Fungal successions on pine needles fallen at different seasons: The succession of interior colonizers

Experimental studies were carried out to investigate seasonal effects on the fungal succession in the interior or decaying pine needles. At different seasons, the needles fallen for a short period were collected and marked, then placed on the surface of the O horizon in a pine forest. The needles were removed at intervals and their interior fungal communities were examined by using a surface sterilization technique. The successions of interior colonizers observed on the fallen needles at four different times are roughly divided into three groups based on the composition of species colonizing from litter. Seasonal shifts in the species combination were discussed with climatic and biotic factors. As a result, temperature at the surface of litter appeared to be a cardinal factor contributing to these seasonal changes in the succession of interior colonizers. Contributions to Sugadaira Montane Research Center, No. 164.     

Fungal succession on pine needles in Germany

The mycofloral succession on the needles ofPinus sylvestris was investigated in Tübingen, southwest Germany. Dead needles attached to the branches (D-type), those caught on branches (C-type) and three types of fallen needles, i.e., freshly fallen (L-type), slightly discolored (OL-type) and almost black needles (F-type) were examined for their fungal flora. Common primary saprophytes were rich on the dead needles on the tree, and on the L-type needles. They were replaced by successive species that contained the well-known species preferring pine needles as their substratum, such asVerticicladium trifidum orSympodiella acicola. Their ecological niches in pine leaf litter and their distribution patterns from a biogeographical viewpoint were discussed.     

Mycofloral succession onPinus densiflora needles on a moder site

Mycofloral succession on decaying pine needles in aPinus densiflora forest on a moder site was investigated in Sugadaira, Nagano Pref., central Japan. Dead needles on the tree, fallen needles obtained from two recognizable sublayers of the L layer and the upper sub-layer of the F₁ layer in the organic horizon were examined for their fungal flora using both washing and surface sterilization techniques. The major interior colonizer in freshly fallen needles varied with the season:Chaetopsina fulva in summer andSelenosporella curvispora in the other seasons.Thysanophora penicillioides was a remarkable external colonizer of freshly fallen needles in summer, while soil fungi were external colonizers of such needles in the other seasons. A possible successional change of major fungi with the needle decay was suggested. The observed seasonal alternation of the species colonizing freshly fallen needles was discussed in relation to climatic conditions. Contributions from Sugadaira Montane Research Center, No. 152.     

Fungal succession in the early decomposition process of pine cones on the floor ofPinus densiflora forests

The fungal succession on pine cones on the floor ofPinus densiflora forest was investigated in the early decomposition process (within ca. 30% decrease in dry weight). The fungal flora was examined by both washing and surface-sterilization methods on artificially placed cones and naturally fallen cones. The decomposition rates of artificially placed cones were 0.081–0.082 yr^–1. On withered cones still attached to the tree,Pestalotiopsis spp. were dominant. These fungi also occurred with higher frequencies after cones had lain on the floor and on cones in the L and FH horizons.Xylaria sp. andPhomopsis sp., which seem to colonize the interior of the tissue, occurred with higher frequencies on the cones on the tree, but their occurrence frequencies decreased after cones had lain on the forest floor. Conversely,Mortierella spp. andTrichoderma spp. newly occurred or their occurrence frequencies increased on lying cones. Of these,Trichoderma koningii increased rapidly and showed high occurrence frequencies.Thysanophora penicillioides, which prefers coniferous substrates, showed higher occurrence frequencies in the early stages of lying on the forest floor. On cones lying on the floor, the fungal flora did not significantly change during the investigation period.     

A Markov approach for describing post-fire succession of vegetation

Vegetation dynamics in the coastal area of the Seto Inland Sea region in Japan, where wild fires occur frequently, were described using a stationary Markov model. In this region, vegetation types ofMiscanthus-Pleioblastus grassland,Lespedeza-Mallotus scrub,Pinus-Rhododendron forest andCrassocephalum-Erechtites community have been identified, and these show cyclic succession under the influence of fires. The model uses parameters determining fire frequency and rate of successional change to analyze the effect of variation in these parameters on the areal ratio of each vegetation type at equilibrium and on the time taken for one vegetation type to succeed another (elapsed successional time). The effect of fire frequency differs between hypothetical habitats with high and low productivity. A policy for vegetation management in areas of high and low productivity is proposed. The advantages and limitations of applying Markov models to studies of vegetation succession are also discussed.     

