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PREFACE This proceedings is the result of an international symposium that was held February 2-6, 1994 in Maui, Hawaii. It was organized under the guidelines of the International Union of Forestry Research Organizations (IUFRO) by the North Central Forest Experiment Station of the U.S. Forest Service, and the Finnish Forest Research Institute. Two IUFRO Working Parties ($7.01-02, Mechanisms and genetics of plant resistance against insects, and $7.01-03, Mechanisms and genetics of plant resistance against mammals) convened for this international conference on the dynamics of herbivory and the elucidation of plant defenses against herbivores. The proceedings is published and distributed by the USDA Forest Service, North Central Forest Experiment Station in recognition and unstinting endorsement of the IUFRO goals of promoting strong world partnerships, particularly in forest science, for the sustained management, conservation, and the betterment of the World's forest and woodland ecosystems. ACKNOWLEDGMENTS The editors gratefully acknowledge the generous support of the North Central Forest Experiment Station and the Finnish Forest Research Institute for underwriting the costs of organizing, assembling, editing, printing, and distributing this publication. The editors also wish to thank all of the particpants whose penetrating and spirited contributions helped make this scientific exchange highly worthwhile.
TABLE OF CONTENTS Seeking The Rules of Plant-Herbivore Interactions l. Why are tree responses to herbivory so variable? E. Haukioja and T. Honkanen .............................................................................................................................................. 1 2. Damage-induced nutritional changes in pine foliage: an overview. R Lyytikainen ..................................................................................................................................................................... I1 3. Stand and landscape diversity as a mechanism of forest resistance to insects. T. Schowalter ...................................................................................................................................................................... 21 4. On neighbor effects in plant-herbivore interactions. J. Tuomi, M. Augner, and R Nilsson .................................................................................................................................. 28 5. Defense theories and birch resistance. M. Rousi, J. Tahvanainen, W. Mattson, and H. Sikanen .................................................................................................... 39 6. A redox-based mechanism by which environmental stresses elicit changes in plant defensive chemistry. D.M. Norris ........................................................................................................................................................................ 46 7. Fungal endophytes: Contrasting effects in trees and grasses. S.H. Faeth ........................................................................................................................................................................... 58 8. Propagating the effects of herbivore attack in an object-oriented model of a tree. J. Perttunen, H. Saarenmaa, R. Siev_inen, H. Salminen, A. Pouttu, and J. V_ikev_i............................................................ 60 Variable Plants and Herbivores: Idiosyncrasies of Individual Cases 9. Defoliation of Fagus crenata affects the population dynamics of the beech caterpillar, Quadricalcarifera punctatella. N. Kamata, Y. Igarashi, and S. Ohara ................................................................................................................................ 68 10. The impact of flowering on the suitability of balsam fir for the spruce budworm varies with larval feeding behavior. E. Bauce, and N. Carisey .................................................................................................................................................... 86 11. Staminate flowering and tree phenology affect the performance of the spruce budworm. W.J. Mattson, B.A. Birr, and R.K. Lawrence ..................................................................................................................... 97 12. Folivore feeding on male conifer flowers: defence avoidance or bet-hedging? T.S. Jensen ........................................................................................................................................................................ 104 13. The blackmargined aphid as a keystone species: a predator attractor redressing natural enemy imbalances in pecan systems. M. K. Harris and T. Li ...................................................................................................................................................... 112 14. Ponderosa pine response to nitrogen fertilization and defoliation by the pandora moth, Coloradia pandora Blake (Lepidoptera: Saturniidae). B.E. Wickman, R.R. Mason, and H.G. Paul ..................................................................................................................... 118 15. Phytochemical protection and natural enemies: what regulates a root and stem boring swift moth? A.V. Whipple, and D.R. Strong ........................................................................................................................................ 127
Page 1 and 2: "
Page 3: DYNAMICS OF FOREST HERBIVORY: QUEST
Page 7 and 8: 32. Companion planting of the nitro
Page 9 and 10: GRIMALSKY, ¥.I., Research Institut
Page 11 and 12: POUTTU, ANTTI, Finnish Forest Resea
Page 13 and 14: persist in mature leaves and affect
Page 15 and 16: THE SINK-SOURCE HYPOTHESIS: TYPE AN
Page 17 and 18: late season defoliation decreases t
Page 19 and 20: BLANCHE, C.A., LORIO, EL., Jr., SOM
Page 21 and 22: NIEMELA, R, TUOMI, J,, and LOJANDER
Page 24: induced reactions have been studied
Page 28 and 29: summer, but the other sawfly specie
Page 30 and 31: LARSSON, S., BJ(_RKMAN, C., and GRE
Page 32 and 33: STAND AND LANDSCAPE DIVERSITY AS A
Page 34 and 35: not related to resistance to the se
Page 36 and 37: In contrast to the situation in eas
Page 38 and 39: ROOT, R.B. 1973. Organization of a
Page 40 and 41: We tlhus neglect two potential sour
Page 42 and 43: c__ c__ 1T1 Ill a) a) 0 0 0 e 1 o e
Page 44 and 45: c_ >l.s S S S m N < DN DN < D c DN
Page 46 and 47: :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
Page 52 and 53: 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
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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
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- = 120 N /'_" =40 1/ _ j Days 0 _
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BIGGS, A.R. 1985. Suberized boundar
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DIFFERENTIAL SUSCEPTIBILITY OF WHIT
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Table 2.--Mortality of white fir pr
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Results from the geographic range p
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esulting seedlings were then rooted
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-0.2 0 5 82. _j -0,4 e a a a D4 E I
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1960, Kalkstein 1976). Increased su
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RYAN, B.F., JOtNER, B.L., and RYAN,
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It is seldom feasible to control wa
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15 15 u. 10 |O O N tal 5 5 g ao o 4
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Only recently have we conducted stu
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esistance to beetle attack. Another
Page 218 and 219:
LORIO, EL., Jr'. and HODGES, J.D. 1
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EFFECTS OF ROOT INHABITING INSECT-F
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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
Page 240 and 241:
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
Page 282 and 283:
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
Page 296:
) i,i_i)i_)i_i_ U.S. Departme_]t of
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