温度对纵坑切梢小蠹伴生菌——云南半帚孢生长的影响

本文研究了纵坑切梢小蠹伴生菌云南半帚孢Leptographium yunnanense在不同温度下，在PDA培养基，云南松木段和活树上的生长变化情况，结果表明：在5－20℃温度范围内，真菌的生长随温度的升高而加快，云南松韧皮反应区长度逐渐增大；在20－35℃温度范围内，真菌的生长随温度的升高而逐渐减弱，韧皮反应区逐渐减小，说明温度变化使真菌的生长力与云南松的抗性之间的对比力量改变，从而导致真菌的生长长度和韧皮反应区发生相应变化。     

混交林抑制纵坑切梢小蠹危害作用研究

80年代以来，云南松遭受纵坑切梢小蠹的猖獗危害，造大量死亡。1992年在昆明市现有的云南松与栎类、云南松与华山松、云南松与滇油杉混交林及云南松纯林中设置11块定位观察样地，经过四年的观察，结果混交林平均每株受害率为10．9％，纯林均株受害率为48．95，混交林比纯林的纵坑切梢小蠹危害率降低38．0％，由于混交林内乔、灌、草种类的多样性，带来纵坑切梢小蠹天敌种类的多样性，有效地抑制了纵坑切梢小蠹的发生与发展，形成有虫不成灾的生态环境。     

混交林抑制纵坑切梢小蠹危害作用研究

８０年代以来，云南全省云南松林区遭受纵坑切梢小蠹的猖獗危害，造成大量林木死亡。１９９２年在昆明市现有的云南松与栎类、云南松与华山松、云南松与滇油杉混交林及云南松纯林中，设置１１块定位观察样地，观察云南松混交林蠹害发生情况，１９９２～１９９５年经过４年的观察，混交林平均株受害率１０．９％，纯林平均株受害率４８．９％。混交林比纯林降低纵坑切梢小蠹危害率３８．０％。混交林内由于乔、灌、草种类的多样性，带来纵坑切梢小蠹天敌种类的多样性，有效地抑制纵坑切梢小蠹的发生与发展，形成有虫不成灾的生态环境     

混交林抑制纵坑切梢小蠹危害作用研究

80年代以来,云南松遭受纵坑切梢小蠹的猖獗危害,造成大量死亡。1992年在昆明市现有的云南松与栎类、云南松与华山松、云南松与滇油杉混交林及云南松纯林中设置11块定位观察样地,经过四年的观察,结果混交林平均每株受害率为10.9％,纯林均株受害率为48.9％。混交林比纯林的纵坑切梢小蠹危害率降低38.0％。由于混交林内乔、灌、草种类的多样性,带来纵坑切梢小蠹天敌种类的多样性,有效地抑制了纵坑切梢小蠹的发生与发展,形成有虫不成灾的生态环境。     

混交林抑制纵坑切梢小蠹危害作用研究

８０年代以来，云南全省云南松林区遭受纵坑切梢小蠹的猖獗危害，造成大量林木死亡。１９９２年在昆明市现有的云南松与栎类，云南松与华山松，云南松与滇油杉混交林及云南松纯林中，设置１１块定位观察样地，观察云南松混交林蠹害发生情况，１９９２￣１９９５年经过４年的观察，混交林平均株受害率１０．９％，纯林平均株受害率４８．９％，混交林比纯林降低纵坑切梢小蠹危害率３８．０％，混交林内由于乔，灌，草种类的多样性，带     

云南松纵坑切梢小蠹种群生态研究现状

通过对现有文献的考察，从影响云南松纵坑切梢小蠹的危害因素，小蠹的生活史，蛀干蛀梢期、种群动态，生命表，空间分布格局以及虫情预测预报等方面阐述了云南松纵坑切梢小蠹种群生态的研究进展状况，以此对松小蠹的综合治理提出几点建议。     

纵坑切梢小蠹伴生菌危害机理的研究

该文对纵坑切梢小蠹伴生菌Leptographium yunnanense的危害机理作了初步研究，结果表明，L.yunnanense在与小伴随侵入后，产生致病毒素对寄主树林进行危害，从而辅助纵坑切梢小蠹成功定殖于寄主树木内，该毒素在低浓度条件下仍维持一定的致病力，进一步表明伴生菌在小蠹虫成功蛀害的过程中具有重要的作用。     

