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OGAWA, A., NAGATA, Y., and SUKENO, S. 1992. Effects of solvent extraction on resistance of the hybrid between Japanese larch and Kurile larch to voles. Proc. Hokkaido Branch, Japan Wood Res. Soc. 24:61-64 (in Japanese). RADWAN, M.A. 1972. Differences between Douglas-fir genotypes in relation to browsing preference by black-tailed deer. Can. J. For. Res. 2: 250-255. RADWAN, M.A., CROUCH, G.L., HARRINGTON, C.A., and ELLIS, W.D. 1982. Terpenes of ponderosa pine and feeding preferences by pocket gophers. J. Chem. Ecol. 8:241-253. REICHARDT, RB., BRYANT, J.P., CLAUSEN, T.R, and WlELAND, G.D. 1984. Defence of winter-dormant Alaska paper birch against snowshoe hares. Oecologia 65: 58-69. ROUSI, M., TAHVANAINEN, J., and UOTILA, I. 1989. Inter- and intraspecific variation in the resistance of winterdormant birch (Betula spp.) against browsing by the mountain hare. ttolar. Ecol. 12: 187-1192. ROUSI, M., TAHVANAINEN, J., and UOTILA, I. 1991. A mechanism of resistance to hare browsing in winter-dormant European white birch (Betula pendula). Amer. Natur. 137: 64-82. ROUSI, M., TAHVANAINEN, J., and UOTILA, I. 1993. Effects of shading and fertilization on resistance of winter- dormant birch (Betula pendula)to voles and hares. Ecology 74: 30-38. SNYDER, M.A. 1992. Selective herbivory by Abert's squirrel mediated by chemical variability in Ponderosa pine. Ecology 73: 1730-1741. TAHVANAINEN, J., HELLE, E., JULKUNEN-TIITTO, R., and LAVOLA, A. 1985. Phenolic compounds of willow bark as deterrents against feeding by mountain hare. Oecologia 65:319-323. 165
GENETIC AND PHENOTYPIC VARIATION IN THE INDUCED REACTION OF SCOTS PINE TO LEPTOGRAPHIUM WINGFIELDII: REACTION ZONE LENGTH AND FUNGAL GROWTH F. LIEUTIER 1,B. LANGSTROM z, H. SOLHEIM 3 C. HELLQVIST 2, and A. YART l _Station de Zoologie Foresti_re, Institut National de la Recherche Agronomique, Ardon, 45 160 Olivet, France 2Division of Forest Entomology, Swedish University of Agricultural Sciences, S-77073 Garpenberg, Sweden 3Department of Forest Ecology, Norwegian Forest Research Institute, As-NLH, N 1432, Norway. INTRODUCTION The induced defensive reaction of phloem generally plays a decisive role in the resistance of conifers to attack by bark beetles and their associated fungi. It is thought to be induced by fungi (Reid et al. 1967, Berryman 1972, Raffa and Berryman 1983, Christiansen and Horntvedt 1983, Matson and Hain 1985, Christiansen et al. 1987, Lfingstr6m et al. 1992), or by the boring activity of the beetle (Lieutier et al. 1988, Lieutier 1993). In most cases, by stimulating that energy demanding response, the fungi can lower the threshold of attack density above which the tree is overwhelmed and beetle attacks are successful. In all situations, the individual reaction is the basic mechanism of tree resistance. The between-tree variability in the threshold of attack density (Waring and Pitman 1980, Mulock and Christiansen 1986), as well as the between-tree variability in the induced reaction itself (Shrimpton and Reid 1973, Peterman 1977, Lieutier and Ferrell 1989, Lieutier et al. 1993), has been linked to tree vigor. However, the genetic contribution to this variability has never been studied. In Scots pine, Pinus sylvestris L., an induced reaction is essential to resist attacks by lps sexdentatus Boern. and Tomicus piniperda L., and their associated fungi (Lieutier et al. 1988, 1989a; Solheim and L_ngstr6m, 1991; L_ngstr_m et al. 1992). The blue-staln fungus, Leptographium wing_i'eldii Morelet, associated with T. piniperda, does not seem to play a role in the development of the induced reaction and in the success of attacks (Lieutier et al. 1988, 1989b). However, artificial inoculation of its sporulated cultures into Scots pine always incites strong defensive reactions (Lieutier et al. 1989a, Solheim and L_ngstr6m 1991) and the tree's response depends on the number of spores present in the inoculum (Lieutier et al. 1989b). Additionally, it may play a role in tree mortality, since weakened Scots pine can be killed by a high density of artificial inoculations (Solheim et al. 1993). The present study is part of a cooperative project with the following objectives: (1) to study the variation in the induced reactions (reaction zone formation, sapwood occlusion) between Scots pine provenances in two different climatic regions, (2) to study the performance of a French and a Swedish strain of L. wingfieldii in these provenances under different climatic conditions, and (3) to study the defensive chemistry involved in the induced reactions, particularly the roles of resin acids and phenolics, and to search for chemical markers for resistance among these groups of chemicals. This study addresses the first two issues. Mattson, W.J., Niemelfi, P., and Rousi, M., eds. 1996. Dynamics of forest herbivory: quest for pattern and principle. USDA For. Serv. Gen. Tech. Rep. NC-I83, N.C. For. Exp. Sta., St. Paul, MN 55108. 166
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DYNAMICS OF FOREST HERBIVORY: QUEST
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TABLE OF CONTENTS Seeking The Rules
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32. Companion planting of the nitro
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GRIMALSKY, ¥.I., Research Institut
