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HUNTER, M.D. and PRICE, P.W. 1992. Playing chutes and ladders: heterogeneity and the relative roles of bottom-up and top-down forces in natural communities. Ecology 73: 724-732. KAYA, H.K. t990. Soil ecology, p. 93-11 I. In Gaugler, R. and Kaya, H.K., eds. Entomopathogenic Nematodes in Biological Control. CRC Press, Boca Raton. LANE, G.A., SUTHERLAND, O.R.W., and SKIPP, R.A. 1987. Isoflavonoids as insect feeding deterrents and antifungal components from root of Lupinus augustijblius. J. Chem. Ecol. 13" 771-783. MARTIN, M.M. 1991. The evolution of cellulose digestion in insects. Phil. Trans. R. Soc. Lond. B. 333: 281-288. MCNAUGHTON, S.J., RUESS, R.W., and SEAGLE, S.W., 1988. Large mammals and process dynamics in African ecosystems. Bio Science 38 794-800. OKSANEN, L., FRETWELL, S.D., ARRUDA, J., and NIEMELA, P. 1981. Exploitation ecosystems in gradients of primary productivity. Am. Nat. 118" 240-261. OPLER, RA. 1968. Unusual numbers of Hepialus sequoiolus Behrens in Sonoma County. Pan. Pac. Entomol. 44: 83. PRICE, RW. 1991. The plant vigor hypothesis and herbivore attack. Oikos 62:244-251. PRICE P.W. 1992. Plant resources as the mechanistic basis for insect herbivore population dynamics, p. 139-173. In Hunter, M.D., Ohgushi, T., and Price, RW., eds. Effects of Resource Distribution on Animal-Plant Interactions. Academic Press, San Deigo. SINGER, M.C. and PARMESAN, C. 1993. Sources of variations in patterns of plant-insect association. Nature 361:251- 253. STRONG, D.R. 1992. Are trophic cascades all wet? Differentiation and donor-control in speciose ecosystems. Ecology 73: 747-754. STRONG, D.R., MARON, J.L., and CONNORS RG. 1996. Phmt competition, endogenous defenses, and natural enemies in terrestrial food webs: an example from the bush lupine community. In Polls, G. and Winemiller, K., eds. Food Webs: integration of patterns and dynamics. Chapman and Hall, NY. 472 p. STRONG, D.R., MARON, J.L., HARRISON, S., CONNORS, RG., and JEFFRIES, R.L. 1995. High mortality, fluctuation in numbers, and heavy subterranean insect herbivory in bush lupine, Lupinus arboreus, ms to be submitted to Oecologia. WAGNER, D.L. 1986. The biosystematics of Hepialus F.s. lato with special emphasis on the californicus-hectoides species group. PhD. Thesis in Entomology, University of California, Berkeley. WILLIAMS, F.X. 1905. Notes on the life history of Hepialus sequoiolus Behrens. Entomol. News. 16: 283-287. WHITE, T.C.R. 1993. The Inadequate Environment: Nitrogen and the Abundance of Animals. Springer-Verlag, New York. WINK, M. 1992. The role of quinolizidine alkaloids in plant-insect interactions, p. 131-166. In Bernays, E., ed. Plant-Insect Interactions. vol. IV. CRC Press, Boca Raton, FI. WINK, M. and WITTE, L. 1984. Turnover and transport of quinolizidine alkaloids. Diurnal fluctuations of lupanine in the phloem sap, leaves, and fruits ofLupinus albus L. Planta 161: 519-524. 131
RESISTANCE OF HYBRID AND PARENTAL WILLOWS TO HERBIVORES: HYPOTHESES AND VARIABLE HERBIVORE RESPONSES OVER 3 YEARS ROBERT S. FRITZ l, BERNADETTE M. ROCHE I, and COLIN M. ORIANS 2 _Department of Biology, Vassar College, Poughkeepsie, New York 12601, USA 2Department of Biology, Williams College, Williamstown, Massachusetts 01267, USA INTRODUCTION Analysis of herbivory on hybrid plants in natural populations has recently gained attention (Whitham 1989, Keim et al. 1989, Boecklen and Spellenberg 1990, Paige et al. 1991, Whitham et al. 1991, Aguilar and Boecklen 1992, Fritz et al. 1994, Whitham et al. 1994), and it may be important for understanding plant-herbivore interactions (Whitham 1989). To assess the significance of interspecific hybridization of plants for plant-herbivore interactions, it will be important to know how interspecific hybridization of plants influences resistance of hybrid plants in relation to resistance of pure parents. It has been hypothesized that hybrid plants may act as an "ecological sink" for herbivorous insects (Whitham 1989). In mixed stands of parental and hybrid plants, this hypothesis predicts that a large fraction of the herbivore population would reside on hybrid individuals (Drake 1981, Whitham 1989, Floate et al. 1993). Whitham (1989) referred to highly susceptible hybrid poplars as being an ecological sink for a galling aphid, but in terms of metapopulation dynamics, hybrid plants could function as a "source" rather than a sink (e.g., Ericson et al. 1993). Hybrid plants have also been hypothesized to act as "evolutionary sinks" to pests (Whitham 1989). The "evolutionary sink" idea predicts that if herbivore fitness was higher on hybrids compared to their fitness on parental plants, there would be selection for specialization on the hybrids, perhaps preventing selection tbr herbivore virulence on the parental species. If so, then hybrid plant-herbivore interactions may be critical to understanding why plants seem to maintain pest resistance in the face of a greater evolutionary potential for virulence in pests. Finally, Floate and Whitham (1993) suggest that hybrid plants may facilitate host shifts of herbivores onto new plants, because they could act as a selective "bridge" favoring adaptation of an herbivore to previously non-host species. Tests of these hypotheses will require knowing the general patterns of herbivore responses to hybrid versus pure parental species. Hypotheses Fritz et al. (1994) proposed several alternative hypotheses of phytophage response to hybrid plants. Those hypotheses assume that pure parent species are being compared to F1 hybrids. The reason to focus on F1 hybrids is that this comparison could suggest: (1) how resistance mechanisms are inherited from parental species, and (2) how these traits may be expressed upon backcrossing. The Additive hypothesis (Fig. 1A) predicts that hybrids do not differ from the mean of the resistances of the two parents (i.e., the midparent value). Thus, F1 hybrids are intermediate between the resistances of the parental species, which suggests that hybrid resistances are due to the additive inheritance of resistance traits from both parents. The second hypothesis is the Dominance hypothesis (Fig. 1B). If hybrid resistance differs significantly from the mean resistance of both parents but does not differ significantly from that of one of the parents, the Dominance hypothesis would be supported. Hybrid resistance could resemble that of either the more resistant or the more susceptible parent. (Dominance in this context refers to phenotypic similarity between a parent and hybrids, not necessarily genetic dominance, though it may imply dominant inheritance of resistance traits.) Herbivore densities could be intermediate between the Additive and Dominance patterns, which would support an hypothesis of partial dominance. The third hypothesis is the Hybrid Susceptibility hypothesis (the Hybrids-as-Sinks hypothesis of Whitham 1989; see also Keim et al. 1989, Boecklen and Spellenberg 1990). This hypothesis predicts higher herbivore densities and/or higher herbivore pertbrmance on hybrids compared to parental taxa (Fig. 1C). The Hybrid Susceptibility hypothesis can be distinguished from the Dominance hypothesis in that the susceptibility of the hybrid must exceed that of the most susceptible parent. Mattson, W.J., Niemelfi, 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. 132
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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
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 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 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
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
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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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