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FLOATE, K.D. and WHITHAM, T.G. 1993. The "hybrid bridge" hypothesis: host shifting via plant hybrid swarms. Am. Nat. 141 : 651-662. FRITZ, R.S., NICHOLS-ORIANS, C.M., and BRUNSFELD, S.J. 1994. Interspecific hybridization of plants and resistance to herbivores: Hypotheses, genetics, and variable responses in a diverse community. Oecologia 97:106-117. HALL, R.W. and TOWNSEND, A.M. 1987. Suitability of Ulmus wilsoniana, the 'Urban' elm, and their hybrids for the ehn leaf beetle, Xa_lthogaleruca luteola (Mtiller) (Coleoptera: Chrysomelidae). Environ. Entomol. 16: 1042-1044. HANHIM_.KI, S., SENN, J., and HAUKIOJA, E. 1994. Performance of insect herbivores on hybridising trees: the case of the subarctic birches. J. Anita. Ecol. in press. HUESING, J., JONES, D., DEVEERNA, J., MYERS, J., COLLINS, G., SEVERSON, R., and SISSON, V. 1989. Biochemical investigations of antibiosis material in leaf exudate of wild Nicotiana species and interspecific hybrids. J. Chem. Ecol. 15: 1203-1217. KEIM, R, PAIGE, K.N., WHITHAM, T.G., and LARK, K.G. 1989. Genetic analysis of an interspecific swarm of Populus: Occurrence of unidirectional introgression. Genetics 123: 557-565. LEVY, A. and MILO, J. 1991. Inheritance of morphological and chemical characters in interspecific hybrids between Pat)aver bracteatum and Papaverpseudo-orientale. Theor. Appl. Genet. 81: 537-540. MEIER, B., BETTSCHART, A., SHAO, Y., and LAUTENSCHLAGER, E. 1989. Einsatz der modernen HPLC for chemotaxonomische Utersuchungen morphologisch schwer zu differenzierender Salix-Hybriden. Planta Medica 55: 213-214. MOSSELER, A. and PAPADOPOL, C.S. 1989. Seasonal isolation as a reproductive barrier among sympatric Salix species. Can. J. Bot. 67: 2563-2570. O'DONOUGHUE, L.S., RAELSON, J.V., and GRANT, W.E 1990. A morphological study of interspecific hybrids in the genus Lotus (Fabaceae). Can. J. Bot. 68: 803-812. PAIGE, K.N., CAPMAN, W.C., and JENNETTEN, R 1991. Mitochondrial inheritance patterns across a cottonwood hybrid zone: Cytonuclear disequilibria and hybrid zone dynamics. Evolution 45: 1360-1369. PAIGE, K.N. and CAPMAN, W.C. 1993. The effects of host-plant genotype, hybridization and environment on gall aphid attack and survival in cottonwood: The importance of genetic studies and the utility of RFLP's. Evolution 47: 36-45. RICE, W.R. 1989. Analyzing tables of statistical tests. Evolution 43: 223-225. SAS INSTITUTE. 1985. SAS user's guide: Statistics, Version 5 Edition. SAS Institute, Inc. Cary, NC. SOETENS, R, ROWELL-RAHIER, M., and PASTEELS, J.M. 1991. Influence of phenolglucosides and trichome density on the distribution of insects herbivores on willows. Entomol. Exp. Appl. 59: 175-187. WHITHAM, T.G. 1989. Plant hybrid zones as sinks for pests. Science 244: 1490-1493. WHITHAM, T.G., MORROW, RA., and POTTS, B.M. 1991. Conservation of hybrid plants. Science 254: 779-780. WHITHAM, T.G., MORROW, RA., and POTTS, B M. 1994. Plant hybrid zones as centers of biodiversity: The herbivore community of two endemic Tasmanian eucalypts. Oecologia, in press. 141
F1 HYBRID SPRUCES INHERIT THE PHYTOPHAGOUS INSECTS OF THEIR PARENTS WILLIAM J. MATTSON, ROBERT K. LAWRENCE, and BRUCE A° BIRR USDA Forest Service, North Central Forest Experiment Station B-7 Pesticide Research Center, Michigan State University, East Lansing, MI 48824, USA INTRODUCTION Interspecific hybridization in plants is a widespread and important natural phenomenon. In fact, Stace (1987) estimates that at least 50% of the extant angiosperm species have been derived via hybridization. Natural hybrids are also common among the gymnosperms (Wright 1955, Hanover and Wilkinson 1969, Schmidt-Vogt 1977, Zobel and Talbert 1984). In spite of the ecological and evolutionary importance of plant hybrid zones (Remington 1968, Barton and Hewitt 1985), little is yet known about how they affect the dynamics of plant/herbivore relationships. Whitham (1989) speculated that hybrids may be particularly prone to herbivory because usual resistance mechanisms derived from the parents are poorly expressed. For example, polygenic