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1.
The process of lesion formation and host response to natural infection by Armillaria ostoyae were studied in the roots of western larch (Larix occidentalis) and Douglas‐fir (Pseudotsuga menziesii ssp. glauca) trees in the three age classes, 6–8, 18–19 and 85–95 years. The characteristics of lesions on infected roots were recorded and bark samples were dissected from infection points and lesion margins in the field and stored in liquid nitrogen for macroscopic study in the laboratory. Infection in the roots of 6‐ to 8‐year‐old trees advanced freely, overcoming any host resistance, quickly girdling the root collar and killing the trees. In 18‐ and 19‐year‐old trees, however, 43% of infections on western larch and 27% of the infections on Douglas‐fir roots were confined to lesions bounded by necrophylactic periderms with multiple bands of phellem. Host response was similar in 85‐ to 95‐year‐old trees, but the percentage of confined lesions was higher than in younger trees. The results suggest that larch shows resistance to A. ostoyae at a younger age and with greater frequency than Douglas‐fir. 相似文献
2.
Structural responses in the bark and wood were described following penetration by Armillaria ostoyae in the roots of 20‐ to 30‐year‐old Douglas‐fir, western hemlock and western redcedar trees. Tissue necrosis presumably caused by fungal exudates was commonly observed at inoculum contact. In Douglas‐fir and western hemlock, A. ostoyae interfered with the initiation of active defence mechanisms involving the development of a lignified zone of impervious tissue (IT), necrophylactic periderm (NP) formation and compartmentalization of infected woody tissue. Breaching of IT and NP barriers was frequent, particularly around the clusters of sclereid cells in western hemlock. In western redcedar, the IT zone was inconspicuous. Induced rhytidome formation occurred in western redcedar either simultaneously with or after completion of NP development. The formation of this tissue facilitated en masse sloughing of infected tissue from the surface of roots. In western redcedar, traumatic phloem resin ducts formed in tangential bands surrounding the margin of expanded lesions. Effective compartmentalization in western redcedar was achieved by a barrier zone comprised of a higher‐than‐average number of axial parenchyma that accumulated polyphenolic deposits. A combination of host‐mediated defence mechanisms in western redcedar resulted in a significantly higher frequency of effective resistance reactions than in western hemlock or Douglas‐fir. 相似文献
3.
D. J. Morrison 《Forest Pathology》2011,41(1):31-40
Results are presented from several studies on the epidemiology of Armillaria ostoyae in Douglas‐fir plantations in the interior cedar‐hemlock (ICH) biogeoclimatic zone of British Columbia. Two plantations were monitored for mortality by A. ostoyae and other agents for 35 years after establishment. In these and other plantations ranging in age from 7 to 32 years, one or more of the following factors were determined: source of inoculum, mode of spread and characteristics of lesions on roots of excavated trees; symptom expression in relation to tree age and damage to the root system and years from initial infection to death on trees killed by the fungus. Mortality from A. ostoyae began in both plantations about 5 years after planting, reaching 30% in one and 11% in the other after 35 years. The spatial pattern of mortality was similar to that reported from New Zealand, France and South Africa; however, the temporal pattern differed, beginning later and, instead of declining, continuing at a nearly constant rate to the present. To age 10, nearly all infections were initiated by rhizomorphs; as plantations aged, the proportion of infections occurring at root–root contacts increased. In seven plantations, in moist and wet subzones of the ICH, from 23 to 52% of Douglas‐firs had root lesions, with the higher incidences occurring on moist sites. The occurrence of aboveground symptoms, reduced leader growth and basal resinosis, was related to the percentage of root length colonized by A. ostoyae on trees with more than 30% of root length killed. Average time from infection to death increased from 1 to 2 years at age 6 to 22 years at age 33. The outlook for timber yield from Douglas‐fir plantations in the cedar‐hemlock zone is discussed. Management alternatives for reducing damage from A. ostoyae when regenerating sites are reviewed. 相似文献
4.