纵坑切梢小蠹伴生菌--云南半帚孢毒素作用机理的初步研究

用纵坑切梢小蠹伴生菌——云南半帚孢Laeptographium yunnanense的毒素处理后，云南松和华山松体内与抗病性密切相关的过氧化物酶、多酚氧化酶和苯丙氨酸解氨酶的活性都明显地下降，表明该毒素可以通过抑制寄主体内与抗性有关的酶的活性来削弱寄主的抗性，从而有利于菌在寄主体内侵染生长。云南松的酶活性下降的程度较大，而华山松的酶活性较高，毒素处理后活性受抑制的程度相对较小，这表明华山松对云南半帚孢的抗御能力较云南松的强，这也是云南境内云南松受害严重而华山松受害轻微的重要原因之一。     

云南松纵坑切梢小蠹种群生态研究现状

通过对现有文献的考察 ,从影响云南松纵坑切梢小蠹的危害因素 ,小蠹的生活史、蛀干蛀梢期、种群动态、生命表、空间分布格局以及虫情预测预报等方面阐述了云南松纵坑切梢小蠹种群生态的研究进展状况。以此对松小蠹的综合治理提出几点建议     

纵坑切梢小蠹种群变动及危害规律研究

以弥勒县的新哨和石林县的长湖受蠹害严重的林区的1000株云南松为定期观察样树,分别不同树木长势级别和受害木等级进行调查统计分析．结果表明,纵坑切梢小蠹在云南1年发生1代,且存在姐妹代现象,姐代种群占79．3％．妹代种群占18．6％．纵坑切梢小蠹对云南松的攻击有很强的选择性,优先攻击长势差的衰弱木．蠹害的危害从零星和簇状分布开始向大片状发展,待树木衰弱后进行蛀干危害．对蠹害木数量变化规律分析结果表明,树木受害越重,转化为枯死木的比例越高,当树木受害等级达4级以上时,树木很难再恢复生长．     

Fungi associated with Ips typographus on Picea abies in southern Poland and their succession into the phloem and sapwood of beetle‐infested trees and logs

This study dealt with the species distribution and frequency of fungi associated with the bark beetle Ips typographus (Scolytidae) on spruce trees of various states of health in southern Poland. The spruce trees were assessed by their degree of defoliation and damage of their crowns and trunks after attack by I. typographus. The state of health of trees from which samples were obtained was related to varying stages of brood development of I. typographus. Fungi were isolated from phloem taken from and around insect galleries and the sapwood underneath brood systems. Samples were taken from ‘healthy‐looking’, weakened, wind‐fallen and wind‐broken trees as well as from trap trees. The mycobiota associated with I. typographus was quite diverse in respect of the number of detected species, 65 fungal taxa were obtained from the phloem of trees infested by I. typographus, and 36 taxa occurred in the sapwood underneath insect galleries. The spectrum of fungi mainly consisted of ascomycetes and anamorphic fungi. The ophiostomatoid fungi were represented by 14 species and were the most numerously represented group in all niches examined. The most frequent ophiostomatoid species were Ceratocystis polonica, Ophiostoma ainoae, O. bicolor, O. penicillatum, O. piceae and O. piceaperdum. The frequency of occurrence of ophiostomatoid fungi differed significantly between the phloem and sapwood as well as in relation to the varying states of health of the spruce trees. These quantitative differences in the mycobiota of I. typographus between spruce trees belonging to different health categories can be explained by successional patterns of fungal colonization of host tissues following attack by I. typographus. The pathogenic species C. polonica was the primary invader, occurring most frequently in the sapwood of ‘healthy‐looking’ trees. Ophiostoma bicolor, O. penicillatum and O. piceaperdum also occurred during the early stages of brood development of I. typographus on ‘healthy‐looking’ trees, but they mainly colonized the phloem. In contrast, O. ainoae, O. minuta and O. piceae likely follow the aforementioned species as secondary and tertiary invaders into the phloem and the sapwood of spruce trees.     

