Vergleichende Feinwurzeluntersuchungen an gesunden und erkrankten Altfichten auf zwei Standorten in Bayernein Beitrag zur Waldsterbensforschung

Fine root studies in two old growth spruce forests affected by “Wadsterben” in Bavaria. Fine roots of healthy and declining Norway spruce (Picea abies) were investigated. We observed fewer active myeorrhizac associated with affected spruce in two old growth Forests. It has been hypothesised that air pollution is dimaging the needles resulting in less photosynthate and that the reduction of active mycorrhizae is due to such an initial weakening of the tree.     

Analysis of Norway spruce dieback phenomenon in Latvia – a belowground perspective

In 2010, dieback of Norway spruce (Picea abies) was observed in Latvia. As mostly stands on seasonally or permanently water-saturated soils were damaged, we hypothesized that the recorded foliar damage might be associated with belowground factors. We established 48 sample plots in stands with damage symptoms for soil parameter evaluation and fine root sampling to determine the number of fine roots in different growth stages and to describe the associated fungal community. Among sample plots on organic soils, there were fewer viable older fine roots and greater number of dead fine roots in more damaged sites than in less damaged. These root parameters were significantly correlated with higher groundwater level and soil pH. The fungal community was dominated by ectomycorrhizal (ECM) species (the most common was Tylospora asterophora), the endophyte Oidiodendron maius and saprotrophic species Cryptococcus magnus. Saprotrophic species had higher abundance in more damaged sites, suggesting a shift in fungal communities from ECM fungi. In conclusion, the results of this work suggest Norway spruce root system response to short-term climatic stress, which should be considered when planning spruce forest management, especially on organic soils with a high groundwater level.     

Intracellular infection by fungi in mycorrhizae of damaged spruce trees

Intracellular infection was found in the vascular tissue and meristem of mycorrhizae of a severely damaged stand of spruce and fir trees in the Northern Black Forest. Two fungal species could be isolated from surface sterilized mycorrhizae: Mycelium radicis atrovirens and Cryptosporiopsis cf. abietina. Infection tests with spruce seedlings revealed that mainly Cryptosporiopsis cf. abietina must be held responsible for infection of the vascular tissue. Less infectious was Mycelium radicis atrovirens, which infected only cortex cells.     

Comparative studies on the fine root status of healthy and declining spruce and beech trees in the Bavarian Alps and occurrence of Phytophthora and Pythium species

Investigations on root and crown status of spruce and beech were carried out on selected trees in the ‘Werdenfelser Land’ area (Bavarian Alps, Southern Germany). In addition, the association of fine root pathogens of the genera Phytophthora and Pythium with the trees’ rhizosphere was studied. In a variety of stands representing various site conditions, soil and root samples were taken from 12 spruce (Picea abies) pairs and eight beech (Fagus sylvatica) pairs. Each pair consisted of a healthy and a declining tree as indicated by crown transparency. The root status was characterized using a set of parameters, and correlations between crown and fine root status were observed. In spruce, most parameters decreased significantly with increasing crown transparency, whereas in beech, correlations were less pronounced. The total number of lateral roots per cm small root (diameter 2–5 mm) was significantly lower in both species for declining trees compared to healthy trees. Pythium spp. were isolated from 15 of 24 soil samples taken from under spruce, and from eight of 16 samples from under beech. Phytophthora citricola was found in two beech stands only. Among the isolated species, Pythium anandrum, Pythium inflatum and Pythium acrogynum were identified according to morphological features. After polymerase chain reaction‐restriction fragment length polymorphism analysis, residual Pythium isolates were assigned to four different groups. No crown transparency‐dependent differences in isolation frequency were found. In soil infestation tests, all species tested caused root damage on both young spruce and beech plants, with P. citricola being the most aggressive pathogen. Additionally, Pythium‐infected beeches showed severe leaf chloroses and necroses. Due to their low isolation frequency, Phytophthora spp. are not considered to play a major role in the decline of spruce and beech in the investigated area. Pythium spp., however, were isolated frequently, showed pathogenicity towards the fine roots of spruce and beech, and are therefore considered to be at least contributing factors in the decline of Bavarian mountain forests.     