The frequency of infection, lesion characteristics and anatomical changes in phloem and cambial tissues caused by Armillaria sinapina were studied on inoculated trees of Douglas‐fir, western hemlock and western redcedar and compared with results of A. ostoyae inoculations previously reported on the same host species. Similar percentages of inoculations resulting in infection of roots on the three hosts indicate that A. sinapina and A. ostoyae are equally pathogenic. Armillaria ostoyae was more virulent than A. sinapina as demonstrated by fungal exudates from A. ostoyae inoculum blocks, which appeared to cause lesions on roots; the higher frequency at which lignified impervious tissue (IT) and necrophylactic periderm (NP) developed in bark and following cambial invasion, compartmentalization; the large proportion of roots that showed no visible host response; the large zones of IT formed under continuous stimulation by A. ostoyae advancing in inner bark; and the high frequency of breaching of NP barriers. Spread of A. sinapina mycelium in host species appeared slower than that of A. ostoyae, particularly in Douglas‐fir and western hemlock. In western redcedar, A. ostoyae induced stronger host responses than those following invasion by A. sinapina, which included further expansion of the induced rhytidome response, traumatic phloem resin duct formation and higher numbers of polyphenolic parenchyma comprising its barrier zone. Where damage by A. sinapina ensued, it was always associated with high inoculum potential. The ecology of virulent and less virulent species of Armillaria in natural forests is discussed. 相似文献
5.
Periderm formation was studied in bark samples collected from the roots of western larch (Larix occidentalis) and Douglas‐fir (Pseudotsuga menziesii) trees infected with Armillaria ostoyae. Necrophylactic periderms were formed in advance of infection and successfully restricted continued fungal spread in 68 and 45% of the samples collected from 10‐ and 27‐year‐old western larch, respectively. However, all periderms formed in 11‐ and 25‐year‐old Douglas‐fir had been breached by the advancing fungus. In both species, necrophylactic periderms were commonly breached at the junction of the periderm with the vascular cambium. In western larch, stone phellem often comprised the external phellem layer of necrophylactic periderms with multiple bands of phellem. In 27‐year‐old western larch, infection was often confined to discrete lesions bounded by multiple periderms with multiple bands of phellem. In both tree species, phellem production was greater in response to infection than in response to abiotic wounding. 相似文献
6.
The sequence of events leading to necrophylactic periderm formation was studied throughout the year following the abiotic wounding of the non‐infected roots of 10‐ and 27‐year‐old western larch (Larix occidentalis) and 11‐ and 25‐year‐old Douglas‐fir (Pseudotsuga menziesii) trees that were infected with Armillaria ostoyae. The sequence was the same for both ages and species of trees. Wound repair was more rapid in the summer compared with the spring and autumn. Following cell hypertrophy, a zone of lignified impervious tissue was in the initial stages of formation within 10 days of wounding in the summer and 14 days in the spring or autumn. The new phellogen produced a layer of phellem three to four rows of cells thick after 20 days in the summer or 40 days in the spring. Modified cells abutting the inner boundary of the impervious zone frequently developed thick lignified abaxial walls and thin suberized adaxial walls. A typical exophylactic periderm in healthy root bark tissue of both western larch and Douglas‐fir consisted of stone phellem one to four rows of cells thick and a layer of thin‐walled phellem three to six rows of cells thick in western larch and two to three rows thick in Douglas‐fir, a single row of phellogen cells and one to three rows of phelloderm cells. Mature thin‐walled phellem cells had pigmented contents, red in western larch and light brown in Douglas‐fir. In response to wounding, 27‐year‐old western larch and 25‐year‐old Douglas‐fir developed necrophylactic periderms with annual bands of phellem. The bands included a layer of phellem that was six to 12 and nine to 15 rows of cells thicker than the layer of phellem observed in the respective naturally developed exophylactic periderms. Fifty days following wounding in the summer, stone phellem, one to three rows of cells thick, was observed in the necrophylactic periderm of 10‐year‐old trees. When fully developed, the necrophylactic periderm in 27‐year‐old western larch also had a layer of stone phellem three to five rows of cells thick in each band. Stone phellem development was only sporadic in 25‐year‐old Douglas‐fir. Wounds in the winter showed no signs of activity associated with repair until dormancy broke in the spring. 相似文献
7.
