Ectomycorrhiza communities of red oak (<Emphasis Type="BoldItalic">Quercus rubra</Emphasis> L.) of different age in the Lusatian lignite mining district,East Germany

Ectomycorrhizal (ECM) communities were assessed on a 720 m² plot along a chronosequence of red oak (Quercus rubra) stands on a forest reclamation site with disturbed soil in the lignite mining area of Lower Lusatia (Brandenburg, Germany). Adjacent to the mining area, a red oak reference stand with undisturbed soil was investigated reflecting mycorrhiza diversity of the intact landscape. Aboveground, sporocarp surveys were carried out during the fruiting season in a 2-week interval in the years 2002 and 2003. Belowground, ECM morphotypes were identified by comparing sequences of the internal transcribed spacer regions from nuclear rDNA with sequences from the GenBank database. Fifteen ECM fungal species were identified as sporocarps and 61 belowground as determined by morphological/anatomical and molecular analysis of their ectomycorrhizas. The number of ECM morphotypes increased with stand age along the chronosequence. However, the number of morphotypes was lower in stands with disturbed soil than with undisturbed soil. All stands showed site-specific ECM communities with low similarity between the chronosequence stands. The dominant ECM species in nearly all stands was Cenococcum geophilum, which reached an abundance approaching 80% in the 21-year-old chronosequence stand. Colonization rate of red oak was high (>95%) at all stands besides the youngest chronosequence stand where colonization rate was only 15%. This supports our idea that artificial inoculation with site-adapted mycorrhizal fungi would enhance colonization rate of red oak and thus plant growth and survival in the first years after outplanting.     

Influence of soil nutrients on ectomycorrhizal communities in a chronosequence of mixed temperate forests

Many factors associated with forests are collectively responsible for controlling ectomycorrhizal (ECM) fungal community structure, including plant species composition, forest structure, stand age, and soil nutrients. The objective of this study was to examine relationships among ECM fungal community measures, local soil nutrients, and stand age along a chronosequence of mixed forest stands that were similar in vegetation composition and site quality. Six combinations of age class (5-, 26-, 65-, and 100-year-old) and stand initiation type (wildfire and clearcut) were replicated on four sites, each representing critical seral stages of stand development in Interior Cedar-Hemlock (ICH) forests of southern British Columbia. We found significant relationships between ECM fungal diversity and both available and organic P; available P was also positively correlated with the abundance of two ECM taxa (Rhizopogon vinicolor group and Cenoccocum geophilum). By contrast, ECM fungal diversity varied unpredictably with total and mineralizable N or C to N ratio. We also found that soil C, N, available P, and forest floor depth did not exhibit strong patterns across stand ages. Overall, ECM fungal community structure was more strongly influenced by stand age than specific soil nutrients, but better correlations with soil nutrients may occur at broader spatial scales covering a wider range of site qualities.     

Host preferences and differential contributions of deciduous tree species shape mycorrhizal species richness in a mixed Central European forest

Mycorrhizal species richness and host ranges were investigated in mixed deciduous stands composed of Fagus sylvatica, Tilia spp., Carpinus betulus, Acer spp., and Fraxinus excelsior. Acer and Fraxinus were colonized by arbuscular mycorrhizas and contributed 5% to total stand mycorrhizal fungal species richness. Tilia hosted similar and Carpinus half the number of ectomycorrhizal (EM) fungal taxa compared with Fagus (75 putative taxa). The relative abundance of the host tree the EM fungal richness decreased in the order Fagus > Tilia >> Carpinus. After correction for similar sampling intensities, EM fungal species richness of Carpinus was still about 30–40% lower than that of Fagus and Tilia. About 10% of the mycorrhizal species were shared among the EM forming trees; 29% were associated with two host tree species and 61% with only one of the hosts. The latter group consisted mainly of rare EM fungal species colonizing about 20% of the root tips and included known specialists but also putative non-host associations such as conifer or shrub mycorrhizas. Our data indicate that EM fungal species richness was associated with tree identity and suggest that Fagus secures EM fungal diversity in an ecosystem since it shared more common EM fungi with Tilia and Carpinus than the latter two among each other.     