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POUTTU, ANTTI, Finnish Forest Resea
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persist in mature leaves and affect
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THE SINK-SOURCE HYPOTHESIS: TYPE AN
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late season defoliation decreases t
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BLANCHE, C.A., LORIO, EL., Jr., SOM
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NIEMELA, R, TUOMI, J,, and LOJANDER
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induced reactions have been studied
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summer, but the other sawfly specie
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LARSSON, S., BJ(_RKMAN, C., and GRE
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STAND AND LANDSCAPE DIVERSITY AS A
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not related to resistance to the se
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In contrast to the situation in eas
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ROOT, R.B. 1973. Organization of a
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We tlhus neglect two potential sour
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c__ c__ 1T1 Ill a) a) 0 0 0 e 1 o e
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c_ >l.s S S S m N < DN DN < D c DN
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:hen k > 0 suggest that the cue or
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oth as protective weapons and as po
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DEFENSE THEORIES AND BIRCH RESISTAN
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P 45 A L 40 A T 35 A B 30 I L 25 I
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However, hares showed strong prefer
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DUGLE, J.A. 1966. A taxonomic study
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accommodate findings related to mic
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Table l.--Mean ±S.E. area eaten by
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Sulfhydryl-Disulfide Mechanisms In
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Iraa prior experimental analysis of
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FRAZIER, J.L. and HEITZ, J.R. 1975.
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REICHARD, R 1993. From RNA to DNA,
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; trees, and between and within ind
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Encapsulated objects are data struc
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_re3.--The Lignum model can be cont
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SEVERE DEFOLIATION Tree Fine Root +
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RUMBAUGH, J., BLAHA, M., PREMERLANI
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oo0 a b ol ¢' _achiman_i ()® 1,eA
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1000 Conspicuous Defo i at ion 100
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2.oo 2.oo o t 4 t ¢._ e'- 1.80 1.8
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Qualitative Changes in [,eaves and
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t00 2 Yr 1 Yr o_ 80 Treatment Treat
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Table 2.--Body size of Q. purzctate
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06 b AddedBSA < 04 _ 2rag -'=' i:3
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Severe Defol iat ion Site-A 1993 _
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KAMATA, N. and IGARASHI, M. 1995b.
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enclosure %r 20 post-diapausing sec
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2,500 F 000 F NF NF b _7 a _;_ a _7
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8O D 09 i 4,'.) F AS4L a Z___7 2o !
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2,500 _ st2-st4 El st5 0 st6 [_ pup
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96 SLANSKY, E, Jr. 1990. Insect nut
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later, we selected one more floweri
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Table 2.---Mean spruce budworm perf
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Table 3..--Mean spruce budworm perf
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FOLIVORE FEEDING ON MALE CONIFER FL
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Table 3.--Lymantria monacha prefere
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500- 400- i 300 v ¢ m a 200- I00 L
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From the point of view of trees, th
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THE BLACKMARGINED APHID AS A KEYSTO
Page 124 and 125: Wood et al. (1987) report that M. c
Page 126 and 127: MIZELL, R.E 1991. Pesticides and be
Page 128 and 129: METHODS Study Site The study took p
Page 130 and 131: the samples. Fertilization had subs
Page 132 and 133: Table 2.--Percent nutrient content
Page 134 and 135: Some tree responses to fertilizatio
Page 136 and 137: PHYTOCHEMICAL PROTECTION AND NATURA
Page 138 and 139: Indirect effects of abiotic factors
Page 140 and 141: HUNTER, M.D. and PRICE, P.W. 1992.