The sequence of events leading to the restoration of an intact periderm surface in the stem bark of Pinus sylvestris following wounding and challenge with Armillaria ostoyae was determined. A sub-erized impervious tissue (SIT) was produced outside the necrophylactic periderm (NP). Al-though A. ostoyae initially infected wounded bark with intracelfularly growing hyphae, these died before penetrating deeply into the bark. Later, intercellularly growing rhizomorphs, which had also developed from the inoculum, penetrated the NP and infected the functional phloem/ cambial zone. Compared to the reactions in uninfected wounded bark, the A. ostoyae infection delayed the formation of NP and reduced the frequency of cells involved in lignification and in SIT, but increased the number of lignified cell layers. The rhizomorph infection reached deeper and lignification was more pronounced in defoliated seedlings than in non-defoliated ones. The importance of structural responses in defence against A. ostoyae is discussed. 相似文献
8.
In this study the effects of thinning, fertilization and pruning on the vigour of Douglas fir (Pseudotsuga menziesii) and its susceptibility to Armillaria root disease were explored. Tree vigour was defined as the relative capacity for tree growth, expressed as the above-ground biomass increment per unit of photosynthetic tissue, or growth efficiency (GE). It has been hypothesized that trees with higher GE can better resist pathogen attack, and that GE can be used as a predictor of tree susceptibility to disease. In a previous study, four Douglas fir plantations were thinned, fertilized and pruned in all combinations, and the effects of these treatments on tree vigour were measured after 10 years. Root disease was not a factor in the initial study design, and mortality was ignored until 8 years after the treatments were applied. The results of an earlier study were utilized and the correlation between Armillaria root disease incidence and the effects of earlier stand treatments on tree growth was tested. Armillaria ostoyae was the primary mortality agent in the study area. Disease incidence on infested subplots ranged from 2 to 20%. Armillaria ostoyae incidence was the highest at medium tree density (6.1%), slightly lower on the low density (5.6%), and lowest on the unthinned plots (3.8%). There were no significant correlations between disease incidence and previous tree growth. The vigour of trees that became symptomatic or died by 1993 was not significantly different in 1983–85 from the vigour of trees that remained asymptomatic. On these sites, in areas of infestation, A. ostoyae is killing the largest, fastest growing trees, as well as less vigorous trees. Armillaria continues to cause mortality, regardless of the growth efficiency or growth rate of the host. 相似文献
9.
Commercial‐aged Douglas‐fir trees mechanically damaged during commercial thinning activities were investigated to quantify fungal taxa present 14 years after thinning. A total of 235 wood cores from 18 damaged and 18 non‐damaged control trees were obtained. Fungi were isolated on malt extract agar and molecular methods were used to identify 48 taxa. Damaged Douglas‐fir trees were more than twice as likely to contain wood‐inhabiting fungi as undamaged trees (p = 0.07) and contained more than 2.5 times more fungi (p = 0.006). However, typical Douglas‐fir stem‐decay fungi were not isolated. Fungi present were mostly endophytes and no significant differences were found in frequencies of decay/non‐decay soft‐rot fungi between damaged and control trees (p = 0.67). Likewise, no significant differences were found for the trend in number of fungi isolates over radial depth into the stem between damaged and control trees (p = 0.29). Frequency of fungal isolates was non‐linearly related to depth in the tree bole. Significant differences between damaged and control trees (p = 0.03) was influenced by the data in the outermost 3 mm. With the exception of these data, there were no significant differences between damaged and control trees in the radial trend in fungal frequency (p = 0.37). Results of this study suggest that wounding in commercial‐aged Douglas‐fir may not contribute significantly to occurrence and species composition of wood inhabiting fungi over the period of a short‐rotation final harvest. However, damage mitigation is recommended supported by a related study showing vertical discolouration affecting all wounded stems. 相似文献
10.
R. Siepmann 《Forest Pathology》1979,9(2):70-78
Stem and root rot of Douglas fir, Pseudotsuga menziesii (Mirb.) Franco. In 20 year old Douglas fir most of the butt rot was caused by Fomes annosus. Stem decay was central as well as eccentric reaching the sap wood in either case. The side roots of 20 year old Douglas fir were compared with those of a 40 year old stand with butt rot. In individual trees with decay there were less roots with Fomes annosus decay in the younger stand. Calocera viscosa was more abundant in the older stand. Soil conditions which might have favoured root rot in the young stand are discussed. 相似文献
11.
