Theworksho pwa sorganize db y the InternationalUnio no fForestr yResearc h Organizations and sponsoredb yth eNort hAtlanti c Treaty Organization,th e Heidemij (RoyalNetherland s LandDevelopmen t andReclamatio n Society)an d theMinistr y ofAgricultur e andFisherie s ofth eNetherlands . 'O Resistance to diseases and pests infores t trees Proceedings of the Third International Workshop on the Genetics of Host-Parasite Interactions in Forestry, Wageningen,th e Netherlands, 14-21 September 1980 Editedb y H.M. Heybroek, B.R.Stepha n and K.vo nWeissenber g X pudoc Y Centre for Agricultural Publishing and Documentation Wageningen - 1982 YS "v lb1-*. "2- 2-fc? - -£> ' PrecedingWorkshops : Genetic improvement fordiseas e andinsec tresistanc e of foresttrees , 30Augus tt o 11Septembe r 1964;a NAT O andNS FAdvance d Study Institute. Pennsylvania StateUniversity ,Universit y Park,Pennsylvania ,USA . Proceedings:H.D . Gerhold,E.J . Schreiner,R.E .McDermot t& J.A .Winiesk i (Eds):Breedin gPest-Resistan t Trees.Pergamo nPress ,Oxford . 1966.50 5p . Basicbiolog y and international aspectso frus tresistanc e infores t trees, 17-24Augus t 1969;a NATO-IUFR OAdvance d Study Institute. University of Idaho,Moscow , Idaho,USA . Proceedings:R.T .Bingham , R.J.Hof f& G.I .McDonal d (Eds):Biolog y ofRus t Resistance inFores tTrees .USD A ForestService ,Miscellaneou s Publication No 1221.1972 .68 1p . ISBN 9022 0 07944 © Centre forAgricultura l Publishing andDocumentatio n (Pudoc), Wageningen,1982 . 'Nopar to fthi spublicatio nma yb ereproduced , stored ina retrieva l sys­ tem,o rtransmitte d inan y formo rb y anymeans ,mechanical ,photocopying , recording,o rotherwise ,withou tth eprio rwritte npermissio n ofth e publisherPudoc ,P.O . Box4 ,670 0A AWageningen ,Netherlands . Printed inth eNetherlands . Contents Breeding for balance? An introduction ix TOPIC 1: INTRODUCTORY B.J. Zobel: Theworld' s need forpest-resistan t foresttree s 1 I.A.S. Gibson, J. Burleu <£ M. R. Speight: The adoptiono f agricultural practices forth e developmento fheritabl e ' resistance topest s andpathogen s in forestcrop s 9 TOPIC 2: HOST-PARASITE INTERACTION AND THE ENVIRONMENT W.H. Parry: Therol eo fth eenvironmen t inhost-insec t interactions 22 W.H. Parry: Variability inconife rneedl e freezing asa resistance factort o aphids 32 P. Schutt: Trees,fungi ,an denvironmen t 39 M. Ridé: Environmental factors affecting host-bacteria inter­ actions inbacteria l diseases of foresttree s and shrubs 48 H.S. McNabb Jr., R.B. Hall &M.E. Ostry: Biological and physical modifications ofth eenvironmen t andth eresult ­ ing effectupo nth ehost-parasit e interactions inshort - rotation treecrop s 60 TOPIC 3: HOST-PARASITE INTERACTIONS AND GENETICS ON THE INDIVIDUAL PLANT LEVEL M.K. Harris: Genes forresistanc e to insects inagricultur e witha discussio n ofhost-parasit e interactions in Carya 72 R.R. Nelson: Ongene s fordiseas e resistance inplant s 84 TOPIC 4: RESISTANCE MECHANISMS AND THEIR GENETIC CONTROL G.S. Puritch <£G.D. Jensen: Non-specifichost-tre eprocesse s occurring inbar k inrespons et odamag e andthei rrol e indefens e 94 A.L. Shigo: Treesresistan tt osprea d ofdeca y associated withwound s 103 