<Emphasis Type="Italic">Polygonum viviparum</Emphasis> mycobionts on an alpine primary successional glacier forefront

Polygonum viviparum is one of the first ectomycorrhizal (EM) plant species colonising primary successional sites at the Rotmoos glacier forefront (Tyrolean Alps, Austria). On a site with soil development of about 150 years (2,400 m above sea level), mycobionts of P. viviparum were identified by morphotyping and fungal ribosomal deoxyribonucleic acid internal transcribed spacer sequencing. For studying seasonal dynamics and spatial heterogeneity, ectomycorrhizae were sampled on five plots during all seasons. P. viviparum root tips were always EM. In total, 18 mycobiont taxa of the following genera were identified: Cenococcum (1), Cortinarius (2), Helvella (1), Inocybe (3), Russula (1), Sebacina (2), Thelephora (2) and Tomentella (6). All were non-specific EM partners of EM plants. As early as 2 weeks after spring snow melt, EM were well developed, vital and showed high mycobiont diversity. The relative abundance of senescent root tips was lowest in spring and increased throughout the year, with a maximum in winter (frozen soil). Thus, mycobiont growth and physiological activity obviously start when soil is still under snow cover: We speculate that water availability is one important initiation factor for mycorrhizal development under snow cover, when temperatures still range around the freezing point. Irrespectively of the season, most abundant mycobionts at this primary successional site belonged to the genera Tomentella, Sebacina and Cenococcum, also in frozen soil. Spatial heterogeneity was high when considering species composition and diversity indices. Overall mycobionts species richness was restricted at this site, probably because of the limited availability of fungal partners. We regard the presence/absence of fungal partner and limiting abiotic impacts of the environment as key factors for the symbiotic status of P. viviparum. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Ectomycorrhizal fungal succession in mixed temperate forests

Ectomycorrhizal (ECM) fungal communities of Douglas-fir (Pseudotsuga menziesii) and paper birch (Betula papyrifera) were studied along a chronosequence of forest development after stand-replacing disturbance. Previous studies of ECM succession did not use molecular techniques for fungal identification or lacked replication, and none examined different host species. Four age classes of mixed forests were sampled: 5-, 26-, 65-, and 100-yr-old, including wildfire-origin stands from all four classes and stands of clearcut origin from the youngest two classes. Morphotyping and DNA sequences were used to identify fungi on ECM root tips. ECM fungal diversities were lower in 5-yr-old than in older stands on Douglas-fir, but were similar among age classes on paper birch. Host-specific fungi dominated in 5-yr-old stands, but host generalists were dominant in the oldest two age classes. ECM fungal community compositions were similar in 65- and 100-yr-old stands but differed among all other pairs of age classes. Within the age range studied, site-level ECM fungal diversity reached a plateau by the 26-yr-old age class, while community composition stabilized by the 65-yr-old class. Simple categories such as 'early stage', 'multi stage', and 'late stage' were insufficient to describe fungal species' successional patterns. Rather, ECM fungal succession may be best described in the context of stand development.     

Belowground interspecific competition in mixed boreal forests: fine root and ectomycorrhiza characteristics along stand developmental stage and soil fertility gradients

We studied fine roots and ectomycorrhizas in relation to aboveground tree and stand characteristics in five mixed Betula pendula Roth, Picea abies (L.) H. Karst., and Pinus sylvestris L. stands in Southern Finland. The stands formed gradients of developmental stage (15-, 30-, and 50-year-old stands) in the stands of medium fertility, and of site fertility in the young stands (30-year-old fertile, medium fertile, and least fertile stands). The biomass of the external hyphae of ectomycorrhizas (ECM) was the highest, and the diversity of the fungal community the lowest, in the most fertile stand. The vertical distributions of fine roots of the three tree species were mostly overlapping, indicating high inter-specific belowground competition in the stands. We did not find any clear trends in the fine root biomass (FRB) or length across the stand developmental stages. The FRB of the conifers varied with site fertility, whereas in B. pendula it was almost constant. In contrast to the conifers, the specific root length (SRL) of B. pendula clearly increased from the most fertile to the least fertile stand. This indicates differences in the primary nutrient acquisition strategy between conifers and B. pendula.     