Climate change poses severe pressures to European conifer forests. Using non-native tree species, such as Douglas fir (Pseudotsuga menziesii), is one proposed strategy to circumvent adverse effects for forest management. However, novel forest health risks can impair the cultivation of non-native trees. In 2022, we observed large Douglas fir trees (approximately 40–50 years old, diameter at breast height (dbh) 21–41 cm) that had recently died in spring or summer 2022 in three forest stands in Eastern Austria. Intensive resin flow, blue-staining of the sapwood and the absence of bark- and wood-boring insects indicated a fungal infection. Isolations from blue-stained sapwood of the dead trees consistently yielded cultures of the opportunistic pathogen Diplodia sapinea. In a greenhouse wound inoculation experiment, seven D. sapinea isolates obtained from Douglas fir caused phloem necrosis, blue-staining of sapwood and mortality and thus displayed pathogenicity towards seedlings of both Ps. menziesii and its common host, Scots pine (Pinus sylvestris). Diplodia sapinea produced significantly longer areas of blue-stain as well as higher and faster mortality in Ps. menziesii compared to P. sylvestris. We conclude that D. sapinea substantially contributed to the death of seven of the 13 examined large Douglas fir trees. While this fungus has been described as a pathogen of young Douglas fir trees before, this is the first report that it can potentially kill large individuals of this conifer species under drought conditions. Thus, our results indicate that D. sapinea could represent a severe threat to the cultivation of Ps. menziesii in European forestry. 相似文献
12.
Plant lesions affect disease impact, progression and host resistance. Root lesions caused by Armillaria ostoyae (Romagn.) Herink were inspected on 24‐ to 34‐year‐old planted Douglas‐fir (Pseudotsuga menziesii (Mirb.) Franco) pulled from the soil in five locations. Four hundred seventy‐nine lesions were measured for length (mm), expansion rate (mm/year) and lesion type. Lesion types were patch lesions on a root side, girdled lesions covering the root circumference and root collar lesions on the stem below ground. Non‐linear mixed modelling revealed that fungal lesion length at girdled and collar lesions increased but the expansion rate slowed as a power function of time, indicating increasing secondary inoculum and host resistance. Average fungal lesion expansion rate in girdled root and collar lesions was 127 mm the first year which then declined non‐linearly to 26 mm/year after 10 years. Fungal lesion length in patch lesions showed little change after the first year of infection. For girdled and collar lesions, fungal lesion length increased with tree size at the time of infection. Sites with a longer frost‐free period had shorter girdled root and stem lesions, probably because of more active host defence. Results indicate that mortality would be rare from one infection event. Furthermore, patch lesions pose a lower risk of mortality than other lesions because of a lower fungal lesion expansion rate and secondary inoculum. Tree size, lesion position, lesion type and fungal lesion expansion rate need to be considered when assessing the risk of lesions to individual tree survival. Techniques to minimize the disease impact are discussed. 相似文献
13.
Plants utilize two general strategies to cope with pathogen attack. They either limit or resist the pathogen (termed ‘resistance’) or they cope with the disease by surviving and growing (termed ‘tolerance’). Both strategies tend to increase plant fitness; however, there are possible costs, trade‐offs and interactions associated with each strategy. This study focused on five half‐sib interior Douglas‐fir families [Pseudotsuga menziesii var. glauca (Beissn.) Franco] that were putatively classified as either resistant or susceptible to Armillaria ostoyae in a previous greenhouse study of seedling families challenged with A. ostoyae. We compared their survival rates in the greenhouse study with results of juvenile trees from the same five families that were artificially inoculated in field conditions. Both resistance and tolerance appeared to be operating in the field test trees, and a possible trade‐off between resistance and tolerance was detected. Significant differences were detected among the five families for stem radial growth following infection at the tree root collar. Compared with the putatively susceptible families, resistant families had smaller lesions and lower proportional root collar girdling. Tolerant families generally had larger lesions but demonstrated better growth when diseased than resistant families. One tolerant family that was a good survivor in the greenhouse survival study presented vertically shaped lesions that were large in area but had greatly reduced proportional root collar girdling. While a second family showed tolerant traits in the field study, its poor survival in the greenhouse study agreed with the large horizontally spreading lesions associated with high root collar girdling in the field study trees. Survival rankings of the five families in the greenhouse study mostly agreed with results from the field study based on the proportion of collar girdling among families. These host responses are discussed with respect to stability, quantity and quality of stands and products. 相似文献
14.