Initial nitrogen content and topographic moisture effects on the decomposition of pine needles

The effects of litter quality and site characteristics on the decomposition process were investigated using a litterbag method. Pine needle litters with differing nitrogen concentrations (0.8, 0.6 and 0.4%) were placed on the upper and lower slopes of a Pinus thunbergii Parl. plantation. After both 3 and 6 months, the mass of decomposing litter with the lower nitrogen concentration was larger than the litter with higher nitrogen concentrations. After 9 months, there were no significant differences in the litter mass remaining, regardless of the initial nitrogen concentration. Moisture content in the litter was always higher on the lower slope, although the mass of litter was smaller. Nitrogen concentration of the decomposing litter increased linearly with accumulated mass loss. The increase in nitrogen concentration of decomposing litter was greater on the lower slope, but this increase did not differ between initial nitrogen concentrations. The nitrogen release from the decomposing litter with higher initial nitrogen concentration was larger than the release from litter bags with lower nitrogen concentrations. This result suggests that there may be positive feedback between soil nutrient availability, litter quality and nutrient release from decomposing litter at the intraspecific level.     

Ethylene production by different age class ponderosa and Jeffery pine needles as related to ozone exposure and visible injury

Summary Ethylene produced by different needle age classes representing natural populations of two ponderosa pine varieties [Pinus ponderosa var. arizonica (Engelm.) Shaw and var. ponderosa Dougl. ex Laws.] and Jeffery pine (Pinus jeffreyi Grev. and Balf.) was characterized using mercuric perchlorate traps. All populations contained individual trees which were either symptomatic or asymptomatic with respect to visible ozone injury. Ethylene production by different needle age classes was also characterized in P. ponderosa var. ponderosa seedlings grown in open top ozone fumigation chambers. Older age class needles produce significantly (P>0.05) more ethylene than younger age class needles. Needles of both P. ponderosa var. ponderosa and P. jeffreyi exhibiting ozone injury in the field produced significantly (P>0.05) higher levels of ethylene than asymptomatic conspecific trees. Seedlings exposed to the highest level of ozone in the fumigation study produced the highest levels of ethylene, followed by fumigation with medium and low ozone concentrations and carbon filtered air. These data indicate that the measurement of ethylene in conifer needles, as a measure of stress, needs to be calibrated for needle age class. It also suggests that the sensitivity of a tree to ozone injury may be regulated by the inherent ability of the individual to produce ethylene.     

Successful inoculation of mature pine with Tricholoma matsutake

Our finding demonstrates, for the first time, that the roots of mature pine trees can be successfully inoculated with a symbiotic ectomycorrhizal fungus, the valuable matsutake mushroom. Long root segments (ca. 5–10 mm in diameter, ca. 50 cm in length) of 50-year-old Pinus densiflora trees were excavated, washed, auxin-treated (2–5 mg indole butyric acid, IBA, per root) and incubated in moist Spagnum moss. Twelve months later, short roots were regenerated, of which approximately 90% were free of mycorrhizae. Mycorrhiza-free short roots were inoculated with mycelial pieces of Tricholoma matsutake and incubated further in a sterilized substrate. Four-and-a-half months later, roots putatively colonized by Matsutake were sampled near the inoculation points. A T. matsutake-specific ITS-rDNA fragment was amplified by nested PCR from approximately 80% of the root samples analyzed, whereas approximately 66% of the root samples processed for staining with Chlorazol black E displayed characteristic T. matsutake Hartig net structures. These results confirm the symbiotic infection of mature P. densiflora roots by matsutake.     