resistance traits might be intermediate in F1 hybrids, whereas monogenic and oligogenic traits might be fully expressed or not at all depending on the mode of gene action (e.g., dominant or recessive, etc.), and on whether there is genetic dysfunction preventing normal gene expression (Fritz et al. 1994, Whitham et al. 1994). Furthermore, to compound these problems, hybrids may attract specialist phytophages from both parent species, thereby being predisposed to a much larger pool of potential consumers than either of the pure parent types (Whitham et al. 1994). Insect Loading in Relation to a Plant's A/R Vectors and S/D Vectors To examine the issue of hybrid resistance to insects in a more mechanistic framework, we consider that there are at least two sets of plant traits affecting insect loading on plants: (1) the mix of properties which elicit insect attraction/ arrestmenffrepulsion (e.g., morphology, electromagnetic and biochemical cues, abiotic requirements, etc.) and thus govern the number of insect species that attempt to colonize it, and (2) the mix of properties that are sustaining/defensive for each of the would be colonizers as they attempt to feed, oviposit, and reproduce. These two sets of properties, which together set the potential for insect loading (total insect biomass or numbers), can be represented as two vectors: the species attraction/ repulsion (A/R) vector, [a_,a2,a3..an],and the suitability/defense (S/D) vector [Sl,S2,S3..s]. Each species of insect, Si, is represented by an element (a)in the species A/R vector, having a numerical value (0-1) that is proportional to the plant's average capacity to attract and arrest individuals of the particular insect species. Likewise, the S/D vector contains specific elements (s) that correspond to and represent the plant's average suitability, ranging from zero, where an insect colonizes a plant but cannot complete development on it (immunity), to 1 where the plant fulfills all of the insect's requirements, for each species in the species A/R vector. The product of the two vectors, [ayL+a2s2+..as ], represents the plant's overall potential "resistance expression", or potential insect loading per plant. To give an extreme example for one insect: the white pine weevil, Pissodes strobi, is highly attracted to Picea glauca shoots as evidenced by the great number of feeding scars it produces (i.e., a = 1), but it can only rarely reproduce in these shoots (i.e., s = 0), with the consequence that P glauca is considered "resistant" to the weevil (Lanier 1983). Actual insect loading depends, in addition, on the quantity and genetic diversity of insects (N) that continually attempt colonization. The total number of non-zero elements (all those a_> 0) in an individual plant's A/R or species vector (its species richness or species loading) is positively linked to the size and density of its population, the total areal distribution of the species, and negatively to its biochemical uniqueness wherein it occurs (for a review, see Tahvanainen and Niemela 1987). For many natural hybrids, their populations and geographical distributions are usually small (compared to their parents) thereby exposing them to fewer potential colonists, but their low biochemical uniqueness or high similarity to the two nearby parent species will likely have a counteracting influence making them highly susceptible to the reservoirs of consumers fiom two plant species (Whitham et al. 1994). Mattson, W.J., Niemel_, R, 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. 142 ;fl ¸_
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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
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 106 and 107: 96 SLANSKY, E, Jr. 1990. Insect nut
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 152 and 153: Elements of the Suitability/Defense
Page 154 and 155: 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
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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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