R. Siepmann 《Forest Pathology》1981,11(3):162-169
On the regeneration and infection of roots of Douglas fir (Pseudotsuga menziesii [Mirb.] Franco) by fungi present in the soil. 167 roots in 4 stands were severed 40–60 cm from the base of the tree. At the proximal cut ends of the thick roots of the 40 years old trees on loamy sand, either replacement roots or callous tissue were formed (only a small percentage of those roots was invaded by rot causing Basidiomycetes), or no new roots and no callus were formed and root rot fungi (Fomes annosus, Coniophora puteana, Resinicium bicolor) had invaded the root sections from the soil. On the root sections of the 35 and 55 years old Douglas firs on sand, with the exception of 2 roots, neither replacement roots nor callus were formed; Fomes and Coniophora had invaded the thicker roots of the older trees from the soil. 相似文献
15.
Conidia of Leptographium wageneri var. ponderosum, Leptographium wageneri var. pseudotsugae and Leptographium wageneri var. wageneri germinated on wounded roots of ponderosa pine (Pinus ponderosa), Douglas fir (Pseudotsuga menziesii), and pinyon (Pinus edulis), but not on unwounded roots. On wounds, germination occurred regardless of the combination of pathogen variety and plant species. When wounded roots of ponderosa pine and Douglas fir seedlings were inoculated with conidia of L. w. var. ponderosum and L. w. var. pseudotsugae, respectively, 87-100% of the seedlings became infected. When unwounded roots were inoculated with wooden blocks colonized with L. wageneri, 60-66% of the seedlings became infected. The development of L. w. var. ponderosum and L. w. var. pseudotsugae on agar was compared with their development on agar overlaid with either cellophane or Nucleopore membranes. Undulating hyphae were found commonly in and on the agar, but less frequently on the membranes. On both membranes, mycelia formed mats which were tightly appressed to the surface; mats were not formed on the agar. On both agar and the membranes, two or more hyphae often formed a strand. When L. w. var. ponderosum mycelia were allowed to colonize the surface of an unwounded pine root, strands, undulating hyphae and poorly formed mats were seen. 相似文献
16.
R. Von Siepmann 《Forest Pathology》1976,6(4):203-210
Polyporus (Phaeolus) schweinitzii and Sparassis crispa as frequent decay fungi in Douglas fir. Of fifty, 55 ears old trees in a Pseudotsuga menziesii stand. 90% showed decay of the heart- wood of the butt. The possibility of transmission of the most frequent decay fungi, P. schwei- nitzii and S. crispa, which also are frequent parasites in the roots and butts of pine, from the roots of the Pinus sylvestris trees of the previous crop into the roots of the Douglas fir is discussed. A first identification of the fungi was made possible by storing the stem discs under moist conditions. The mycelia of P. schweinitzii and S. crispa grew out of the decayed wood and possessed a characteristic form and colour. 相似文献
17.