Succession of mycorrhizal fungi in stands of Pinus sylvestris in the Netherlands

The mycorrhizal mycoflora was investigated in 35 stands of Pinus sylvestris in three types of young (4-13 yr) and three of old (50-80 yr) stands in the Netherlands, differing in number of rotations and soil type. A plot of 1050 m² (30 m x 35 m) within each stand was searched for carpophores during the autumns of 1986 and 1987. 10 soil samples per plot were taken in October 1987 in order to assess the mycorrhizal status of the tree roots. The composition of mycorrhizal mycoflora in the different plots was subjected to TWINSPAN cluster analysis and Detrended Correspondence Analysis. Plot groupings generated by these analyses largely parallelled the stand types, indicating that each stand type has its own mycoflora. Differences in myco-floristic composition between stand types were parallelled by differences in the composition of green vegetation. The young stand types had 3.5–27 x more carpophores and 1.4–6.8 x more species than two of the old stand types One old stand type was intermediate. Considerable differences in species composition between the young stand types were observed. It is concluded that the succession of mycorrhizal fungi is not primarily influenced by ageing of the trees, but rather by changes in the soil. The results were compared with data on changes in the occurrence of fruiting species of mycorrhizal fungi in the Netherlands during this century. It appeared that species which have declined according to these data were more frequent in the young plots than in the old plots. However, these species are reported to be frequent in old stands of P. sylvestris in Estonia and Finland. It is argued that this difference is related to the high nitrogen deposition in the Netherlands.     

Ectomycorrhizae and ectomycorrhizal fungal fruit bodies in pine stands differentially damaged by pine wilt disease

We surveyed ectomycorrhizae, ectomycorrhizal fungal fruit bodies, and soil physical properties in one heavily damaged and two lightly damaged pine stands on Mt. Tsukuba, central Japan. The rate of ectomycorrhizal root tips was not different between heavily and lightly damaged pine stands. For ectomycorrhizae, Cenococcum geophilum had high relative abundance in the heavily damaged pine stand. The number of ectomycorrhizal fungal fruit bodies in the heavily damaged pine stand was much lower than that in the lightly damaged pine stands.     

Dynamics of ectomycorrhizae in Abies amabilis stands: The role of Cenococcum graniforme

The growth and development of the predominant mycorrhizal fungus infecting the root systems of subalpine Pacific silver fir was examined seasonally in 23 and 180 yr old stands. All data was obtained by sorting roots from intact soil cores separated into forest floor and mineral soil horizons.
The predominant mycorrhizal fungus infecting the root systems of 23 and 180 yr old Pacific silver fir was Cenococcum graniforme. C. graniforme comprised 36–97% and 40–96% of the total mycorrhizal root tips in the soil profile in the young and old stands, respectively. C. graniforme root biomass reached a maximum in autumn in the young stand, and in winter in the old. Dual mycorrhizal associations were commonly observed on the same root tip during the spring and autumn periods of root elongation in both stands. Over 80% of the C. graniforme roots were located in the forest floor and A-horizon throughout the study in both stands, Sclerotia were predominantly located in the A-horizon (38-70%) in both stands.     

Impact of <Emphasis Type="Italic">Heterobasidion</Emphasis> root-rot on fine root morphology and associated fungi in <Emphasis Type="Italic">Picea abies</Emphasis> stands on peat soils