F.F. Jewell, D.C. Jewell & C.H. Walkinshaw: Histopathology ofanatomica lmechanism s forresistanc e to fusiform rusti nslas hpin e 110 B. B. Kinloch: Mechanisms and inheritance ofrus t resistance inconifer s 119 R. Siwecki, A. Werner, Z. Krzan & F. Miodzianowski: Resistance mechanisms in interactionsbetwee npoplar s andrus t 130 D.M. Elgersma: Resistance mechanisms ofelm st oDutc hel m disease 143 Y. Mamiya: Pinewil tan dpin ewoo dnematode : histopathological aspects ofdiseas e development 153 TOPIC 5: BIOCHEMICAL ASPECTS IN RESISTANCE BREEDING M.R. Bridgen & J.W. Hanover: Indirect selection for pest resistance usingterpenoi d compounds 161 H.J. Schuck: Monoterpenes andresistance.o fconifer s to fungi 169 F. Scholz & B.R. Stephan: Buffering ofp H inplan torgan s and resistance against fungi 176 Z.C. Chen & P.D. Manion: Histoenzymologicalapproac hfo rde ­ tectinghost-parasit e interactions in foresttree s caused byheart-ro t diseases 187 G.I. Forrest: Preliminary work onth erelatio nbetwee nresis ­ tancet o Fomes annosus andth emonoterpen e composition ofSitk a spruce resin 194 E.I. Ladejtschikova & G.M. Pasternak: Biochemical aspects of the resistance of Pinus sylvestris to Fomes annosus 198 P.J. Charles, A. Delplanque, A. Marpeau, C. Bernard-Dagan & M. Arbez: Susceptibility ofEuropea nblac kpin e (Pinus nigra) toth eEuropea npin e shootmot h (Rhyacionia buoliana): variations of susceptibility atth eprovenanc e andindividua l levelo fth epin e andeffec to fterpen e composition 206 TOPIC 6: GENETIC VARIATION IN FUNGAL PATHOGENICITY CE. Caten: Genetic basis forvariatio n in fungi 213 CM. Brasier: Genetics ofpathogenicit y in Ceratocystis ulmi andit ssignificanc efo rel mbreedin g 224 H. R. Powers, Jr. :Pathogeni cvariabilit ywithi nth egenu s Cronartium 236 G.A. Snow, W.L. Nance & E.B. Snyder: Relativevirulenc e of Cronartium quercuum f. sp. fusiforme on loblollypin e fromLivingsto nparis h 243 R.D. Raabe: Variationi npathogenicit yan dvirulenc ei n Fomes annosus and Armillaria mellea 251 L. Dimitri: Somehost/parasit e relationshipsbetwee nNorwa y spruce (Picea abies) and Fomes annosus 260 C. Delatour: Behaviouro f Fomes annosus inth este mo fNorwa y sprucean di nth elaborator y 268 C.H. Driver, D.R. Morse & R.L. Edmonds: Detectiono fdiseas e resistancei nDouglas-fi r seedlingsan dvariatio ni n pathogenicityi n Phellinus weirii bymonitorin gwate r stress after inoculation 275 L.D. Dwinell & J.B. Barrows-Broaddus: Variability ofhos tan d pathogeni nth epitc h canker complex 283 TOPIC 7: POPULATION GENETICS AND BREEDING FOR BALANCE N. Lorimer: Pestoutbreak sa sa functio no fvariabilit yi n pestsan dplant s 287 W.J. Mattson, N. Lorimer &A.A . Leary: Roleo fplan tvariabil ­ ity (traitvecto r dynamicsan ddiversity )i nplant/herbi ­ vore interactions 295 E.G. Brittain: Stabilityi na multi-componen thost-pathoge n model 304 D. Krusche: Selectioni nhost-parasit e systems 312 C. Person, R. Fleming & L. Cargeeg: Non-specific interaction basedo npolygene s 318 H.M. Heybroek: Monocultureversu smixture :interaction s between susceptiblean dresistan ttree si na mixe d stand 326 W.J. Libby: Whati sa saf enumbe ro fclone spe rplantation ? 