Protective Sheath Highly reactive because of parenchyma cells Stronger than reaction zones Most decay fungi can t breach barrier zone Shigo
64 Biological Health Barrier Zone Strength Importance of Plant Health Care Breaching the Barrier Zone can Spread Decay Tom Smiley, Bartlett Tree Experts 64
65 Depends on the Fungus Present Barrier Zone Breach Kersten and Schwarze 2004 Inonotus hispidus Decay grows in then out to cambium New Barrier zone forms!!! With Woundwood! 65
66 CODIT In Order of Increasing Strength! Walls 1-3 Reactions zones or Boundary layers Wall 1 = Vessels and tracheid plugging Wall 2 = Heartwood and inner growth rings Wall 3 = Rays Strongest Formed by Cambium after Wounding Wall 4 = Barrier zone Wall 1 CODIT Details Wall 1 Weakest wall Tyloses and parenchyma Decay spreads up and down fastest Up> Wall 1 can reform as decay spreads Axial and radial parenchyma 66
67 Wall 2 CODIT Details Second weakest wall Is the reaction zone when Wall 4 fails Annual rings Latewood Thick walled and highly lignified Heartwood 67
68 Wall 2 Wall 2 Heartwood Exposed High probability of decay No living tissues to respond 68
69 Wall 3 CODIT Details Ray Parenchyma Radial Axial Second strongest wall Phenolics and free radical Will reform along with Wall 1 and 4 if breached Strength directly affected by tree health? Reaction Zones Wall 2: previous growth ring Wall 1: Plugs in Vessels & tracheids Wall 3: Rays Wall 4: Cells produced Right after wounding Reaction Zones of a good compartmentalizer G. W. Hudler 69
70 Columns of Discoloration Drilling Tom Smiley Bartlett Tree Experts Reaction zones Strength of the RZ Directly tied to tree biological health Stored reserves in ray parenchyma Discolored wood may or may not be decayed Pearce
71 Ash max 12 inches Toole and Gammage, J Forestry 71
72 72
73 Tom Smiley Bartlett Tree Experts Tom Smiley Bartlett Tree Experts Wounding or Breaching of Walls 1-4 Reaction Zones Formed by parenchyma cells High concentrations of polyphenolics 73
74 Reaction Zone Formation Whenever Old Barrier Zone or Reaction Zone is Breached Wall 1, Wall 2, Wall 3 Parenchyma of sapwood Enzymatic reaction of heartwood Critical Assessment in Decay Evaluation Change in decay amount over time Stability of Barrier zone and Reaction zone 74
75 Increasing trunk diameter Amount of sound wood remains static Tree becomes more stable Moment of Resistance MR= pi * D4-d4 32 * D Trunk Diameter Same Load Carrying Capacity -Amount of Sound Wood Decreases -Trunk Diameter Increases Rinn
76 Biology Trumps Biomechanics Tipping Point Biology Trumps Biomechanics Tipping Point We need to know Change in amount of sound wood relative to change in diameter 76
77 Decay Assessment-Presence/Absence of Woundwood Response 77
78 Diagnostics Absence of WW Southern Red Oak Armillaria sp. 78
79 Fungal Identification Matters Type of decay Location of decay How it interacts with barrier zone Future potential decay progression DNA identification of fungi decayed wood Vetdna.com Jack O Lantern Fungus Obligate saprophyte 79
80 Molecular Diagnostics vetdna.com $18 per sample Test wood shavings samples or fruiting structures Results immediately DNA testing of wood or conk Armillaria ssp. (all species) Armillaria gallica Armillaria mellea Armillaria ostoyae Armillaria tabescens Bacterial Leaf Scorch (X. fastidiosa) Bondarzewia berkeleyi Ceratocystis ssp. Dothistroma pini Dothistroma septosporum Dutch Elm Disease (O. ulmi/o. novoulmi) Elm Yellows (Ca. Phytoplasma ulmi) Emerald Ash Borer (Agrilus planipennis) Fire Blight (Erwinia amylovora) Ganoderma applanatum Ganoderma lucidum Ganoderma zonatum Grifola frondosa Kretzschmaria deusta Laetiporus sulphureus Lecanosticta acicola Oak Wilt (C. fagacearum) Phaeolus schweinitzii Pholiota ssp. (all species) Phytophthora ssp. (all species) Phytophthora palmivora Phytophthora ramorum Phytoplasma ssp. (all species) Pinewood Nematode (B. xylophilus) Pleurotus ssp. Pythium genus (all species) Rose Rosette Virus Sparassis spathulata Thousand Canker Disease (Geosmithia morbida) Verticillium genus (all species) Verticillium albo-atrum Verticillium dahlia 80
81 Decay Fungi Mode of Action Dead wood only (Obligate Saprophytes) Large areas due to wounding or branch death Large number of fungi Anatomical deadwood (Obligate Saprophytes) Heartwood primarily Living sapwood (Facultative Pathogens) Break Barrier zones and Reaction zones Cambium killers and Sapwood decayers (Strong Facultative Pathogens) Very small number of fungi Terms Obligate saprophyte- can only live on dead tissues Dead tissues may be in a living tree Facultative parasite or pathogen can infect and live on live and dead tissues Obligate parasite- must have living host 81
82 True Heart Rots Obligate Saprophytes Cannot Breach Wall 4 Perenniporia fraxinophila 82
83 Decay May Produce No Biological Health Symptoms 83
84 Heart Rot Fungi- Some are Weak Facultative Pathogens Usually limited to heartwood Dead sapwood Gain entrance through heartwood wounds May weakly attack living sapwood Host interactions 84
85 Obligate Saprophytes Living Trees Decaying heartwood/dead sapwood Polyporus squamosus? No Worries as Long as Tree Growth/Health is Maintained Wound Closure Continues Wall 4 and/or Reaction Zone Intact 85
86 Mode of Action Facultative Pathogens -attack both dead and living tissues Weak Strong Facultative Pathogens Sapwood Decayers Establish on dead sapwood Can attack living bark/wood Smaller group Schizophyllum commune Cerrena unicolor Trametes versicolor 86
87 Facultative Pathogens-Strong Heart Rot and Sapwood Rot Breach Wall 4 and Reaction Zones Can attack healthy sapwood More aggressive decayers Small group of fungi Kretzschmaria deusta Ganoderma lucidum Armillaria mellea Ganoderma applanatum Canker Rots Ganoderma lucidum 87
88 Ganoderma lucidum aka Reishi Cambium killer! Ustulina (Hypoxylon deustum) Kretzschmaria deusta Burnt Crust Fungus 88
89 Will kill Woundwood, penetrate Barrier zones and Reaction Zones Fungal Identification Matters Jack O Lantern Fungus Obligate saprophyte 89
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