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96 SLANSKY, E, Jr. 1990. Insect nutritional ecology as a basis for studying host plant resistance. Florida Entomol. 73" 360- 378. VOLNEY, W.J.A. 1988. Analysis of historic jack pine budworm in the Prairie province of Canada. Can. J. For. Res. 18: 1152-1158. WELLINGTON, W.G. 1950. Effects of radiation on the temperatures of insectan habitats. Sci. Agr. 30: 209-234.
STAMINATE FLOWERING AND TREE PHENOLOGY AFFECT THE PERFORMANCE OF THE SPRUCE BUDWORM WILLIAM J. MATTSON, BRUCE A. BIRR, and ROBERT K. LAWRENCE USDA Forest Service, North Central Forest Experiment Station Pesticide Research Center, Michigan State University, E. Lansing, M148824, USA INTRODUCTION The important role of staminate flowers in the ecology and population dynamics of spruce budworm has been the subject of speculation for more than 50 years (Blais 1952, Greenbank 1963). This suspicion has been fueled by the fact that high populations of budworm have historically been associated with mature, flower-bearing trees (Mott 1963). Moreover, overwintering budworm larvae typically spin their silk shelter in the old staminate flower bracts, and spring emerging 2nd instars prefer to mine and feed in staminate flower buds rather than old needles. Budworms emerge 1 to 4 weeks before vegetative buds have begun to expand and are suitable for attack, whereas flower buds, when present, are always more advanced and thus better synchronized with the early, vernal budworms. Hence, second-stage larvae are invariably concentrated in the host's expanding flower buds. On the other hand, whenever budworm emergence is retarded relative to tree phenology, then larvae tend to go directly to expanding foliage rather than the staminate flower buds (Greenbank 1963). Obviously, then, the importance of flower buds to budworms may be linked to the phenology of the hosts' vegetative buds relative to the timing of spring emergence of the budworms. Larvae that feed within staminate flower clusters may actually grow faster than foliage-feeding larvae because such clusters are miniature greenhouses, trapping more radiant energy than comparable pure foliage environments. However, there is little other experimental evidence to support the hypothesis that staminate flowers can enhance budworm performance and thus contribute to outbreak development. For example, Jaynes and Speers (1949) and Blais (1952) reported that there is no difference in fecundity of budworms having fed on pollen or new foliage. Herms and Mattson (1992) hypothesized that plant reproduction can also indirectly affect herbivore performance through its strong impact on the allocation of plant nutrients and photosynthates. When plants produce large crops of pollen and/or seeds, the reproductive organs may preferentially receive scarce nutrients and energy that might otherwise have gone to plant defenses or even new growth on which phytophages depend. Hence, staminate flowering could indirectly impact budworms by reducing the concentrations of nutrients and perhaps even altering the levels of secondary metabolites in plant foliage. This study was undertaken to simulate and explore the consequences of abundant staminate flower production on the performance of pre-outbreak spruce budworm populations. We hypothesized that budworm larval performance would depend not just on the availability of staminate flower but also on the phenology of a tree's vegetative buds. We predicted that the presence of staminate flowers would be most important on very late flushing trees, and least important on very early flushing trees. METHODS In 1989 and 1991 we selected 20 half-sibling balsam fir, Abies balsamea, that were flowering at the Kellogg Experi- mental Forest (Michigan State University) near Augusta, Michigan. Trees were planted in 1970 and about 6-8 m tall at the time of the experiments. Before budbreak (April 21, 1989, and April 11, 1991), we selected per tree two flowering and two nonflowering branches near midcrown, which were then enclosed with a fine-mesh, cloth sleeve cage wherein we placed about 20 ready to emerge second instar budworm larvae still in their silken hibernacula on a gauze patch. Exactly 1 week 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. 97
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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
Page 56 and 57: DUGLE, J.A. 1966. A taxonomic study
Page 58 and 59: accommodate findings related to mic
Page 60 and 61: Table l.--Mean ±S.E. area eaten by
Page 62 and 63: Sulfhydryl-Disulfide Mechanisms In
Page 64 and 65: Iraa prior experimental analysis of
Page 66 and 67: FRAZIER, J.L. and HEITZ, J.R. 1975.
Page 68 and 69: REICHARD, R 1993. From RNA to DNA,
Page 70 and 71: ; trees, and between and within ind
Page 72 and 73: Encapsulated objects are data struc
Page 74 and 75: _re3.--The Lignum model can be cont
Page 76 and 77: SEVERE DEFOLIATION Tree Fine Root +
Page 78 and 79: RUMBAUGH, J., BLAHA, M., PREMERLANI
Page 80 and 81: oo0 a b ol ¢' _achiman_i ()® 1,eA
Page 82 and 83: 1000 Conspicuous Defo i at ion 100
Page 84 and 85: 2.oo 2.oo o t 4 t ¢._ e'- 1.80 1.8
Page 86 and 87: Qualitative Changes in [,eaves and
Page 88 and 89: t00 2 Yr 1 Yr o_ 80 Treatment Treat
Page 90 and 91: Table 2.--Body size of Q. purzctate
Page 92 and 93: 06 b AddedBSA < 04 _ 2rag -'=' i:3
Page 94 and 95: Severe Defol iat ion Site-A 1993 _
Page 96 and 97: KAMATA, N. and IGARASHI, M. 1995b.
Page 98 and 99: enclosure %r 20 post-diapausing sec
Page 100 and 101: 2,500 F 000 F NF NF b _7 a _;_ a _7
Page 102 and 103: 8O D 09 i 4,'.) F AS4L a Z___7 2o !
Page 104 and 105: 2,500 _ st2-st4 El st5 0 st6 [_ pup
Page 108 and 109: later, we selected one more floweri
Page 110 and 111: Table 2.---Mean spruce budworm perf
Page 112 and 113: Table 3..--Mean spruce budworm perf
Page 114 and 115: FOLIVORE FEEDING ON MALE CONIFER FL
Page 116 and 117: Table 3.--Lymantria monacha prefere
Page 118 and 119: 500- 400- i 300 v ¢ m a 200- I00 L
Page 120 and 121: From the point of view of trees, th
Page 122 and 123: 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
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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
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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
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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
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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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