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SOLHEIM, H., LANGSTROM, B., and HELLQVIST, C. 1993. Pathogenicity of the blue-stain fungi Leptographium wingfieldii and Ophiostoma minus to Scots pine: effect of tree pruning and inoculum density. Can. J. For. Res. 23" 1438-1443. SHRIMPTON, D.M. and REID, R.W. 1973. Change in resistance of lodgepole pine to mountain pine beetle between 1965 and 1973 Can. J. For. Res. 3:430-432 • WARING, R.H. and PITMAN, G.B. 1980. A simple model of host resistance to bark beetles. Oreg. State Univ. for. Res. Lab. Res. Note 65. 2 p. WONG, B.L. and BERRYMAN, A.A. 1977. Host resistance to the fir engraver beetle. 3. Lesion development and contain ment of infection by resistant Abies grandis inoculated with Trichosporium symbioticum. Can. J. Bot. 55: 2358- 2365. 177
CAN PHLOEM PHENOLS BE USED AS MARKERS OF SCOTS PINE RESISTANCE TO BARK BEETLES? F. LIEUTIER _,F. BRIGNOLAS l, V. PICRON t, A. YART l, and C. BASTIEN 2 tStation de Zoologie Foresti_re, Institut National de la Recherche Agronomique Ardon, 45 160 Olivet, France 2Station d'Am61ioration des Arbres Forestiers, Institut National de la Recherche Agronomique Ardon, 45 160 Olivet, France INTRODUCTION Bark beetle damage is considerable in all coniferous forests in the temperate zone. In most cases, it results from beetle attacks surpassing the critical density (numbers/m 2)below which trees can successfully reject them. Above this threshold, natural mechanisms of tree resistance are overcome by the beetles and fungi which become successfully established. The level of this critical threshold of attack density is a fundamental expression of the strength or capacity of tree resistance (Berryman 1978, 1982; Christiansen et al. 1987). The basic phenomenon involved in the resistance is a phloem reaction induced by the invaders and localized around them. It consists of a rapid synthesis of chemicals, including secondary metabolites such as terpenes and phenols, which impregnate tree tissues (Reid et al. 1967, Berryman 1972, Shrimpton 1973, Christiansen and Horntvedt 1983, Christiansen et al. 1987, Lieutier et al. 1988, Lfmgstr6m et al. 1992, Lieutier 1993). The critical threshold of attack density depends on the quantity of energy the tree is able to rapidly mobilize for various syntheses in the reaction zones, and this quantity is necessarily limited (Christiansen et al. 1987). Thus, the threshold can be high if the lag time for stopping an aggressor at each point of attack is very short and hence the energy mobilized for that purpose in each reaction zone is low. Under these conditions of arrested attacks, a short reaction zone has been assumed to be more effective than an extended one (Lieutier et al. 1993). Markers of conifer resistance to bark beetles could be very useful; e.g., forecasting susceptibility to beetle attacks and beetle outbreaks, and aiding in selection of highly resistant trees. Because the concentrations of secondary metabolites vary considerably during tree responses to aggression, they merit a special interest. Preliminary studies on phloem phenolics have suggested dramatic changes in Scots pine, Pinus sylvestris L., in response to aggression (Lieutier et al. 1991a). This paper presents the results from two complementary approaches recently developed in France concerning the possible role of phenols in arresting bark beetle invasion and their use as biochemical markers of resistance. Detailed methods and other findings are presented elsewhere (Lieutier et al. 1995). METHODS Experimental Devices In our first experiment, six clones located in the INRA's nursery in Orlfians (France), each represented by one to three 10-year-old trees, were used to test if a relationship existed between reaction zone length and phenolic composition. Trees were each inoculated with a malt agar sporulated culture of Ophiostoma brunneo-ciliatum Math.-K., a fungus previously isolated from lps sexdentatus Boern., and thought to play a role in the establishment of this beetle in Scots pine (Lieutier et al. 1988). Each tree received three inoculations. Three weeks later, the reaction zone around each inoculation point was measured and sampled. Three samples of unwounded phloem were also taken from each tree. Mattson, W.J., Niemel_i, P., and Rousi, M., eds. 1996. Dynamics of forest herbivory: quest for pattern and principle. USDA For. Serv. Gen. Tech. Rep. NC-183, N.C. For. Exp. Sta., St. Paul, MN 55108. 178
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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
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Wood et al. (1987) report that M. c
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MIZELL, R.E 1991. Pesticides and be
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METHODS Study Site The study took p
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the samples. Fertilization had subs
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Table 2.--Percent nutrient content
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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 174 and 175: OGAWA, A., NAGATA, Y., and SUKENO,
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 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
Page 224 and 225: ]Predisposition of Root-infected Tr
Page 226 and 227: chemistry associated with root colo
Page 228 and 229: Implications to Natural Resource Ma
Page 230 and 231: KLEPZIG, K.D., RAFFA, K.F., and SMA
Page 232 and 233: RAFFA, K.F., PHILLIPS, T., and SALO
Page 234 and 235: 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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