Page 142 and 143: A° ADDITIVE C. HYBRID SUSCEPTIBILI
Page 144 and 145: Galls, leaf miners, or leaf folds w
Page 146 and 147: Table 2.--Summary of the hypotheses
Page 148 and 149: the inheritance patterns of seconda
Page 150 and 151: FLOATE, K.D. and WHITHAM, T.G. 1993
Page 152 and 153: Elements of the Suitability/Defense
Page 154 and 155: Table l.--Comparing the mean number
Page 156 and 157: ACKNOWLEDGEMENTS The authors sincer
Page 158 and 159: SINGH, D.R 1986_ Breeding for resis
Page 160 and 161: The objectives of this presentation
Page 162 and 163: Table 1.--Average weevil attack on
Page 164 and 165: 1977, Kline and Mitchell 1979, Wood
Page 166 and 167: Further research is underway to cla
Page 168 and 169: A SUGI CLONE HIGHLY PALATABLE TO HA
Page 170 and 171: RESULTS Seasonal Stability of Palat
Page 172 and 173: -
Page 176 and 177: MATERIALS AND METHODS Field Work In
Page 178 and 179: Laboratory Procedures The week afte
Page 180 and 181: E 120 Nogent-sur-Vernisson Remnings
Page 182 and 183: Interdependence Between Reaction Zo
Page 184 and 185: alance hypothesis (Lorio 1986) cann
Page 186 and 187: SOLHEIM, H., LANGSTROM, B., and HEL
Page 188 and 189: In a second experiment, three 30-ye
Page 190 and 191: Considering biochemical pathways, i
Page 192 and 193: - = 120 N /'_" =40 1/ _ j Days 0 _
Page 194 and 195: BIGGS, A.R. 1985. Suberized boundar
Page 196 and 197: DIFFERENTIAL SUSCEPTIBILITY OF WHIT
Page 198 and 199: Table 2.--Mortality of white fir pr
Page 200 and 201: Results from the geographic range p
Page 202 and 203: esulting seedlings were then rooted
Page 204 and 205: -0.2 0 5 82. _j -0,4 e a a a D4 E I
Page 206 and 207: 1960, Kalkstein 1976). Increased su
Page 208 and 209: RYAN, B.F., JOtNER, B.L., and RYAN,
Page 210 and 211: It is seldom feasible to control wa
Page 212 and 213: 15 15 u. 10 |O O N tal 5 5 g ao o 4
Page 214 and 215: Only recently have we conducted stu
Page 216 and 217: esistance to beetle attack. Another
Page 218 and 219: LORIO, EL., Jr'. and HODGES, J.D. 1
Page 220 and 221: EFFECTS OF ROOT INHABITING INSECT-F
Page 222 and 223: OCCURRENCE OF ROOT AND STEM SUBCORT
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]Predisposition of Root-infected Tr
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chemistry associated with root colo
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Implications to Natural Resource Ma
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KLEPZIG, K.D., RAFFA, K.F., and SMA
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RAFFA, K.F., PHILLIPS, T., and SALO
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METHODS The study was installed in
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FURNISS, M.M., LIVINGSTON, R.L., an
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METHODS In the spring and summer of
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The importance of a compound as a r
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WERNER, R.A. 1995. Toxicity and rep
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41 40 42 \ 4t 42 41 42 ",,,\ 42 40
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Table 1.--Summary data for the 26 p
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One weakness of using data from gen
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Lower Peninsula (populations 14-26)
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AUCLAIR, A.ND., WORREST, R.C., LACH
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KELLOMAKI, S., HANNINEN, H., and KO
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WARGO, RM. and HAACK, R.A. 1991. Un
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investigation. Sampling was done on
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Figure 2.--Changes in protein preci
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q) J 400 T J ! i i i i o control tr
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attacked trees than in control tree
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ACIDIC DEPOSITION, DROUGHT, AND INS
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Table 1.--Impact of acidic fog upon
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MENGEL, K., BREININGER, T., and LUT
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THE RESISTANCE OF SCOTCH PINE TO DE
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EXPERIMENTAL METHODS Experiments we
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C w Q. 1,200 1,000 o 800 C_ --- 600
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0 .................................
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FONTAtNE, R.G. 1985. Forty years of
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Host Defenses An important componen
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Functional Heterogeneity of Forest
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The Connectivity Index (CI). The CI
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FUNCTIONAL HETEROGENEITYANALYSIS :
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The Modified Hazard Map. The next l
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systems facilitate mapping of fores
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KOLASA, J. and ROLLO, C.D. 1991. In
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) i,i_i)i_)i_i_ U.S. Departme_]t of
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