Symptoms and signs associated with root rot caused by Heterobasidion annosum or Armillaria ostoyae in mountain pines (Pinus mugo ssp. uncinata) were investigated in the Swiss Alps. A sample of dying or recently dead mountain pine trees (≥12 cm d.b.h.) and saplings (<1.3 m height) was assessed for root pathogen infection by taking root samples followed by isolations in the laboratory. From a subsample, an additional core was taken from the butt of each tree and evaluated in the same fashion. A total of 157 dying or recently dead mountain pine trees and 184 saplings with roots infected by either of the two pathogens or which lacked infection were analyzed using logistic regression models. The main objectives were to determine the most prominent symptoms induced by the fungi (resinosis), signs of the fungi (mycelia, fruiting bodies and rhizomorphs), and tree characteristics (d.b.h./height and evidence of wounds) that would allow an easy and reliable determination of H. annosum and/or A. ostoyae infection of mountain pines in the field. Heterobasidion annosum caused both root and butt rot on mountain pine, whereas A. ostoyae was mostly restricted to the root systems of the trees sampled. The most discriminating sign for the presence of A. ostoyae infection was the presence of characteristic mycelial fans, and for H. annosum root rot the presence of H. annosum mycelia (sheets of paper‐thin mycelium and mycelial pustules). In addition, resinosis was a powerful predictor for A. ostoyae in trees. Symptoms and signs indicating A. ostoyae or H. annosum infections were more reliable for saplings than for mature trees. Armillaria rhizomorphs were not useful in detecting A. ostoyae infection and, if present, were often formed by saprophytic Armillaria species. Heterobasidion annosum fruiting bodies were rarely observed and poorly reflected the widespread occurrence of this pathogen in the mountain pine forests. 相似文献
18.
A parallel investigation to characterize host reactions involved in wound repair following abiotic injury and challenge with Armillaria ostoyae in three conifer species (Douglas‐fir, western hemlock and western redcedar) was conducted in the southern Interior of British Columbia. In this study, we characterize wound healing in roots following freezing injuries to the bark. Five weeks following wounding, all conifers developed lignified impervious tissue (IT) around killed tissue, and a necrophylactic periderm (NP) formed internally abutting IT, although IT was difficult to discern in western redcedar. Phellogen restoration in the secondary phloem of some western hemlock roots was retarded around large clusters of sclereids. Displacement of phloem fibre cells occurred in western redcedar but did not appear to delay NP formation. In roots exhibiting expansion of necrosis to the vascular cambium, callus tissue was evident at the margin of the wound. Novel host reactions in western redcedar involving induced rhytidome formation and the formation of traumatic phloem resin ducts are described. 相似文献
19.
A. Zamani R. Sturrock A. K. M. Ekramoddoullah S. B. Wiseman M. Griffith 《Forest Pathology》2003,33(5):299-316
Extracellular proteins were extracted from Phellinus weirii infected Douglas‐fir (Pseudotsuga menziesii var. menziesii) roots and needles to examine endochitinase activity. Chitinases have been associated with the plant's defence response against fungal attack because they hydrolyse chitin, a structural component of fungal cell walls. Protein separation using sodium dodecyl sulphate‐polyacrylamide gel electrophoresis (SDS‐PAGE) followed by Western immunoblot analysis using a polyclonal antibody specific to an endochitinase‐like protein (ECP) resulted in the detection of up to three polypeptides between 27 and 30 kDa in size. Two‐dimensional gel electrophoresis (2‐D PAGE) followed by Western immunoblot analysis revealed that the apoplastic fluid contained multiple ECP isoforms with isoelectric points (pIs) ranging from 5.3 to 5.8 and molecular masses of 27–30 kDa. Chitinase activity in needle and root tissues was measured spectrophotometrically using a colorimetric assay. A gel overlay technique using glycol chitin as a substrate for endochitinase was applied to confirm that the ECP antibody detected an enzymatically active protein. The apoplastic fluid collected from P. weirii‐infected winter Douglas‐fir needles showed anti‐freeze activity and seasonal analysis of needle tissue showed some evidence of ECP accumulation in winter months. ECP was distributed systemically throughout the tree. Increased levels of endochitinase activity in the region of P. weirii infection supports a physiological role for ECP in the plant defence response. 相似文献
20.
R. Von Siepmann 《Forest Pathology》1988,18(2):93-97
Intersterility groups and clones of Heterobasidion annosum isolates from root and butt rots of conifers 69 H. annousum heterokaryons from scots pine, Douglas fir, Norway spruce and larch belonged to the P group, three heterokaryons from Norway spruce to the S group. The results of a study with clones of H. annosum isolates from roots of Scotch pine trees in close neighbourhood suggest colonisation by H. annosum (partly) via root contacts. 相似文献