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1960, Kalkstein 1976). Increased susceptibility may be due to a reduced yield of constitutive resins. However, this explanation may not hold when drought is only mild (cf. Dunn and Lorio 1993). It seems likely that the degree of stress to which trees are exposed will influence their resistance. Assuming that a higher level of stress would have made the trees more susceptible, it is relevant to ask how often it occurs in the field under natural conditions. In this study the sheltered trees were deprived (depending on their root systems) of as much as half a year's precipitation (c. 400 mm) prior to inoculation, and a further 200 rnm were lost thereafter. Such an abrupt loss of water does not seem likely to occur under field conditions in Norway. At the norrnat time of mass flight by I. typographus (i.e., in May), soil water reserves are not likely to be seriously depleted. In the present case it can be hypothesized that the relativley mild drought may have triggered a mobilization of some defence mechanism in Clone 194; it may have induced tile accumulation of defensive compounds or have elicited the development of defence-related cell types (V.R. Franceschi, pets. comm.). A more severe drought might have prevented the mobilization of such defences by critically reducing the availability of some important resources, but this remains to be proved. SUMMARY Young Norway spruce trees of two different clones were artificially deprived of precipitation water over a period of 8 months, starting in late October. In late June, pre-dawn water potentials of the experimental trees were significantly lower than those of unstressed control trees, but the stress was still tlairly mild. At this time both categories of trees were inoculated with a standard dose of the blue-stain fungus Ophiostoma polonicum, a usual associate of the spruce bark beetle Ips typographus. The stressed trees of one clone proved to be significantly more resistant to infection than unstressed controls of the same clone. The other clone showed a similar but much less pronounced trend. It is suggested that the mild stress has triggered the mobilization of some defence mechanism in the former clone. ACKNOWLEDGEMENTS The study was supported by a grant from the Agricultural Research Council of Norway (Contract no. 205030). Thanks are due to Tore SkrOppa and colleagues for providing experimental trees; to Line Hamar, Heidi Knutsen, Olaug Olsen, and Torfinn Smther for field assistance; to Olaug Olsen for fungal reisolation; to Olaug Olsen and Halvor Solheim for fungal identification; and to Halvor Solheim and Ye Hui for critical examination of the manuscript. LITERATURE CITED BERRYMAN, A.A. 1972. Resistance of conifers to invasion by bark beetle-fungus associations. BioScience 22: 598-602. CATES, R.G. and ALEXANDER, It. 1982. Host Resistance and Susceptibility, p. 212-263. In Mitton, J.B. and Sturgeon, K.B., eds. Bark Beetles in North American Conifers. University of Texas Press, Austin. CHRISTIANSEN, E. 1985a. Ceratocystis polonica inoculated in Norway spruce: Blue-staining in relation to inoculum density, resinosis, and tree growth. Eur. J. For. Pathol. 15' 160-167. CHRISTIANSEN, E. 1985b. lps/Ceratocystis-infection of Norway spruce: What is a deadly dosage? Z. Angew. Entomol. 99:6-11 CHRISTIANSEN, E. 199 I. Ips typographus and Ophios'toma polonicum versus Norway spruce: Joint attack and host defence, p. 321-324. In Baranchikov, Y., Mattson, W.J., Hain, F., and Payne, T., eds. Forest insect guilds: patterns of interaction with host trees. Gen. Tech. Rep. NE-153. Radnor, PA: U.S. Department of Agriculture, Forest Service: 321-334. 197
