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1.
Christiansen E Krokene P Berryman AA Franceschi VR Krekling T Lieutier F Lönneborg A Solheim H 《Tree physiology》1999,19(6):399-403
Norway spruce trees (Picea abies (L.) Karst.) pretreated by wounding and fungal infection showed highly enhanced resistance to a subsequent challenge inoculation with the pathogenic bluestain fungus Ceratocystis polonica (Siem.) C. Moreau. This is the first time the effectiveness of the constitutive and inducible defenses has been shown to depend on prior wounding and infection in conifers, although such acquired resistance has previously been found in several angiosperms. Trees that were pretreated with a combination of 12 bark wounds (1.6 x 10 cm), four fungal inoculations and four sterile inoculations 1-15 days before mass inoculation with C. polonica at 400 inoculations per square meter over a 0.8 m stem section had significantly shorter necroses in the phloem, less bluestained sapwood, and less dead cambium than untreated control trees. Pretreatment with four fungal or sterile inoculations alone did not lead to enhanced resistance. Pretreatment by bark wounding alone seemed to provide an intermediate degree of resistance compared to bark wounding, fungal inoculations and sterile inoculations combined. All trees had a marked increase in the number of resin ducts in the year of inoculation compared with previous years, suggesting that formation of traumatic resin ducts play an important role in the development and maintainance of enhanced resistance. 相似文献
2.
To study the effect of chemical pretreatment on conifer resistance, 13-year-old Norway spruce (Picea abies (L.) Karst.) trees were treated with methyl jasmonate (MJ) or oxalic acid (OxA) on the outer bark and inoculated with the pathogenic blue-stain fungus Ceratocystis polonica (Siem.) C. Moreau 4 weeks later. Both chemicals significantly reduced symptoms of fungal infection, but MJ was more effective than OxA (51 versus 18% reduction in length of necrotic lesions in the phloem relative to untreated control trees). Anatomical examination of treated stem tissues showed that MJ induced extensive formation of traumatic resin ducts in the xylem and extra polyphenolic parenchyma (PP) cells in the secondary phloem between the cambium and the regular annual PP cell layer. No traumatic resin ducts were formed after treatment with OxA, and the coverage of extra PP cells in OxA-treated tissues was not significantly higher than in the controls. The anatomically based defense reactions induced by MJ were similar to the reactions observed after pathogen infection, mechanical wounding and bark beetle attack. Neither MJ nor OxA had apparent phytotoxic effects on Norway spruce at the concentrations used, with needle and stem tissues of all trees appearing normal without visible symptoms of toxicity. However, trees treated with MJ had 30% less radial sapwood growth than control trees. In conclusion, MJ treatment of Norway spruce appears to have practical potential as a tool for increasing plant resistance to fungal infection, but with a modest reduction in sapwood growth. 相似文献
3.
Villari C Battisti A Chakraborty S Michelozzi M Bonello P Faccoli M 《Tree physiology》2012,32(7):867-879
Conifer bark beetles are often associated with fungal complexes whose components have different ecological roles. Some associated species are nutritionally obligate fungi, serving as nourishment to the larvae, whereas others are pathogenic blue-stain fungi known to be involved in the interaction with host defenses. In this study we characterized the local and systemic defense responses of Scots pine (Pinus sylvestris L.) against Ophiostoma brunneo-ciliatum Math. (a blue-stain pathogen) and Hyalorhinocladiella macrospora (Franke-Grosm.) Harr. (a nutritional fungus). These fungi are the principal associates of the pine engraver beetle, Ips acuminatus (Gyll.). Host responses were studied following inoculation with the fungi, singly and as a fungal complex, and by identifying and quantifying terpenoids, phenolic compounds and lignin. Although the length of the necrotic lesions differed between control (wound) and fungal treatments, only two compounds (pinosylvin monomethyl ether and (+)-α-pinene) were significantly affected by the presence of the fungi, indicating that Scots pine has a generic, rather than specific, induced response. The fact that both nutritional and blue-stain fungi triggered comparable induced defense responses suggests that even a non-pathogenic fungus may participate in exhausting host plant defenses, indirectly assisting in the beetle establishment process. Our findings contribute to the further development of current theory on the role of associated fungal complexes in bark beetle ecology. 相似文献
4.