Fine root biomass in relation to site and stand characteristics in Norway spruce and Scots pine stands

Variations in fine root biomass of trees and understory in 16 stands throughout Finland were examined and relationships to site and stand characteristics determined. Norway spruce fine root biomass varied between 184 and 370 g m(-2), and that of Scots pine ranged between 149 and 386 g m(-2). In northern Finland, understory roots and rhizomes (< 2 mm diameter) accounted for up to 50% of the stand total fine root biomass. Therefore, the fine root biomass of trees plus understory was larger in northern Finland in stands of both tree species, resulting in a negative relationship between fine root biomass and the temperature sum and a positive relationship between fine root biomass and the carbon:nitrogen ratio of the soil organic layer. The foliage:fine root ratio varied between 2.1 and 6.4 for Norway spruce and between 0.8 and 2.2 for Scots pine. The ratio decreased for both Norway spruce and Scots pine from south to north, as well as from fertile to more infertile site types. The foliage:fine root ratio of Norway spruce was related to basal area and stem surface area. The strong positive correlations of these three parameters with fine root nitrogen concentration implies that more fine roots are needed to maintain a certain amount of foliage when nutrient availability is low. No significant relationships were found between stand parameters and fine root biomass at the stand level, but the relationships considerably improved when both fine root biomass and stand parameters were calculated for the mean tree in the stand. When the northern and southern sites were analyzed separately, fine root biomass per tree of both species was significantly correlated with basal area and stem surface area per tree. Basal area, stem surface area and stand density can be estimated accurately and easily. Thus, our results may have value in predicting fine root biomass at the tree and stand level in boreal Norway spruce and Scots pine forests.     

Intersterilitätsgruppen und Klone von Heterobasidion annosum-Isolaten aus Koniferen- Wurzel- und -Stammfäulen

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.     

Moderate drought alters biomass and depth distribution of fine roots in Norway spruce

A rain shelter experiment was conducted in a 90‐year‐old Norway spruce stand, in the Kysucké Beskydy Mts (Slovakia). Three rain shelters were constructed in the stand to prevent the rainfall from reaching the soil and to reduce water availability in the rhizosphere. Fine root biomass and necromass were repeatedly measured throughout a growing season by soil coring. We established the quantities of fine root biomass (live) and necromass (dead) at soil depths of 0–5, 5–15, 15–25 and 25–35 cm. Significant differences in soil moisture contents between control and drought plots were found in the top 15 cm of soil after 20 weeks of rainfall manipulation (lasting from early June to late October). Our observations show that even relatively light drought decreased total fine root biomass from 272.0 to 242.8 g m^?2 and increased the amount of necromass from 79.2 to 101.2 g m^?2 in the top 35 cm of soil. Very fine roots (VFR), that is, those with diameter up to 1 mm, were more affected than total fine roots defined as 0–2 mm. The effect of reduced water availability was depth‐specific; as a result, we observed a modification of vertical distribution of fine roots. More roots in drought treatment were produced in the wetter soil horizons at 25–35 cm depth than at the surface. We conclude that fine and VFR systems of Norway spruce have the capacity to re‐allocate resources to roots at different depths in response to environmental signals, resulting in changes in necromass to biomass ratio.     

Intersterilitätsgruppen und Klone von Heterobasidion annosum in einem 31jährigen Fichtenbestand1

Intersterility groups and clones of Heterobasidion annosum in a 31 years old Norway spruce stand . H. annosum isolates from roots of 13 spruce trees growing in close neighbourhood be longed to the P group (from five roots) and to the S group (from 44 roots). The clone identifications revealed colonisation via root contacts.     

Interspecific competition impacts on the morphology and distribution of fine roots in European beech (Fagus sylvatica L.) and Norway spruce (Picea abies (L.) Karst.)

Morphology and vertical distribution patterns of spruce and beech live fine roots (diameter ≤2 mm) were studied using a soil core method in three comparable mature stands in the Solling: (1) pure beech, (2) pure spruce and (3) mixed spruce–beech. This study was aimed at determining the effects of interspecific competition on fine root structure and spatial fine root distribution of both species. A vertical stratification of beech and spruce fine root systems was found in the mixed stand due to a shift in beech fine roots from upper to lower soil layers. Moreover, compared to pure beech, a significantly higher specific root length (SRL, P<0.05) and specific surface area (SSA, P<0.05) were found for beech admixed with spruce (pure beech/mixed beech SRL 16.1–23.4 m g⁻¹, SSA 286–367 cm² g⁻¹). Both indicate a flexible ‘foraging’ strategy of beech tending to increase soil exploitation and space sequestration efficiency in soil layers less occupied by competitors. Spruce, in contrast, followed a more conservative strategy keeping the shallow vertical rooting and the root morphology quite constant in both pure and mixed stands (pure spruce/mixed spruce SRL 9.6/7.7 m g⁻¹, P>0.10; SSA 225/212 cm² g⁻¹, P>0.10). Symmetric competition belowground between mixed beech and spruce was observed since live fine roots of both species were under-represented compared to pure stand. However, the higher space sequestration efficiency suggests a higher competitive ability of beech belowground.     