Mineral nutrition,resin flow and phloem phytochemistry in loblolly pine

Southern pine beetles and associated pathogenic fungi represent the largest biotic threat to pine forests in the southeastern USA. The two primary defensive mechanisms of the tree to the beetle-fungal complex are the primary oleoresin flow and the concentrations of preformed and induced secondary compounds. We compared oleoresin flow and concentrations of phloem nutrients, soluble sugars, starch, total phenolics and proanthocyanidins in Pinus taeda L. trees in fertilized and control plots in the Sandhills region of North Carolina. Four blocks of 10 trees per treatment were sampled on five dates from May to November 1995. Phloem nitrogen and potassium concentrations were elevated in trees on fertilized plots, whereas phloem calcium concentrations were decreased. Fertilization significantly enhanced (10-20%) concentrations of phloem phenolics and proanthocyanidins. In contrast, phloem soluble sugars and starch concentrations were up to 30% lower in fertilized trees than in control trees. Increased phenolic concentrations and lower nonstructural carbohydrates should correlate with reduced tissue palatability and decreased pathogen susceptibility in fertilized trees; however, resin flows were significantly lower (30-100%) in fertilized trees compared with control trees, which may facilitate pine bark beetle establishment. Furthermore, fertilization-induced increases in phloem nitrogen concentration may be more important than tissue carbohydrate or phenolic content in determining tissue palatability.     

Seasonal water stress and the resistance of Pinus yunnanensis to a bark-beetle-associated fungus

We examined the influence of seasonal water stress on the resistance of Pinus yunnanensis (Franch.) to inoculation with Leptographium yunnanense, a pathogenic fungus associated with the aggressive bark beetle, Tomicus n. sp. Experiments took place between October 1997 and November 1999 in two plots located at the top and at the foot of a hill in Shaogiu, China, a region characterized by dry winters and wet summers. Following isolated and mass fungal inoculations, we observed the reaction zone length, fungal growth in the phloem, and the occlusion, blue-staining and specific hydraulic conductivity of the sapwood. Measurements of soil and needle water contents and predawn needle water potentials confirmed that trees were subject to mild water stress during winter, especially at the drier hilltop site. Measures of tree resistance to fungal infection of phloem and sapwood were congruent and indicated that trees were most susceptible to inoculation during the wet summer, especially at the lower-elevation plot. Specific hydraulic conductivity decreased after inoculation in summer. The results indicate that mild seasonal water stress is not likely responsible for the recent extensive damage to young P. yunnanensis stands by Tomicus n. sp. in the vicinity of our study plots. Rather, the results suggest that mild water stress enhances tree resistance to fungal pathogens associated with Tomicus n. sp.     

Biocontrol of decay in seasoning utility poles. I. Growth rate and colonizing ability of bluestain and decay fungi in vivo and in vitro

Seasoning (air drying) of utility poles for 6–12 months is essential before preservative treatment can be achieved. However, during seasoning, pine sapwood is often colonized by decay fungi, thereby compromising the performance and service life of the poles. This study investigated the potential of bluestain fungi to act as short‐term biocontrol agents against decay during seasoning. An important attribute for biocontrol is rapid growth, so growth rates of common bluestain (Ceratocystis coerulescens, Ophiostoma minus, Ophiostoma piceae, Ophiostoma piliferum, Sphaeropsis sapinea) and decay fungi (Heterobasidion annosum, Phlebiopsis gigantea, Stereum sanguinolentum) were compared on agar medium and pine in logs at various temperatures. On agar, the growth temperature optimum of most bluestain fungi and all the decay fungi was ～25°C, with little growth at ≤5°C or above 32.5°C. Overall, the fastest growing were S. sapinea and O. minus. In logs, the most effective colonizers were S. sapinea and O. minus with pathogenic abilities that made them well fitted to colonize the sapwood of freshly felled pine. Within these species, certain isolates produced much larger lesions in phloem and the sapwood tangential plane than all the decay fungi. Notably, there was significant variation in colonizing ability between different isolates within a species, emphasizing the need for testing a range of isolates when selecting a potential biocontrol agent.     

Effects of drought stress and high density stem inoculations with Leptographium wingfieldii on hydraulic properties of young Scots pine trees

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.     

Oak decline: I. Fungi associated with various disease symptoms on overground portions of middle-aged and old oak (Quercus robur L.)

Fungi colonizing overground portions of Quercus robur art presented separately for each of the following types of disease symptoms: leaves with atrophy and necrotic areas, dead branches, dead tops of branches, dead water sprouts, local necrotic areas in bark and phloem, locally dead and discoloured sapwood in debarked places, local discolorations inside sapwood, extensive nec-rotic areas on stems, dead trees. Over 80 species of fungi were found in all, mainly from As-comycotina and Deuteromycotina. Connected with each type of symptom there were several species of fungi which were more frequent than the remaining ones. In most cases fungi seem to be of secondary importance in development of symptoms characteristic for oak decline.     