We examined differences in fine root morphology, mycorrhizal colonisation and root-inhabiting fungal communities between Picea abies individuals infected by Heterobasidion root-rot compared with healthy individuals in four stands on peat soils in Latvia. We hypothesised that decreased tree vitality and alteration in supply of photosynthates belowground due to root-rot infection might lead to changes in fungal communities of tree roots. Plots were established in places where trees were infected and in places where they were healthy. Within each stand, five replicate soil cores with roots were taken to 20 cm depth in each root-rot infected and uninfected plot. Root morphological parameters, mycorrhizal colonisation and associated fungal communities, and soil chemical properties were analysed. In three stands root morphological parameters and in all stands root mycorrhizal colonisation were similar between root-rot infected and uninfected plots. In one stand, there were significant differences in root morphological parameters between root-rot infected versus uninfected plots, but these were likely due to significant differences in soil chemical properties between the plots. Sequencing of the internal transcribed spacer of fungal nuclear rDNA from ectomycorrhizal (ECM) root morphotypes of P. abies revealed the presence of 42 fungal species, among which ECM basidiomycetes Tylospora asterophora (24.6 % of fine roots examined), Amphinema byssoides (14.5 %) and Russula sapinea (9.7 %) were most common. Within each stand, the richness of fungal species and the composition of fungal communities in root-rot infected versus uninfected plots were similar. In conclusion, Heterobasidion root-rot had little or no effect on fine root morphology, mycorrhizal colonisation and composition of fungal communities in fine roots of P. abies growing on peat soils.     

Fine-scale distribution of ectomycorrhizal fungi colonizing Tsuga diversifolia seedlings growing on rocks in a subalpine Abies veitchii forest

Numerous species of ectomycorrhizal (ECM) fungi coexist under the forest floor. To explore the mechanisms of coexistence, we investigated the fine-scale distribution of ECM fungal species colonizing root tips in the root system of Tsuga diversifolia seedlings in a subalpine forest. ECM root tips of three seedlings growing on the flat top surface of rocks were sampled after recording their positions in the root system. After the root tips were grouped by terminal-restriction fragment length polymorphism (T-RFLP) analysis of ITS rDNA, the fungal species representing each T-RFLP group were identified using DNA sequencing. Based on the fungal species identification, the distribution of root tips colonized by each ECM fungus was mapped. Significant clustering of root tips was estimated for each fungal species by comparing actual and randomly simulated distributions. In total, the three seedlings were colonized by 40 ECM fungal species. The composition of colonizing fungal species was quite different among the seedlings. Twelve of the 15 major ECM fungal species clustered significantly within a few centimeters. Some clusters overlapped or intermingled, while others were unique. Areas with high fungal species diversity were also identified in the root system. In this report, the mechanisms underlying generation of these ECM root tip clusters in the root system are discussed.     

Abundance,diversity, and vitality of mycorrhizae of Scots pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis> L.) in lignite recultivation sites

Scots pine (Pinus sylvestris L.) stands cover large areas in the Lusatian and the Middle German lignite mining districts. Due to adverse chemical substrate conditions, the root systems of the trees are restricted to the ameliorated top-spoil and the organic forest floor layers. To investigate functioning of fine root systems under the prevailing site factors, we studied mycorrhizal colonization rate and frequency as well as mycorrhizal diversity, vitality and growth phases in Scots pine ecosystems along a chronosequence in both mining districts. Mycorrhizal rate was close to 100% in both districts. Mycorrhizal abundance was higher in the organic forest floor layer than the mineral soil layer. In total, 25 morphotypes were recorded. Diversity differed between the districts. The mycorrhizae of Amphinema byssoides, Tuber puberulum, Pinirhiza discolor, Pinirhiza cf. bicolorata and E-type were present in both mining areas. These morphotypes are typical of nutrient-rich soils with high pH values. Compared with the undisturbed sites, vitality of mycorrhizae was very high at the test sites on spoil substrate, correlating with the high growth dynamics of mycorrhizae at recultivation sites. A relatively high carbon flow to the mycorrhizal root systems at these sites seems likely. Thus, mycorrhizal root systems are able to cope with the ameliorated top-spoil and the organic layer. The main reason for the adaptation is the large number of ectomycorrhizal fungal species available in this area where Pinus sylvestris is indigenous.     