Spatial variation of seedling distribution in an east Mediterranean pine woodland at the beginning of post-fire succession

Most of the area in pine woodlands is occupied by perennial seeders that regenerate from seeds in the first winter after the fire and by annuals. Control of the germination in the regenerating vegetation after wildfire is therefore a primary ecological component of the post-fire succession in this ecosystem. The aim of the study presented here was to determine the distribution of Pinus, Cistus and other plants seeds around burned Pinus halepensis trees, and to measure the conditions related to seed germination in the upper soil layers in the same locations. The study was carried out in a 50-year old planted Pinus halepensis woodland that was burned down by a wildfire in July 1995. The variation of seedbank density was determined by collecting samples under the canopies of burned trees and in a nearby open area. Pine seedbank density decreased and that of Cistus and annuals increased with increasing distance from the burned trunks. Most pine seeds were present in the ash layer while those of the other plants were in the soil. In situ germination experiments showed that seedling density decreased with distance from the burned trunks while the proportion of pines in the seedling population increased. This was a result of seedbank variation and germination inhibition by the high pH conditions caused by the ash. The establishment of sparse pine seedling under the dead tree canopies insured their rapid development without interference by other plants and played a key role in the regeneration and stability of the pine woodland community. The concomitant mass germination of the perennial seeders in the rest of the area prevented invasion by annuals.     

Effects of drought and waterlogging on ultrastructure of Scots pine and Norway spruce needles

Effects of water stress on needle ultrastructure of 2-year-old Scots pine (Pinus sylvestris L.) and 5-year-old Norway spruce [Picea abies (L.) Karst.] seedlings were studied in greenhouse experiments. Drought stress was induced by leaving seedlings without watering, and waterlogging stress was produced by submerging the seedling containers in water. Needle samples for ultrastructural analyses were collected several times during the experiments, and samples for nutrient analyses at the end of the experiments. In drought stress, plasmolysis of mesophyll and transfusion parenchyma tissues, aggregation of chloroplast stroma and its separation from thylakoids and decreased size and abundance of starch grains in needles of both species were observed. The concentration of lipid bodies around the chloroplasts were detected in pine needles. Calcium and water concentrations in spruce needles were lower by the end of the experiments compared to controls. In waterlogging treatment, swelling of phloem cells in pine needles and large starch grains, slight swelling of thylakoids and increased translucency of plastoglobuli in chloroplasts of both species studied were observed. The phosphorus concentration in pine needles was higher while phosphorus, calcium and magnesium concentrations in spruce needles were lower in the waterlogging treatments compared to controls. Typical symptoms induced by drought stress, e. g. aggregation of chloroplast stroma and its separation from thylakoids, were detected, but, in waterlogging stress, ultrastructural symptoms appeared to be related to the developing nutrient imbalance of needles.     

Effect of provenance on free amino acid and chemical composition of Scots pine needles

Free amino acid (16 amino acids) and chemical composition (N, P, K, Mg, Ca, S, Fe, Mn, Cu, Zn) of Scots pine (Pinus sylvestris L.) needles were compared between six provenances in three different experimental areas. The main free amino acids in the needles were in the sequence of quantity; glutamic acid, glutamine, arginine and γ-aminobutyric acid. There were no significant differences in the concentrations of phenylalanine, γ-aminobutyric acid, methionine, proline and threonine in pine needles between the sites or between the provenances. Significant differences in the foliar concentrations of alanine and leucine were found between the sites and in the foliar concentrations of isoleucine, glutamine, glycine and tyrosine between the provenances. The concentrations of aspartic acid, glutamic acid, arginine and lysine were significantly affected by the sites and the provenances. The foliar nutrients, except copper, had statistically significant differences, both between the sites and between the provenances. Calcium did not differ between the provenances. This revised version was published online in June 2006 with corrections to the Cover Date.     