Conifers have defenses such as the production of phenolic compounds and resins that can be induced by bark beetles and other invading organisms, but the signaling agents involved are unknown. The anatomical effects of methyl jasmonate (MJ), a potent inducer of certain plant defenses, were compared with wounding of the bark of 12-15-year-old trees of five conifer species. Wounding in all species resulted in tissue necrosis and wound periderm development immediately around the wound site. One cm from the wound, swelling of phloem polyphenolic parenchyma cells and phenolic accumulation were observed in Pseudotsuga menziesii (Mirb.) Franco, Picea pungens Engelman, Larix occidentalis Nutt. and Pinus monticola Douglas ex D. Don, but not in Taxus brevifolia Nutt. Traumatic resin ducts were formed in response to wounding in three species of Pinaceae, but not in P. monticola, which formed irregular clusters of cells rather than ducts. Taxus brevifolia did not form resin ducts in response to either wounding or MJ treatment. In the Pinaceae species studied, surface application of 100 mM MJ caused similar anatomical changes to those observed in response to wounding, including phenolic accumulation, cell swelling and traumatic resin duct formation, but it did not induce a wound periderm. Traumatic resin ducts differed in size among the study species, ranging from small in L. occidentalis to very large in P. menziesii. In P. menziesii, P. pungens and L. occidentalis, traumatic resin ducts were more abundant after MJ treatment than after wounding. We conclude that the octadecanoid pathway is likely involved in defense responses in stems of the Pinaceae, but not necessarily in other taxa. 相似文献
5.
Tree defense against bark beetles (Curculionidae: Scolytinae) and their associated fungi generally comprises some combination of constitutive (primary) and induced (secondary) defenses. In pines, the primary constitutive defense against bark beetles consists of preformed resin stored in resin ducts. Induced defenses at the wound site (point of beetle entry) in pines may consist of an increase in resin flow and necrotic lesion formation. The quantity and quality of both induced and constitutive defenses can vary by species and season. The inducible defense response in ponderosa pine is not well understood. Our study examined the inducible defense response in ponderosa pine using traumatic mechanical wounding, and wounding with and without fungal inoculations with two different bark beetle-associated fungi (Ophiostoma minus and Grosmannia clavigera). Resin flow did not significantly increase in response to any treatment. In addition, necrotic lesion formation on the bole after fungal inoculation was minimal. Stand thinning, which has been shown to increase water availability, had no, or inconsistent, effects on inducible tree defense. Our results suggest that ponderosa pine bole defense against bark beetles and their associated fungi is primarily constitutive and not induced. 相似文献
6.
Importance of current photosynthate in the regulation of tree defense against the southern pine beetle (SPB),Dendroctonus frontalis Zimm. (Coleoptera: Scolytidae) was investigated in loblolly pine,Pinus taeda L. Downward translocation of carbohydrate was blocked by removing a ring of bark (girdle) at 3.5 m above ground. Beetle colonization success and tree defensive responses measured as resin flow and induced lesion formation to inoculations with the beetle-associated fungus,Ophiostoma minus (Hedge.) H. and P. Sydow., were assessed above and below girdles. We hypothesized that a decrease in a available carbohydrate would lead to a decrease in tree defense and an increase in SPB colonization success below girdles. At 2 weeks, post-girdling carbohydrates were already reduced by 50% below girdles, but no significant differences in beetle colonization success or defensive responses were measured. At 8 weeks, post-girdling starch was reduced by 93%, sucrose by 44% and resin flow by 44% below girdles. Southern pine beetle adults were more successful in colonizing areas below than above girdles. However, no significant differences were detected in the size of the lesions that formed in response to inoculated fungi. This suggests that the size of the induced lesion may not be a reliable indicator of tree resistance to bark beetle attack as previously believed, and that lesion development is not entirely dependent on available carbohydrate or related to tree defense against beetles. Cambial growth was reduced below girdles with no latewood formed below and 25 rows of latewood formed above girdles. Latewood contains much higher densities of resin ducts than earlywood. This may explain why resin flow was lower and beetles were more successful in colonizing areas of the tree below girdles. Therefore, tree defense against bark beetles may be more complex than a simple supply-and-demand relationship for carbohydrate, and changes in source-sink relationships, as influenced by the environment, may be more important than supply. 相似文献
7.