Distribution of lead in mycorrhizal and non-mycorrhizal Norway spruce seedlings

Non-mycorrhizal spruce seedlings (Picea abies Karst.) and spruce seedlings colonized with Lactarius rufus (Scop.) Fr. or two strains of Paxillits involutus (Batsch) Fr. were grown in an axenic silica sand culture system with frequently renewed nutrient solution. After successful mycorrhizal colonization, the seedlings were exposed to 1 μM PbCI₂ for 19 weeks. The degree of infection in all of the mycorrhizal treatments approached 100% during the experiment and was not affected by exposure to Pb. However, the number of root tips per root dry weight and the shoot: root ratio, both in the non-mycorrhizal and the mycorrhizal seedlings, had decreased after the 19 week treatment with PbCl₂ Using X-ray microanalysis, the distribution and concentration of Pb in the tissues of mycorrhizal and non-mycorrhizal root tips were compared. In the mycorrhizae of seedlings exposed to Pb no significant accumulation of Pb in the hyphal mantle or in fungal cell walls of the Hartig net were detected. Lead accumulated primarily in the cortex cell walls both of non-mycorrhizal and mycorrhizal root tips. No significant difference of Pb concentrations in root cortex cell walls of non-mycorrhizal and mycorrhizal seedlings was found; except for seedlings colonized with Paxillus involutus strain 537. However, at the endodermis no effect of mycorrhizal fungal colonization on the Pb tissue concentration was detected. The presence of the fungal sheath did not prevent Pb from reaching the root cortex. The endodermis acted as a barrier to Pb radial transport in both mycorrhizal and non-mycorrhizal seedling roots.     

Biomass and nutrient concentrations of sporocarps produced by mycorrhizal and decomposer fungi in Abies amabilis stands

Summary Sporocarps and sclerotia were collected for a one-year period in 23- and 180-year-old Abies amabilis stands in western Washington. All sporocarps were classified and chemically analyzed for N, P, K, Ca, Mg, Na and Fe. Lactarius sp. and Cortinarius sp. contributed the largest proportion of the total annual epigeous sporocarp production in both stands. Annual epigeous production was 34 kg/ha in the young stand and 27 kg/ha in the mature stand. Hypogeous sporocarp production increased from 1 kg ha^-1 yr^-1 to 380 kg ha^-1 yr^-1 with increasing stand age. High sclerotia biomass occurred in the young (2,300 kg/ha) and mature (3,000 kg/ha) stands. Peak sclerotia and epigeous sporocarp biomass in the young stand and epigeous and hypogeous sporocarp biomass in the mature stand coincided with the fall peak of mycorrhizal root biomass.In the young stand, sporocarps produced by decomposer fungi concentrated higher levels of Ca and Mn than those produced by mycorrhizal fungi. In the mature stand, sporocarps of decomposer fungi concentrated higher levels of N, P, Mn, Ca and Fe than sporocarps of mycorrhizal fungi. Epigeous and hypogeous sporocarps concentrated higher levels of N, P, and K than sclerotia or mycelium. The highest concentration of N (4.36%), P (0.76%), K (3.22%) and Na (1,678 ppm) occurred in epigeous sporocarps. Highest Mn (740 ppm) and Ca (20,600 ppm) concentrations occurred in mycelium, while highest Mg (1,929 ppm) concentrations were in hypogeous sporocarps and highest Fe (4,153 ppm) concentrations were in sclerotia.     

Morphological classification of ectomycorrhizas ofPinus densiflora

Morphological classification of ectomycorrhizas ofPinus densiflora was conducted. Fifty soil samples containing pine ectomycorrhizas, and 40 pine seedlings were collected randomly in two separate reforested stands ofP. densiflora (45 yr old) from May 1992 to October 1994. Fifty-six types of ectomycorrhizas could be classified based upon microscopically observable morphological characteristics. Fifty percent of the types showed cystidia or other specific characteristics such as laticiferous hyphae in the fungal sheaths, verrucose emanating hyphae and a positive hyphal reaction to UV irradiation. Four mycorrhizal types were confirmed to be formed by the fungiRussula delica, R. mariae, R. nigricans, andCenococcum geophilum, respectively. Although the other 52 types were unidentified mycobionts at species level, it was inferred that they were formed by the fungiHebeloma, Lactarius, Russula andTuber. There was a slight difference in the observed mycorrhizal types between the tree ages. Contribution No. 127, Laboratories of Plant Pathology and Mycology, Institute of Agriculture and Forestry, University of Tsukuba.     

