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DUGLE, J.A. 1966. A taxonomic study of Western Canadian species in the genus Betula. Canadian Journal of Botany 44: 929-1007. GILL, R.M.A. 1992a. A review of damage by mammals in north temperate forests: 1. Deer. Forestry 65(2): 145-169. GILL, R.M.A. 1992b. A review of damage by mammals in north temperate forests: 2. Small mammals. Forestry 65(3): 281-308. HERMS, D.A. and MATTSON, W.J. 1992. The dilemma of plants: to grow or defend. The Quarterly Review of Biology 67(3): 283-335. JOHNSSON, H. 1966. Avkommepr6vning av bj6rk. F6reningen Skokstrfidsf6r_idling, Arsbok 1966: 90-135. LEPPIK, E.E. 1970. Gene centers of plants as a source of disease resistance. Annual Review of Phytopathology 8: 323-344. MATTSON, W.J. 1980. Herbivory in relation to plant nitrogen content. Annual Review of Ecology and Systematics 11: 119-161. PAAKKONEN, E., PAASiSALO, S., ttOLOPAINEN, T., and K._RENLAMPI, L. Growth and stomatal responses of birch (Betula pendula Roth.) clones to ozone in open-air and chamber fumigations. New Phytologist 125: 615-623. POTERI, M. and ROUSI, M. 1996. Variation in Melampsoridium resistance among European white birch clones grown in different fertilizer treatments. Eurp. J. For. Pathol.: In Press. RHOADES, D.F. 1979. Evolution of plant chemical defense against herbivores, p. 3-54. In Rosenthal, G.A. and Janzen D.H., eds. Herbivores: their interaction with secondary plant metabolites. Academic Press, Orlando. ROUSI, M. 11988. Resistance breeding against voles in birch: possibilities for increasing resistance by provenance transfers. EPPO (European Plant Protection Organization) Bulletin 18: 257-263. ROUSI, M. 1990. Breeding forest trees for resistance to mammalian herbivores - a study based on European white birch. Acta Forestalia Fennica 210. ROUSI, M., TAHVANAINEN, J., and UOTILA, I. 1989. Inter- and intraspecific variation in the resistance of winterdormant birch (Betula spp.) against browsing by the mountain hare. Holarctic Ecology 12:187-192. ROUSI, M., HENTTONEN, H., and KAIKUSALO, A. 1990. Resistance of birch (Betula pendula and B. platyphylla) seedlots to vole. Scandinavian Journal of Forest Research 5: 427-436. ROUSI, M., TAHVANAINEN, J., and UOTILA, I. 1991. A mechanism of resistance to hare browsing in winter-dormant European white birch (Betula pendula). The American Naturalist 137: 64-82. ROUSI, M., TAHVANAINEN, J., and UOTILA, I. 1993. Effects of shading and fertilization on resistance of winterdormant birch (Betula pendula) to voles and hares. Ecology 74(1): 30-38. ROUSI, M., MATTSON, W.J., TAHVANAINEN, J., HENTTONEN, H., KOIKE, T., and UOTILA, I. 1996a. Growth and hare resistance of birches: testing defense theories. Oikos: In Press. ROUSI, M., TAHVANAINEN, J., HERMS, D.H, HENTTONEN, H., and UOTILA, I. 1996b. Clonal variation in susceptibility of white birches to herbivores. For. Sci.: Submitted. VAVILOV, N.I. 1922. The law of homologous series in variation. Journal of Genetics 12(1): 47-89. 45
A REDOX-BASED MECHANISM BY WHICH ENVIRONMENTAL STRESSES ELICIT CHANGES IN PLANT DEFENSIVE CHEMISTRY DALE M. NORRIS 642 Russell Laboratories, University of Wisconsin, Madison, WI 53706, USA INTRODUCTION It is now clear that most, if not all, lifeforms employ chemicals in communicating with their biotic and abiotic environments. Higher plants and animals, including insects and humans, may even use many of the same compounds (Rodriquez and Levin 1976, Norris and Liu 1992, Raina et aL 1992). Quinol-quinone redox couples, e.g., p-hydroquinone/ p-benzoquinone (Fig. 1), are prime examples of such common messengers. Hundreds of ions, free radicals (Packer and Glazer 1990) and molecules have been shown to be messengers. Alkaloids, flavonoids and terpenoids are families of compounds which are especially important as messengers (Norris 1986). 0 O- OH II II II 0 0 O- OH "A" "B" Figure l.--The classical redox couple in physical chemistry, p-hydroquinone / p-benzoquinone. The development of knowledge about the chemical senses has frequently been quite disconnected among many scientific disciplines. As a result, limited progress has been made in finding unifying principles. However, for several decades, I have developed hypotheses pertaining to such unifying principles and tested them experimentally (Norris 1994). The results of our and others' efforts have allowed me to propose that a common code for chemical communication exists among lifeforms, and them and their abiotic environments. The proposed name for this code is the Environmental Energy Exchange (EEE) Code (Norris 1981, 1986, 1988, 1994; Neupane and Norris 1992; Norris and Liu 1992). In this paper, the essential characteristics of this EEE code, including its underlying sulfhydryl (thiol, -SH) / disulfid_ (-S-S-) -dependent oxidative-reductive, energy-transducing mechanism, are applied to the interpretation of how environmental stresses elicit changes in plant defensive chemistry. Stress Alteration Of Phytoehemieal Messages The chemistry by which plants "communicate" with their herbivores has been classified as either constitutive (i.e., pre-stress) or inducible (i.e., post-stress). Plant pathologists and plant physiologists created this classification to Mattson, W.J., Niemel_i, P., and Rousi, M., eds. 1996. Dynamics of forest herbivory: quest for pattern and principle. USD_-X For. Serv. Gen. Tech. Rep. NC- 183, N.C. For. Exp. Sta., St. Paul, MN 55108. 46 !
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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
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
Page 54 and 55: However, hares showed strong prefer
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
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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
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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
Page 200 and 201:
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
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
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chemistry associated with root colo
Page 228 and 229:
Implications to Natural Resource Ma
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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
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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
Page 248 and 249:
One weakness of using data from gen
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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
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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
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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
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 .................................
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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
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
Page 292 and 293:
systems facilitate mapping of fores
Page 294 and 295:
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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