Chemical composition and biochemical changes in needles of Scots pine (Pinus sylvestris L.) stands at different stages of decline in Bulgaria

Water content was decreased in the needles of Scots pines (Pinus sylvestris L.) with moderate and slight levels of stress-induced decline. In pines with moderate decline a misbalance of foliar nutritional elements was observed. In 1-year-old needles of afflicted pines, the content of potassium and calcium was decreased, but the content of phosphorus, magnesium, iron, manganese and of the heavy metals cadmium and lead was increased. In 2-year-old needles of damaged pines, the content of calcium, iron and of the heavy metal cadmium was decreased, but the content of phosphorus, magnesium, manganese and zinc was increased. Potassium deficiency and zinc levels above toxicity tolerance characterized the nutrient status of declined Scots pines. In the same pines chlorophyll a/b ratio was diminished in 2-year-old needles and the protein content was increased in 1-year-old needles but was decreased in 2-year-old needles.In the second phase of the study 25- and 40-year-old pines with slight decline were examined. An increase in the protein content of 2-year-old needles of 25-year-old declined pines and of both 1- and 2-year-old needles of older 40-year-old pines was observed. A 280% increase of peroxidase activity in 1-year-old and a 178% increase in 2-year-old needles of 25-year-old damaged pines were shown. In older 40-year-old damaged pines a slight increase only in 1-year-old needles was seen. The observed age-related decrease in peroxidase activity in older 40-year-old symptomatic pines might be responsible for the lower resistance to decline in older pines. Our results show that clear but complex chemical and biochemical changes are observed in the needles of Scots pines in Bulgaria and that these changes hold promise for use as indicators of stress.     

Comparison of the initial demographies of pine and oak populations in rural pine forests in Korea and Japan

In order to understand the human impact upon demographic change in plant population in rural forests, we examined the population trends ofPinus densiflora andQuercus serrata inP. densiflora forests viewing their seedlings and saplings in rural Korea and Japan. The most prominent factor affecting the regeneration of the pine and oak was the intensity of management activity which controlled the vegetation stratification and its light environment. Open spaces, such as graveyard or cleaned area that allow the long-term direct daylight in dry season to accelerate the surface heat of soil, were unfavorable habitats for germination and growth of both species. The negative effects of the presence of litter and evergreen trees were related to the failure of early seedling and sapling growth of pine. Cover of litter is, especially, another factor related to the growth of pine saplings. It is considered that successful germination and sapling growth of pine and oak in early successional stages is determined by vegetation structures and light condition.     

Some observations on fungal succession during decomposition of wool in soil

Fungal succession on woolen baits was studied under laboratory conditions for more than one year. It was found that the initial colonizers on woolen baits are non-keratinophilic fungi, while the late colonizers are keratinophilic fungi. Six phases in total were observed during fungal succession. The successional trends obtained during decomposition of wool in soil samples collected from plain and hilly areas were almost the same, except for the dominant colonization in the last phase, which was constituted byChrysosporium tropicum for the plain, butMicrosporum gypseum andM. fulvum for the hilly area.     

Nitrogen availability affects saprotrophic basidiomycetes decomposing pine needles in a long term laboratory study

Fungi, especially basidiomycetous litter decomposers, are pivotal to the turnover of soil organic matter in forest soils. Many litter decomposing fungi have a well-developed capacity to translocate resources in their mycelia, a feature that may significantly affect carbon (C) and nitrogen (N) dynamics in decomposing litter. In an eight-month long laboratory study we investigated how the external availability of N affected the decomposition of Scots pine needles, fungal biomass production, N retention and N-mineralization by two litter decomposing fungi – Marasmius androsaceus and Mycena epipterygia. Glycine additions had a general, positive effect on fungal biomass production and increased accumulated needle mass loss after 8 months, suggesting that low N availability may limit fungal growth and activity in decomposing pine litter. Changes in the needle N pool reflected the dynamics of the fungal mycelium. During late decomposition stages, redistribution of mycelium and N out from the decomposed needles was observed for M. epipterygia, suggesting autophagous self degradation.     

