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The importance of a compound as a repellent is somewhat novel in bark beetle host selection research. Limonene and other terpenes that kill bark beetles and their vectored fungi have been thought to have a mainly short-range effect as feeding deterrents (Smith, R.H. 1975, Coyne and Lott 1976). The majority of work in this field, however, has looked for compounds that increase with stress and are attractive. Few studies have looked for compounds that decrease with stress and are repellent. SUMMARY The significance of these results rests in three principal areas" 1) The large difference in methyl chavicol between healthy and diseased trees provides a clear olfactory sign of stress for bark beetles, far clearer than has been demonstrated for any monoterpene to date. 2) The stresses that produce the large drop in methyl chavicol are the same stresses that are associated with bark beetle infestation, suggesting that methyl chavicol is one of the biochemical links between ecology and beetle behavior. 3) Methyl chavicol reduces the arrestment and landing of bark beetles in the field sufficiently strongly to suggest management applications for several of our most important bark beetle species with a naturally occurring, abundant host compound. Methyl chavicol is the best candidate so far for a compound that responds to stress and to which beetles respond. It may serve as an indicator and reveal, via biochemical linkage, the elements of host defenses that most strongly affect bark beetle success. Further investigation of other host species applying stress of the sort known to favor beetle colonization may determine some new insights for our understanding of the biochemical basis of conifer defenses. Ultimately, we may understand how environmental stresses that favor beetle colonization produce the characteristics of susceptibility which beetles favor. We may be able to provide the biochemical linkage between ecological conditions and host selection. LITERATURE CITED AHMAD, A., MISRA, L.N., and THAKUR, R.S. 1989. Composition of the volatile oil from Feronia limonia leaves. Planta-Medica 55" 199-200. BARDYSEV, I.I., PAPANOV, G.J., and PERCOVSKIJ, A.L. 1970. Chemical composition of the balsams ofPinus sylvestris and P. nigra growing in Bulgaria. Dokl. AN BSSR 14: 539-40. BERGVINSON, D.J. and BORDEN, J.H. 1991. Glyphosate-induced changes in the attack success and development of the mountain pine beetle and impact of its natural enemies. Entomologia Experimentalis et Applicata. 60:203-212. BORDEN, J.H. 1985. Aggregation pheromones. In Kerkut, G.A. and Gilbert, L.I., eds. Comprehensive Insect Physiology, Biochemistry & Pharmacology. Pergamon Press, Oxford. BRIDGES, J.B. 1987. Effects of terpenoid compounds on growth of symbiotic fungi associated with the southern pine beetle. Phytopathol. 77: 83-85. COBB, EW. Jr., WOOD, D.L., STARK, R.W., and MILLER, P.R. 1968. Theory on the relationships between oxidant injury and bark beetle infestation. Hilgardia 39: 141-152. COBB, EW. Jr., ZAVARIN, E., and BERGOT, J. 1972. Effect of air pollution on the volatile oil from leaves of Pinus ponderosa. Phytochemistry 11: 1815-1818. COYNE, J.F. and LOTT, L.H. 1976. Toxicity of substances in Pine oleoresin to southern pine beetles. J. Georgia Entomol. Soc. 11: 301-305. 231
DUKE, J.A. 1985. CRC Handbook of Medicinal Herbs. CRC Press, Boca Raton, FL. 677 p. HAYES, J.L., INGRAM, L.L. Jr., STROM, B.L., ROTON, L.M., BOYETTE, M.W., and WALSH, M.T. 1994a. Identification of a host compound and its practical application: 4-allylanisole as a bark beetle repellent. Proceedings of the 4th Southern Station Chemical Sciences Meeting; Feb. 1-2, 1994. Gen. Tech. Rep. SO-104. Asheville, NC: U.S. Department of Agriculture, Forest Service. HAYES, J.L., STROM, B.L., ROTON, L., and INGRAM, L.L. Jr. 1994b. Repellent properties of the host compound 4allylanisole to the southern pine beetle. J .Chem. Ecol. p. 1595-1615. HAYES, J.L. and STROM, B.L. 1994. 4-Allylanisole as an inhibitor of bark beetle (Coleoptera: Scolytidae) aggregation. J. Econ. Entomol. 87: 1586-1594. HOBSON, K.R. 1992. Studies on host selection and biology of Dendroctonus valens and fungal host interactions of Dendroctonus brevicomis. Ph.D. Dissertation, University of California, Berkeley. HOBSON, K.R,, WOOD, D.L., COOL, L.G., WHITE, RR., OHTSUKA, T., KUBO, I., and ZAVARIN, E. 1993. Chiral specificity :inresponses by the bark beetle D. valens to host kairomones. J. Chem. Ecol. 19: 1837-1846. HUSSEIN, M.Y., LABS, N.H., and ALI, J.H. 1990. Biological and chemical factors associated with the successful introduction of Elaeidobius kamerunicus Faust, the oil palm pollinator in Malaysia. In van Heemert, C. and de Ruijter, A., eds., The Sixth Int'l Syrup. on Pollination. MARR, K.L. and TANG, C.S. 1992. Volatile insecticidal compounds and chemical variability of Hawaiian Zanthoxylum (Rutaceae) species. Biochem. System. and Ecol. 20: 20%217. MASSON, C. and MUSTAPARTA, H. 1990. Chemical information processing in the olfactory system of insects. Physiol. Rev. 70: 199-245. METCALF, R.L. and LAMPMAN, R.L. 1989. Estragole analogues as attractants for Diabrotica species (Coleoptera: Chrysomelidae) corn rootworms. J. Econ. Entomol. 