342 A. Segal, J. Manisterski, J. A. Browning, G. Fischbeck & I. Wahl: Balancei nindigenou splan tpopulation s 361 J. A. Browning, J. Manisterski, A. Segal, G. Fischbeck <S I. Wahl: Extrapolationo fgeneti can depidemiologi c concepts from indigenousecosystem st oagroecosystem s 371 V. Umaerus: Newproblems ,trend san dvision si nagricultura l resistancebreedin g 381 TOPIC 8: PRESENTATION AND ANALYSIS OF CURRENTBREEDING PROGRAMS K. Ohba: Breedingo fpine s forresistanc et oth ewoo dnematode , Bursaphelenchus lignicolus 387 G. Bazzigher: TheSwis s£ndothia-resistanc ebreedin gprogra m 396 G.I. McDonald & R.J. Hoff: Engineeringbliste rrust-resistan t westernwhit epin e 404 C. Delatour & Y. Birot: Theinternationa lIUFR Oexperimen to n resistanceo fwhit epine st obliste r rust (Cronartium ribicola). TheFrenc htria l 412 I. Blada: Relativebliste r rustresistanc eo fnativ eand * introduced whitepine si nRomani a 415 B. Zobel: Developing fusiform-resistant treesi nth esouth ­ easternUnite d States 417 H.R. Powers, Jr., S.D. Hubbard & R.L. Anderson: Testing for resistancet ofusifor m rusto fpin e 427 P. Raddi & A. Ragazzi: Italianstudie so nresistanc et opin e blisterrus t (Cronartium flaccidum) 435 R. Jalkanen: Lophodermella sulcigena inclone s andprogenie s ofScot spin ei nFinlan d 441 I. Baumanis, D. Pirags & Z. Spalvins: Resistance trials of Scotchpin eclone si nth eLatvia nSS R 448 0. Martinsson: Impacto fwester n gall rusti nnatura l stands oflodgepol epin e ' 450 R.J. Dinus: Integrating resistancebreedin g andtre eimprove ­ ment 452 G.I. McDonald: Progeny testingo fDouglas-fi r seedlingsfo r resistancet owester n sprucebudwor m 460 1. Blada: Testing larch clonesfo r Adelges laricis resistance . 466 W.H. Fogal, E.K. Morgenstern, P. Viidik & C.W. Yeatman: Varia­ tioni nsusceptibilit yo fnativ ean dintroduce d coniferous treest osom e insectso feaster nCanad a 472 M. Ridé : Internationalbacteria l cankertestin gprogramm eo n poplar 478 TOPIC 9: CONCLUDING REMARKS H.D. Gerhold: Acoevolutionar y viewo fresistanc e breeding andresearc h 487 Recommendations 496 Index of hosts and parasites 498 Vlll Breeding for balance? An introduction Breeding forresistanc e to aparasit e isbreedin g against aflexibl e adversary: onecanno ttak e itfo rgrante d thatresistance ,onc e obtained, will remain atth e same level.Th eparasit ema y comeu pwit hrar e orne w geneticvariant s thatcircumven tth eresistanc e andrende r thehos tsuscep ­ tible.Therefore ,breedin g forresistanc e toa parasit e is fundamentally different frombreedin g forresistanc e to frost,drought ,sal to rsom e otherenvironmenta l extreme. ' Foresttre ebreeding , at firstsight ,i sparticularl y vulnerablet o changes inth eparasite , asa quic kchang e fromon etre e species to another isdifficul t to achieve infores tmanagemen twithou tbi g losses.Thi si s causedb y the facttha t forestryha s alon gtim e scale compared toagricul ­ ture:th e agriculturalplan tbreede rma yconside r someresistanc e tob e 'stable'i fi tdoe sno terod ewithi ntwent y years,whic h ismor e thanth e normal lifetime ofa cultivar ; formos t foresttrees ,however ,thi s isonl y halfth e lifetime ofa singl ecrop . There isn ocaus e fordespair ,however , forther