Live storage and drying of storm-felled Norway spruce (Picea abies,L. Karst.) and Scots pine (Pinus sylvestris L.) trees

Storm-felled trees left in the forest for a shorter or longer period, affect the quality of the logs. The change in quality is mainly because of attack of fungi and insects, which in turn depends on the moisture content (MC) of the sapwood. The purpose of this study was to receive more knowledge about drying of storm-felled trees by investigating how fast winter storm-felled Norway spruce and Scots pine dried when left in the forest. Sixteen storm-felled spruces with part of the roots still in ground contact were selected from three stands and in addition to 10 pines from one of the stands. The trees were examined for MC in the sapwood until 21 months after the storm. This study indicates that wind-thrown trees with roots still connected to the soil can survive one summer without any value loss caused by draught, fungi and insects. The stand conditions can be of importance as the storm-felled trees in the stand, with scattered windthrow, were in best condition after one year, as they were shadowed by the trees still standing. Comparing spruces and pines with the stand with scattered windthrow, pines were more sensitive to drought and reached critical MC earlier.     

Intraspecific variation in Heterobasidion annosum for growth in sapwood of Picea abies and Pinus sylvestris

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.     

Methyl jasmonate and oxalic acid treatment of Norway spruce: anatomically based defense responses and increased resistance against fungal infection

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.     

Stimulation of tree defenses by Ophiostomatoid fungi can explain attack success of bark beetles on conifers

? Our aim is to present why the hypothesis, that Ophiostomatoid fungi play an important role in the establishment of most bark beetle species on living conifers, is valuable.

? After summarizing knowledge about the relationships of bark beetles with conifers and fungi, we conclude that controversy results from misinterpretations when using fungal pathogenicity to demonstrate the role of Ophiostomatoid fungi in beetle establishment on host trees.

? We demonstrate that fungal pathogenicity is not the right parameter to appreciate the role of fungus in beetle establishment on host trees. We argue that artificial low density inoculations that allow the appreciation of fungus ability to stimulate tree defenses and thus to help beetles in overcoming tree resistance must be used in complement to mass inoculations. In both cases, results must be expressed in terms of tree defense stimulation rather than in terms of tree killing.

1. Fungal species stimulating tree defenses are generally not those that grow the best in the sapwood.
2. We argue that beetle development in the phloem, fungal invasion of the sapwood and phloem, and tree death, occur after tree defenses are exhausted, and that any fungus present in the beetle gallery could thus potentially invade the sapwood after defense exhaustion.

? We conclude that stimulation of the tree defense reactions in both the phloem and the superficial sapwood is a real benefit brought by fungi to the beetles during the first phase of establishment (overcoming tree resistance).

? Considering the origin of the bark beetle fungus associations attacking living trees and their general functioning based on stimulation of tree defenses, we develop three hypotheses:

1. any beetle species would be helped in its establishment in a given tree species by developing an association, even loosely, with a fungus species belonging to the Ophiostomatoid flora of that tree species;
2. the necessity of a considerably low level of tree resistance for fungus extension into the tree is the selection pressure that has led fungi to develop their intrinsic ability to stimulate tree defenses, through their ability to grow into the phloem. This association can be completed by antagonistic fungal species controlling extension of the previous fungal species in the tree tissues;
3. Beetle species using the strategy of overcoming tree resistance are associated with a fungal complex, of which species could assume three roles regarding relationships between beetles and trees: 1- to stimulate tree defenses in the phloem and superficial sapwood, 2- to grow into the sapwood after tree resistance is overcome, and 3- to control phloem extension of the first other two categories. Bringing nutrients to the beetle progeny can be a fourth role.

? We propose that bark beetle — Ophiostomatoid associations can be categorized, based on associations’ frequency and complexity while taking into account beetle aggressiveness. We show that a close correspondence exists between beetles’ aggressiveness and the ability of their main associated fungal species to stimulate the defenses of their host tree.

? We conclude with suggesting that most sapwood invading fungi might be “cheaters” which have taken advantage of the efficiency of the relationship between beetles and fungi that stimulate tree defenses.