Ectomycorrhizal fungal communities in late-successional Swedish boreal forests, and their composition following wildfire

This study was conducted to evaluate the effects of wildfires on ectomycorrhizal (EM) fungal communities in Scots pine ( Pinus sylvestris ) stands. Below- and above-ground communities were analysed in terms of species richness and evenness by examining mycorrhizas and sporocarps in a chronosequence of burned stands in comparison with adjacent unburned late-successional stands. The internal transcribed spacer (ITS)-region (rDNA) of mycobionts from single mycorrhizas was digested with three restriction enzymes and compared with an ITS–restriction fragment length polymorphism (RFLP) reference database of EM sporocarps. Spatial variation seemed to be more prominent than the effects of fire on the EM fungal species composition. Most of the common species tended to be found in all sites, suggesting that EM fungal communities show a high degree of continuity following low-intensity wildfires. Species richness was not affected by fire, whereas the evenness of species distributions of mycorrhizas was lower in the burned stands. The diversity of EM fungi was relatively high considering that there were only three EM tree species present in the stands. In total, 135 EM taxa were identified on the basis of their RFLP patterns; 66 species were recorded as sporocarps, but only 11 of these were also recorded as mycorrhizas. The species composition of the below-ground community of EM fungi did not reflect that of the sporocarps produced. EM fungal species present in our ITS–RFLP reference database accounted for 54–99% of the total sporocarp production in the stands, but only 0–32% of the mycorrhizal abundance.     

Fungal communities in mycorrhizal roots of conifer seedlings in forest nurseries under different cultivation systems, assessed by morphotyping, direct sequencing and mycelial isolation

Fungi colonising root tips of Pinus sylvestris and Picea abies grown under four different seedling cultivation systems were assessed by morphotyping, direct sequencing and isolation methods. Roots were morphotyped using two approaches: (1) 10% of the whole root system from 30 seedlings of each species and (2) 20 randomly selected tips per plant from 300 seedlings of each species. The first approach yielded 15 morphotypes, the second yielded 27, including 18 new morphotypes. The overall community consisted of 33 morphotypes. The level of mycorrhizal colonisation of roots determined by each approach was about 50%. The cultivation system had a marked effect on the level of mycorrhizal colonisation. In pine, the highest level of colonisation (48%) was observed in bare-root systems, while in spruce, colonisation was highest in polyethylene rolls (71%). Direct internal transcribed spacer ribosomal DNA sequencing and isolation detected a total of 93 fungal taxa, including 27 mycorrhizal. A total of 71 (76.3%) fungi were identified at least to a genus level. The overlap between the two methods was low. Only 13 (13.9%) of taxa were both sequenced and isolated, 47 (50.5%) were detected exclusively by sequencing and 33 (35.5%) exclusively by isolation. All isolated mycorrhizal fungi were also detected by direct sequencing. Characteristic mycorrhizas were Phialophora finlandia, Amphinema byssoides, Rhizopogon rubescens, Suillus luteus and Thelephora terrestris. There was a moderate similarity in mycorrhizal communities between pine and spruce and among different cultivation systems.Electronic supplementary material Supplementary material is available for this article at and accessible for authorised users     

Stand age-related effects on soil respiration in a first rotation Sitka spruce chronosequence in central Ireland

The effect of stand age on soil respiration and its components was studied in a first rotation Sitka spruce chronosequence composed of 10‐, 15‐, 31‐, and 47‐year‐old stands established on wet mineral gley in central Ireland. For each stand age, three forest stands with similar characteristics of soil type and site preparation were used. There were no significant differences in total soil respiration among sites of the same age, except for the case of a 15‐year‐old stand that had lower soil respiration rates due to its higher productivity. Soil respiration initially decreased with stand age, but levelled out in the older stands. The youngest stands had significantly higher respiration rates than more mature sites. Annual soil respiration rates were modelled by means of temperature‐derived functions. The average Q ₁₀ value obtained treating all the stands together was 3.8. Annual soil respiration rates were 991, 686, 556, and 564 g C m^?2 for the 10‐, 15‐, 31‐, and 47‐year‐old stands, respectively. We used the trenching approach to separate soil respiration components. Heterotrophic respiration paralleled soil organic carbon dynamics over the chronosequence, decreasing with stand age to slightly increase in the oldest stand as a result of accumulated aboveground litter and root inputs. Root respiration showed a decreasing trend with stand age, which was explained by a decrease in fine root biomass over the chronosequence, but not by nitrogen concentration of fine roots. The decrease in the relative contribution of autotrophic respiration to total soil CO₂ efflux from 59.3% in the youngest stand to 49.7% in the oldest stand was explained by the higher activity of the root system in younger stands. Our results show that stand age should be considered if simple temperature‐based models to predict annual soil respiration in afforestation sites are to be used.     