Resin flow is the primary means of natural defense against southern pine beetle (Dendroctonus frontalis Zimm.), the most important insect pest of Pinus spp. in the southern United States. As a result, factors affecting resin flow are of interest to researchers and forest managers. We examined the influence of fertilization, artificial wounding and fungal inoculation on resin flow in 6- and 12-year-old stands of loblolly pine (Pinus taeda L.) and determined the extent of that influence within and above the wounded stem area and through time. Fertilization increased constitutive resin flow, but only the younger trees sustained increased resin flow after wounding and inoculation treatments. An induced resin flow response occurred between 1 and 30 days after wounding and inoculation treatments. Wounding with inoculation resulted in greater resin flow than wounding alone, but increasing amounts of inoculum did not increase resin flow. Increased resin flow (relative to controls) lasted for at least 90 days after wounding and inoculation. This increase appeared to be limited to the area of treatment, at least in younger trees. The long-lasting effects of fungal inoculation on resin flow, as well as the response to fertilization, suggest that acquired resistance through induced resin flow aids in decreasing susceptibility of loblolly pine to southern pine beetle. 相似文献
8.
Drill wounds in balsam fir and hemlock roots activated the nonspecific resistance mechanisms of compartmentalization in wood and necrophylactic periderm in bark. Tangential bands of resin ducts localized around the wounds constituted the barrier zones in the secondary xylem of conifer roots. Barrier zones were more extensive in roots which showed symptoms characteristic of invasion by fungi and bacteria after wounding. This observation supports an expanded definition of barrier zones; barrier zones may form not only in response to mechanical wounds but also in response to xylem injury caused by pathogens. Multiple bands of resin ducts were common in young xylem when bark lesions developed around wounds. Necrophylactic periderms isolared dead bark from living bark. Development of phellem cells with dark contents and thick suberized walls, typical of exophylactic periderm, followed initiation of necrophylactic periderm. The wound responses were similar in both balsam fir and hemlock. 相似文献
9.
《Scandinavian Journal of Forest Research》2012,27(1-4):300-306
Bark stripping by moose (Alces alces L.) on young Scots pine (Pinus sylvestris L.) was studied at the Grimsö Wildlife Research Area in south‐central Sweden. The investigation was launched to attempt to elucidate the underlying mechanism(s) for this behaviour. To document the seasonal pattern of bark stripping by moose, four young Scots pine plantations were observed between August 1988 and November 1989. In addition, pine bark samples were collected systematically throughout the study period, and the composition of various nutrients, minerals, and fibre fractions and their digestibility were determined. The occurrence of bark stripping was significantly higher in April and May, coinciding with an increase in fibre, K, starch and total carbohydrate concentrations, and decreasing crude protein, P and Mg content. Results are discussed with respect to the potential relationship between bark stripping and nutrient deficiencies in moose and the potential mechanisms inducing bark stripping. 相似文献
10.
Norway spruce (Picea abies (L.) Karst.) trees were preinoculated with a sublethal dose of the blue-stain fungus Ceratocystis polonica Siem. (C. Moreau) 1 to 52 weeks before they were mass inoculated with the same fungus. Trees pretreated 1 week before mass inoculation had similar, severe symptoms of fungal infection as the control trees. Pretreatment 3, 6 or 9 weeks before mass inoculation resulted in effective protection of the trees, reducing pathogenic symptoms by 63-90% relative to the control trees, whereas pretreatment 52 weeks before mass inoculation gave intermediate protection (44-71% reduction in symptoms). Thus, pretreatment induced resistance to the blue-stain fungus in Norway spruce by a process that requires more than 1 week to become activated and protects trees for at least one year after pretreatment. Pretreatment induced formation of traumatic resin ducts (TDs) in the sapwood and swelling and proliferation of polyphenolic parenchyma cells (PP cells) in the phloem. Trees pretreated 3-9 weeks before mass inoculation had more TDs and showed greater swelling of existing PP cells than control trees or trees pretreated 1 week before mass inoculation. We conclude that induced disease resistance in Norway spruce is probably associated with PP cell activation and TD induction, because resistance was enhanced within the same time frame as the induction of these defense responses. 相似文献
11.