Distance effect of co-occurring tree species on pine wilt disease incidence inPinus densiflora seedlings inoculated withBursaphelenchus xylophilus

Field-grownPinus densiflora seedlings were inoculated withBursaphelenchus xylophilus and the incidence of pine wilt disease was analyzed with respect to the spatial relationships between pine seedlings and adjacent seedlings of other tree species in a pure pine stand and three stands mixed withAlnus sieboldiana, Sarothamus scoparius or naturally associated species. The disease incidence was 60.9% in a 0–40 cm distance class from the nearestA. sieboldiana seedlings and then decreased with increasing minimum distance. The mean minimum distance between pine seedlings andA. sieboldiana was significantly shorter in diseased seedlings than in healthy ones. The highest disease incidence (53.9%) was found in a 0–40 cm distance class from the nearestS. scoparius, although the relationship with minimum distance was blurred by a high incidence in the 80-cm distance class. The mean minimum distance between pine seedlings was significantly short in diseased seedlings in a pure stand, whereas it was not so short between pine seedlings and other tree species in the two stands mixed withS. scoparius and the naturally associated species. The distance effect on disease incidence was noticeable inA. sieboldiana andP. densiflora at high density. This was not so clear inS. scoparius and was not found in the naturally associated species.     

不同微生境下齿肋赤藓(Syntrichia caninervis)生理生化特性对不同季节的响应

齿肋赤藓(Syntrichia caninervis)作为典型的耐旱藓类,在古尔班通古特沙漠的藓类结皮中占优势地位。该沙漠的季节气候差异较大,冬季低温湿润,春季干旱,夏季高温且干旱。荒漠藓类植物叶片仅具单层细胞,对外界环境的变化十分敏感。而有关荒漠藓类植物在生理上如何适应这种剧烈环境变化还不得而知。研究测定了生长于两种不同微生境下的齿肋赤藓,经由低温湿润的冬季到干旱的春季再到高温干旱的夏季过程中生理生化变化特征,以探究不同微生境下齿肋赤藓在水热变化剧烈的不同季节的适应机制。研究发现:季节、微生境及二者的交互作用能够显著影响齿肋赤藓的游离脯氨酸、可溶性糖、可溶性蛋白及丙二醛(MDA)含量、过氧化氢酶(CAT)、过氧化物酶(POD)、超氧化物歧化酶(SOD)活性。夏季的高温干旱使齿肋赤藓的脯氨酸、可溶性糖、MDA含量及3种抗氧化酶活性均显著高于冬季及春季,而可溶性蛋白含量却呈相反趋势。干旱的春季齿肋赤藓脯氨酸及可溶性糖显著高于冬季。而在低温湿润的冬季,齿肋赤藓丙二醛含量及过氧化氢酶活性均显著高于春季。表明夏季齿肋赤藓所受胁迫最大,脯氨酸、可溶性糖含量及抗氧化酶活性大幅度提高。同时,在具有积雪覆盖的冬季,两种微生境下的齿肋赤藓生理生化特性无显著差异;而在无积雪覆盖的春夏季节,灌丛的遮阴作用为齿肋赤藓提供了水含量相对较高的良好生活环境,其下齿肋赤藓的渗透调节物质含量和抗氧化酶活性均显著低于裸露地。表明生长在裸露地的齿肋赤藓较活灌丛拥有更强的胁迫耐受性。     

Distribution of Myxomycetes on coarse woody debris of <Emphasis Type="Italic">Pinus densiflora</Emphasis> at different decay stages in secondary forests of western Japan

The relationship between myxomycete species and the decay stage of wood of Pinus densiflora coarse woody debris was investigated in warm temperate secondary forests of western Japan. The number of species and species diversity of the myxomycete community reached the maximum on moderately decayed wood. The 25 dominant species recorded from 8 or more samples of the total 1530 samples were arranged in order of the succession index corresponding to the stage of decay. Species on slightly decayed hard pine wood were characterized by Stemonitis splendens, Enerthenema papillatum, and Physarum viride, whereas species of Cribrariaceae were found on brittle decayed soft wood increasing abundance according to the decay stages. Most of the species occurred where there was sufficient moisture preserved in the environment of the decaying wood, although S. splendens specifically emerged in low-moisture environments. Because the myxomycete species had preference to different decay stages of wood, it appears that they change sequentially during myxomycete community succession on dead pine wood according to the progression of decay.