Post-fire seedlings of Nothofagus alpina in Southern Chile show strong dominance of a single ectomycorrhizal fungus and a vertical shift in root architecture

We investigated belowground responses of Nothofagus alpina seedlings to post-fire conditions during natural regeneration after a wildfire in Chile, focusing on mycorrhizal community and root architecture. The complete root systems of 2-year-old N. alpina seedlings were extracted from a post-fire site with natural regeneration and compared to roots of seedlings from undisturbed forest nearby. Mycorrhizal morphotype richness was determined in each seedling. Morphometric parameters of tertiary root structure and dry biomass of whole root systems were determined in 5 cm vertical intervals and in four lateral root classes. With 43.5% of colonized vital mycorrhizal root tips, the Basidiomycete Descolea antarctica was the most abundant fungal symbiont on post-fire seedlings. Tertiary root morphology of these seedlings was distinct from control plants and characterized by a deep-reaching tap root with rather evenly distributed lateral branches whereas seedlings from the undisturbed site had shallower root systems with most lateral roots concentrated in the upper soil layers. Post-fire seedlings had more mycorrhizal rootlets and mycorrhiza-bearing third order lateral roots than control plants which was expressed in a 34% higher total root number but only a 10% higher total root biomass, although both values were not statistically significant. A major part of these fine roots in seedlings from burnt forest was found in deeper soil horizons, compared to the seedlings from undisturbed forest. According to our results, post-fire conditions clearly favour Descolea antarctica as an early ectomycorrhizal colonizer of Nothofagus seedlings at the studied site. As no significant changes in soil chemistry could be observed at the burnt site, the deep-reaching tertiary root architecture of these seedlings may be interpreted as a response to other abiotic factors like reduced moisture in surface soil.     

Stand fine root biomass and fine root morphology in old-growth beech forests as a function of precipitation and soil fertility

Only very limited information exists on the plasticity in size and structure of fine root systems, and fine root morphology of mature trees as a function of environmental variation. Six northwest German old-growth beech forests (Fagus sylvatica L.) differing in precipitation (520 – 1030 mm year^–1) and soil acidity/fertility (acidic infertile to basic fertile) were studied by soil coring for stand totals of fine root biomass (0–40 cm plus organic horizons), vertical and horizontal root distribution patterns, the fine root necromass/biomass ratio, and fine root morphology (root specific surface area, root tip frequency, and degree of mycorrhizal infection). Stand total of fine root biomass, and vertical and horizontal fine root distribution patterns were similar in beech stands on acidic infertile and basic fertile soils. In five of six stands, stand fine root biomass ranged between 320 and 470 g m^–2; fine root density showed an exponential decrease with soil depth in all profiles irrespective of soil type. An exceptionally small stand fine root biomass (<150 g m^–2) was found in the driest stand with 520 mm year^–1 of rainfall. In all stands, fine root morphological parameters changed markedly from the topsoil to the lower profile; differences in fine root morphology among the six stands, however, were remarkably small. Two parameters, the necromass/biomass ratio and fine root tip density (tips per soil volume), however, were both much higher in acidic than basic soils. We conclude that variation in soil acidity and fertility only weakly influences fine root system size and morphology of F. sylvatica, but affects root system structure and, probably, fine root mortality. It is hypothesized that high root tip densities in acidic infertile soils compensate for low nutrient supply rates, and large necromasses are a consequence of adverse soil chemical conditions. Data from a literature survey support the view that rainfall is another major environmental factor that influences the stand fine root biomass of F. sylvatica.