Oleoresin flow is an important factor in the resistance of pines to attack by southern pine beetle, Dendroctonus frontalis Zimm., and its associated fungi. Abiotic factors, such as nutrient supply and water relations, have the potential to modify this plant-insect-fungus interaction; however, little is known of the effects of inoculation with beetle-associated fungi on oleoresin flow. We observed that constitutive and induced resin yield in loblolly pine, Pinus taeda L., were affected by either fungal inoculation (with the southern pine beetle-associated fungus Ophiostoma minus (Hedgcock) H. & P. Sydow) or silvicultural treatment. The effects of mass wounding (400 wounds m(-2)) and mass wounding and inoculation with O. minus were assessed by comparison with untreated (control) trees. The treatments were applied to trees in a 2 x 2 factorial combination of fertilizer and irrigation treatments. Fertilization did not significantly affect constitutive resin yield. Even as long as 105 days post-treatment, however, mass-inoculated trees produced higher induced resin yields than control or wounded-only trees, indicating a localized induced response to fungal inoculation. We noted no systemic induction of host defenses against fungal colonization. Although beetles attacking previously attacked trees face a greater resinous response from their host than beetles attacking trees that had not been previously attacked, the effect of an earlier attack may not last more than one flight season. Despite mass inoculations, O. minus did not kill the host trees, suggesting that this fungus is not a virulent plant pathogen. 相似文献
12.
Two greenhouse experiments were conducted to study intraspecific variation in growth of the root rot fungus Heterobasidion annosum in living host sapwood. In experiment 1, Scots pine (Pinus sylvestris) and Norway spruce (Picea abies) seedlings were inoculated with H. annosum isolates, 14 each of the S-and P-intersterility groups, collected from various parts of Sweden. In pine, the P-group isolates were more virulent than the S-group isolates both in terms of infection frequency, induced mortality rate (p < 0.05), and fungal growth in sapwood (p < 0.05). In spruce, the P-group isolates were also more virulent on average, but the difference was not statistically significant. Both S and P isolates had a higher infection frequency and a significantly longer sapwood growth on spruce than on pine. The P-group caused higher mortality on pine than on spruce. The length of the lesion in the inner bark was strongly correlated with fungal growth in spruce, but not in pine where the lesions were short or absent. In experiment 2, ten Norway spruce clones were inoculated with 18 S-isolates, originating from nine live-decayed trees and from nine spore-infected stumps in a single Norway spruce stand. The objective was to test whether any selection for growth rate in sapwood was detectable among individuals of H. annosum originating either from stumps or trees. The results gave no support for such selection since no difference in sapwood growth between the two groups of isolates was found. 相似文献
13.
《Scandinavian Journal of Forest Research》2012,27(2):158-165
The short-term effect of thinning on cronartium rust on Scots pine ( Pinus sylvestris L.) was investigated in a stand of Scots pine (artificially seeded and in the pole-stage) severely infected by the stem-rust fungus Cronartium flaccidum (Alb. & Schwein) G. Winter. The disease rate (the number of old and fresh infections) was recorded before thinning, after which all trees with a resin-top as well as all individual branches with sporulating lesions were removed from the thinned plots, while the comparison plots remained non-thinned. The numbers of sporulating lesions were recorded on the plots annually for 2-5 yrs after thinning. Thinning had no significant short-term effect on the year-to-year relative number of sporulating lesions. The relative number of sporulating lesions, however, increased in both thinned and nonthinned trees, varying significantly from year to year. Thinning had no significant effect on the year of formation of the shoot bearing annual sporulating lesions, the duration of sporulation or the growth of the fungus along the infected shoot in perennial lesions during the first 5 yrs after the treatment. 相似文献
14.