CHRISTIANSEN, E. 1992. After-effects of drought did not predispose young Picea abies to infection by the bark beetletransmitted blue-stain fungus Ophiostoma polonicum. Scand. J. For. Res. 7: 557-569. CHRISTIANSEN, E. and BAKKE, A. 1988. The spruce bark beetle of Eurasia, p. 479-503. In Berryman, A.A., ed. Dynamics of Forest Insect Populations. New York and London: Plenum Publishing Corporation. CHRISTIANSEN, E. and ERICSSON, A. 1986. Starch reserves in Picea abies in relation to defence reaction against a bark beetle transmitted blue-stain fungus, Ceratocystispolonica. Can. J. For. Res. 16: 78-83. CHRISTIANSEN, E. and HORNTVEDT, R. 1983. Combined Ips/Ceratocystis attack on Norway spruce, and defensive mechanisms of the trees. Z. Angew. Entomol. 96:110-118. CHRISTIANSEN, E. and SOLHEIM, H. 1990. The bark beetle-associated blue-stain fungus, Ophiostoma polonicum, can kill various spruces and Douglas fir. Eur. J. For.Pathol. 20: 436-446. CHRISTIANSEN, E., WARING, R.H., and BERRYMAN, A.A. 1987. Resistance of conifers to bark beetle attack: Searching for general relationships. For. Ecol. Manage. 22: 89-106. CRAIGHEAD, F.C. 1928. Interrelation of tree-killing bark beetles (Dendroctonus) and blue stains. J. Forestry 26: 886- 887. DUNN, J.R and LORIO, EL. 1993. Modified water regimes affect photosynthesis, xylem water potential, cambial growth, and resistance of juvenile Pinus taeda L. to Dendroctonusfrontalis (Coleoptera: Scolytidae). Environ. Entomol. 22: 948-957. FURNISS, M.M., SOLHEIM, H., and CHRISTIANSEN, E. 1990. Transmission of blue-stain fungi by Ips typographus (Coleoptera: ScoOytidae) in Norway spruce. Ann. Entomol.Soc. Am. 83: 712-716. HAVRANEK, W.M. and BENECKE, U. 1978. The influence of soil moisture on water potential, transpiration and photosythesis of conifer seedlings. Plant and Soil 49:91-103. HORNTVEDT, R. 11988. Resistance of Picea abies to Ips typographus: Tree response to monthly inoculations with Ophiostoma polonicum, a beetle transmitted blue-stain fungus. Scan. J. For. Res. 3" 107-114. HORNTVEDT, R., CHRISTIANSEN, E., SOLHEIM, H., and WANG, S. 1983. Artificial inoculation with lps typographusassociated blue-stain fungi can kill healthy Norway spruce trees. Medd. Nor. inst. skogforsk. 38(4): 1-20. HORNTVEDT, R. and SOLHEIM, H. 1991. Pathogenicity of Ophiostoma polonicum to Norway spruce: The effect of isolate age and inoculum dose. Medd. Skogforsk.44(4): 1-11. HSIAO, T.C. 1973. Plant responses to water stress. Ann. Rev.Plant Physiol. 24: 519-570. KALKSTEIN, L.S. 1976. Effects of climatic stress upon outbreaks of the southern pine beetle. Environ. Entomol.5: 653- 658. MATSON, RA. and HAIN, F.R 1985. Host conifer defence strategies, p. 33-42. In Safranyik, L., ed. The Role of the Host in the Population Dynamics of Forest Insects. Proc. IUFRO Conf., Banff, Alberta, Canada, 4-7 Sept. 1983. Pacific For. Res. Centre, Can. For. Serv., Victoria B.C. MILLER, J.M. and KEEN, F.R 1960. Biology and control of thewestern pine beetle. U.S. Department of Agriculture, 381 p. RAFFA, K.F. and BERRYMAN, A.A. 1983. The role of host plant resistance in the colonization behavior and ecology of bark beetles (Coleoptera: Scolytidae). Ecol. Monogr. 53: 27-49. 198
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DYNAMICS OF FOREST HERBIVORY: QUEST
Page 5 and 6:
TABLE OF CONTENTS Seeking The Rules
Page 7 and 8:
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
Page 19 and 20:
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
Page 30 and 31:
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
Page 36 and 37:
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
Page 42 and 43:
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
Page 48 and 49:
oth as protective weapons and as po
Page 50 and 51:
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
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 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 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 240 and 241: The importance of a compound as a r
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
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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
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
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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
Page 282 and 283:
Host Defenses An important componen
Page 284 and 285:
Functional Heterogeneity of Forest
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The Connectivity Index (CI). The CI
Page 288 and 289:
FUNCTIONAL HETEROGENEITYANALYSIS :
Page 290 and 291:
The Modified Hazard Map. The next l
Page 292 and 293:
systems facilitate mapping of fores
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KOLASA, J. and ROLLO, C.D. 1991. In
Page 296:
) i,i_i)i_)i_i_ U.S. Departme_]t of
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