82: 123-129. MILLER, RR., COBB, EW., Jr., and ZAVARIN, E. 1968. Effect of injury upon oleoresin composition, phloem carbohydrates, and phloem pH. Hilgardia 39: 135-139. MIRO'v; N.T. 1961. Composition of gum turpentines of pines. Tech. Bull. 1239. Washington, De: U.S. Department of Agriculture. NEBEKER, T.E., SCHMITZ, R.E, TISDALE, R.A., and HOBSON, K.R. 1995. Chemical and nutritional status of dwarf mistletoe, armillaria root rot, and comandra blister rust infected trees that may influence tree susceptibility to bark beetle attack. Can. J. Bot. (accepted 11/94, in press) OKUNADE, A.L. and OLAIFA, J.I. 1987. Estragole: an acute toxic principle from the volatile oil of the leaves of Clausena anisata. J. Nat. Prod. 50: 990-991. RUDINSKY, J.A. 1962. Ecology of Scolytidae. Ann. Rev. Entomol. 7: 327-348. SMITH, R.H. 1975. Formula for describing effect of insect and host tree factors on resistance to western pine beetle attack. J. Econ. Entomol. 68(6): 841-844. SMITH, R.M. 1975. Note on gum turpentine of Pinus caribaea from Fiji. N. Zealand J. of Sci. 18: 547-548. TKACZ, B.M. and SCHMITZ, R.E 1986. Association of an endemic mountain pine beetle population with lodgepole pine infected by ArmiUaria root disease in Utah. Res. Note INT-353. Ogden, LIT: U.S. Department of Agriculture, Forest Service. 232
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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
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PHYTOCHEMICAL PROTECTION AND NATURA
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Indirect effects of abiotic factors
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HUNTER, M.D. and PRICE, P.W. 1992.
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A° ADDITIVE C. HYBRID SUSCEPTIBILI
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Galls, leaf miners, or leaf folds w
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Table 2.--Summary of the hypotheses
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the inheritance patterns of seconda
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FLOATE, K.D. and WHITHAM, T.G. 1993
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Elements of the Suitability/Defense
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Table l.--Comparing the mean number
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ACKNOWLEDGEMENTS The authors sincer
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SINGH, D.R 1986_ Breeding for resis
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The objectives of this presentation
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Table 1.--Average weevil attack on
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1977, Kline and Mitchell 1979, Wood
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Further research is underway to cla
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A SUGI CLONE HIGHLY PALATABLE TO HA
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RESULTS Seasonal Stability of Palat
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-
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OGAWA, A., NAGATA, Y., and SUKENO,
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MATERIALS AND METHODS Field Work In
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Laboratory Procedures The week afte
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E 120 Nogent-sur-Vernisson Remnings
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Interdependence Between Reaction Zo
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alance hypothesis (Lorio 1986) cann
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SOLHEIM, H., LANGSTROM, B., and HEL
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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
Page 236 and 237: FURNISS, M.M., LIVINGSTON, R.L., an
Page 238 and 239: METHODS In the spring and summer of
Page 242 and 243: WERNER, R.A. 1995. Toxicity and rep
Page 244 and 245: 41 40 42 \ 4t 42 41 42 ",,,\ 42 40
Page 246 and 247: Table 1.--Summary data for the 26 p
Page 248 and 249: One weakness of using data from gen
Page 250 and 251: Lower Peninsula (populations 14-26)
Page 252 and 253: AUCLAIR, A.ND., WORREST, R.C., LACH
Page 254 and 255: KELLOMAKI, S., HANNINEN, H., and KO
Page 256 and 257: WARGO, RM. and HAACK, R.A. 1991. Un
Page 258 and 259: investigation. Sampling was done on
Page 260 and 261: Figure 2.--Changes in protein preci
Page 262 and 263: q) J 400 T J ! i i i i o control tr
Page 264 and 265: attacked trees than in control tree
Page 266 and 267: ACIDIC DEPOSITION, DROUGHT, AND INS
Page 268 and 269: Table 1.--Impact of acidic fog upon
Page 270 and 271: MENGEL, K., BREININGER, T., and LUT
Page 272 and 273: THE RESISTANCE OF SCOTCH PINE TO DE
Page 274 and 275: EXPERIMENTAL METHODS Experiments we
Page 276 and 277: C w Q. 1,200 1,000 o 800 C_ --- 600
Page 278 and 279: 0 .................................
Page 280 and 281: FONTAtNE, R.G. 1985. Forty years of
Page 282 and 283: Host Defenses An important componen
Page 284 and 285: Functional Heterogeneity of Forest
Page 286 and 287: The Connectivity Index (CI). The CI
Page 288 and 289: 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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