Forty-one 2-year-old clones of Picea sitchensis (Bong.) Carr. from three full-sib families (14 clones from each of two families and 13 clones from a third family) were either wounded and inoculated with an isolate of Heterobasidion annosum (Fr.) Bref. or wounded without inoculation. Lesion lengths on the inner bark from the point of inoculation varied among clones 35 days after treatment. There was no relationship between lesion length and relatedness within families. Two clones (21342 and 25202) with the shortest lesions, tentatively designated as less susceptible to H. annosum, and two clones (21176 and 27166) with the longest lesions, designated more susceptible, were selected for comparison of host anatomical and chemical responses to infection. The position and structure of the ligno-suberized boundary zone (LSZ) in the bark of the clones suggested that the less susceptible clones formed thicker layers of suberized cells in the LSZ following wounding plus inoculation. No LSZ was observed in two ramets of the more susceptible Clone 27166 following wounding and inoculation with H. annosum. Compared with more susceptible genotypes, clones of P. sitchensis with low susceptibility to H. annosum had high relative proportions of (+)-alpha-pinene, (-)-beta-pinene and one unidentified terpene constituent (Unknown-15) in cortical resin sampled 25 cm from the lesions. In contrast, more susceptible clones had higher relative proportions of (-)-limonene, Unknown-16, Unknown-18 and Unknown-19. In the secondary resin produced in bark tissues surrounding the lesions, proportions of several monoterpene constituents varied; these changes included a decrease in the relative amount of beta-phellandrene and corresponding small increases in some minor constituents. The concentrations of the monoterpenes, except a few minor constituents, increased in the infected tissues. Wounding plus inoculation with H. annosum resulted in varied monoterpene responses, with distinct differences between less susceptible and more susceptible clones. In less susceptible clones, Unknown-19 increased following wounding plus inoculation, whereas in more susceptible clones, concentrations of delta-3-carene and Unknown-13 and Unknown-16 increased. Differences in both constitutive and induced resin monoterpene profiles may provide useful markers for resistance to H. annosum in selection and breeding programs. 相似文献
15.
Anatomical and cellular responses of Pinus monticola stem tissues to invasion by Cronartium ribicola
J. W. Hudgins G. I. McDonald P. J. Zambino N. B. Klopfenstein V. R. Franceschi 《Forest Pathology》2005,35(6):423-443
White pine blister rust (Cronartium ribicola) causes extensive damage to white pines and their associated ecosystems across North America. The anatomical and cellular characteristics of C. ribicola colonization in Pinus monticola branch and stem tissues were studied as a basis for understanding host tree reactions that may be related to resistance. Samples examined showed typical fusiform swelling and some had produced aecia. The reaction of phloem and xylem tissues was compared with non-infected tissue using light and electron microscopy. Cortical parenchyma and phloem polyphenolic parenchyma cells underwent mitotic division, cell swelling, and ca sixfold greater accumulation of phenolic compounds in colonized vs. control stems. In the cortex and secondary phloem, haustoria were common in parenchymatous cells, and hyphae were abundant in the intercellular spaces, but cell death was rare, unless aecia had ruptured the stem cortex. Hyphae were also common in xylem rays, tracheids and between tracheids. Disease-induced changes in the cambial zone included development of cambium-derived xylem traumatic resin ducts. Results demonstrate that diverse host defence responses were activated in the bark of apparently susceptible trees, but lack of mechanical damage by C. ribicola to the phenol-containing host cells and the resin duct system allowed extensive colonization and development of aecia despite elicitation of these stem defences. Interactions between P. monticola and C. ribicola are discussed and compared with other conifer–fungus pathosystems. 相似文献
16.
17.
When conifers such as Picea abies Karst. (Norway spruce) are attacked by insects or pathogens, they often respond by producing increased quantities of terpenoid oleoresin. This response can be mimicked in young P. abies seedlings by treatment with methyl jasmonate (MJ). In this study, we determined the effects of MJ on terpenoids and other chemical defenses of mature P. abies, and investigated whether this treatment protected trees against attack by the blue-stain fungus Ceratocystis polonica (Siem.) C. Moreau, the most important fungal associate of the spruce bark beetle Ips typographus L. Methyl jasmonate treatment induced the formation of traumatic resin ducts in the developing xylem, enhanced resin flow and stimulated increased accumulation of monoterpenes, sesquiterpenes and diterpene resin acids. However, only minor changes were detected in terpene composition in response to MJ treatment and no changes in soluble phenolic concentration were measured. There was much variability in the timing and degree of response to MJ among clones. The observed chemical and anatomical changes in response to MJ treatment were correlated with increased resistance to C. polonica, suggesting that terpenoid oleoresin may function in defense against this pathogen. 相似文献
18.
We examined drought-induced changes in susceptibility of potted Scots pine (Pinus sylvestris L.) trees to a bark-beetle associated fungus (Leptographium wingfieldii Morelet, from the bark beetle Tomicus piniperda L.). Five-year-old field-grown trees were transplanted to 50-l pots and grown for 1 year before the treatments were applied. Trees in the drought treatment were subjected to several successive, 3-week-long drought cycles, with predawn water potential dropping below -2 MPa at peak drought intensity. The experimental drought cycles were more severe than the natural drought episodes usually recorded in Scots pine stands. Trees were then mass-inoculated with L. wingfieldii at a density close to the critical threshold density of inoculations (400 m(-2)) above which tree resistance is overcome. Inoculation of well-watered trees resulted in induced reaction zones around the inoculation points and very limited damage (resinosis) in the sapwood. Drought alone had no long-lasting consequences on tree water relations, except for a decrease in hydraulic conductance in the youngest segments of the main stem. However, the combination of mass-inoculation and drought stress after inoculation resulted in a dramatic loss of stem hydraulic conductivity that was paralleled by conspicuous damage to the sapwood (resinosis, drying and blue staining). There was a close correlation between amount of visible sapwood damage and loss of hydraulic conductivity. The intensity of induced reactions in the phloem was unaffected by drought stress. We conclude that tree defence against L. wingfieldii is decreased during severe drought stress, resulting in changes in the spread and action of the fungus in the sapwood but not in the phloem. 相似文献
19.
Induced (traumatic) resin in white spruce (Picea glauca (Moench) Voss) leaders resistant or susceptible to the white pine weevil (Pissodes strobi Peck) was analyzed for volatile terpenes and diterpene resin acids after simulated white pine weevil damage. Leaders from 331 trees were wounded just below the apical bud with a 1-mm diameter drill, coinciding with the natural time of weevil oviposition in the spring. Leaders were removed in the fall, and the bark and xylem from the upper and lower regions of the leader extracted and analyzed by gas chromatography. Unwounded trees had low amounts of resin in xylem compared with bark. In response to wounding, volatile terpenes and diterpene resin acids increased in the upper xylem (area of wounding), with resistant trees showing a greater increase than susceptible trees. Wounding caused monoterpenes in particular to decrease in the lower region of the leader (away from the drilled area) in greater amounts in susceptible trees than in resistant trees. In response to wounding, the proportion of monoterpene to resin acid increased in the upper and lower xylem of resistant trees, and slightly increased in the upper xylem of susceptible trees. Monoterpene-enriched resin is more fluid than constitutive resin, and probably flows more readily into oviposition cavities and larval mines, where it may kill immature weevils. Loss of resin components in the lower xylem suggested catabolism and transport of these materials to the site of wounding; however, energetic and regulatory data are necessary to confirm this hypothesis. This study provides a basis for measuring the ability of a tree to undergo traumatic resinosis that could be used to screen for resistance to white pine weevil. 相似文献
20.
Coniferous trees have both constitutive and inducible defences that deter or kill herbivores and pathogens. We investigated constitutive and induced monoterpene responses of jack pine (Pinus banksiana Lamb.) to a number of damage types: a fungal associate of the mountain pine beetle (Dendroctonus ponderosae Hopkins), Grosmannia clavigera (Robinson-Jeffrey & R.W. Davidson); two phytohormones, methyl jasmonate (MJ) and methyl salicylate (MS); simulated herbivory; and mechanical wounding. We only included the fungal, MJ and mechanical wounding treatments in the field experiments while all treatments were part of the greenhouse studies. We focused on both constitutive and induced responses between juvenile and mature jack pine trees and differences in defences between phloem and needles. We found that phytohormone applications and fungal inoculation resulted in the greatest increase in monoterpenes in both juvenile and mature trees. Additionally, damage types differentially affected the proportions of individual monoterpenes: MJ-treated mature trees had higher myrcene and β-pinene than fungal-inoculated mature trees, while needles of juveniles inoculated with the fungus contained higher limonene than MJ- or MS-treated juveniles. Although the constitutive monoterpenes were higher in the phloem of juveniles than mature jack pine trees, the phloem of mature trees had a much higher magnitude of induction. Further, induced monoterpene concentrations in juveniles were higher in phloem than in needles. There was no difference in monoterpene concentration between phytohormone applications and G. clavigera inoculation in mature trees, while in juvenile trees MJ was different from both G. clavigera and simulated herbivory in needle monoterpenes, but there was no difference between phytohormone applications and simulated herbivory in the phloem. 相似文献