Effects of slash retention and wood ash addition on fine root biomass and production and fungal mycelium in a Norway spruce stand in SW Sweden

In the study reported here we examined the short-term effects (1–3 years) of slash retention (SR) and the long-term effects (13–15 years) of wood-ash application (A) on fine roots and mycorrhizae in a 40-year-old Norway spruce forest in southwest Sweden. Soil cores were used to obtain estimates of the biomass (g m⁻²) of roots in three diameter classes (<0.5, 0.5–1 and 1–2 mm), root length density (RLD), specific root length (SRL) and mycorrhizal root tip density (RTD). Fine root (<1 mm) length production and mortality, and mycelium production, were estimated using minirhizotron and mesh bag techniques, respectively. Compared with the control plots (C), the biomass of fine roots in diameter classes <0.5 mm and 0.5–1 mm was significantly higher in A plots, but lower in SR plots. In addition, RLD was significantly lower in the humus layer of SR plots than in the humus layers of C and A plots, but not in the other layers. None of the treatments affected the SRL. In all soil layers, the SR treatment resulted in significant reductions in the number of ectomycorrhizal root tips, and the mycelia production of fungi in mesh bags, relative to the C treatment, but the C and A treatments induced no significant changes in these variables. Fine root length production in the C, A and SR plots amounted to 94, 87 and 70 mm tube⁻¹ during the 2003 growing season, respectively. Fine root mortality in treated plots did not change over the course of the study. We suggest that leaving logging residues on fertile sites may result in nitrogen mineralisation, which may in turn induce reductions in root biomass, and both root and mycelium production, and consequently affect nutrient uptake and the accumulation of organic carbon in soil derived from roots and mycorrhizae.     

Root distribution of under-planted European beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.) below the canopy of a mature Norway spruce stand as a function of light

Aboveground and belowground biomass of 15-year-old under-planted European beech seedlings (Fagus sylvatica L.) in Norway spruce stand were studied along a light gradient in three plots, in the northern part of Slovenia. Differences in soil water content, aboveground and fine root biomass distribution were confirmed between studied plots. Light had significant effect on the total biomass, root-shoot ratio (0.388 ± 0.076 under canopy, 0.549 ± 0.042 in the edge, 0.656 ± 0.047 in the open), specific root length (SRL) of fine beech roots (561.9 ± 42.2 under canopy, 664.3 ± 51.2 in the edge, 618.2 ± 72.8 in the open) and specific leaf area in beech, indicating morphological adjustment to shade. However, SRL of beech fine roots indicated no change between plots. The correlation between total aboveground and root biomass and light below the mature stand canopy was higher in the case of diffuse light intensity. Most fine roots of spruce were concentrated in the top (0–20 cm) soil layer. Beech fine roots under canopy and edge conditions were also concentrated in top (0–20 cm) soil layer and exhibited shift downwards to deeper soil horizons in open plot. Root proportion between beech and spruce changed with light toward beech with increasing light intensity for both fine and coarse roots.     

Zur Anatomic gesunder und “kranker” Feinstwurzeln von Fichten: Meristem und Differenzierungen in Wurzelspitzen und Mykorrhizen

On the anatomical features of well functioning and “damaged” rootlets of spruce: meristem and differentiation of root apices and mycorrhizae. The root apices of various developmental stages of Picea abies (L.) Karst. are described in detail and the differences between long- and short roots discussed.     

The effect of nitrogen fertilization on fungistatic phenolic compounds in roots of beech (Fagus sylvatica) and Norway spruce (Picea abies)

The effect of nitrogen fertilization on fungistatic phenolic compounds in fine roots of beech and Norway spruce growing in afforestation plots was analysed. The plots were situated at two sites in Switzerland on acidic soil with low base saturation. For 9 years, the trees have been treated with dry ammonium nitrate to give 0, 10, 20, 40, 80, 160 kg N ha^?1 year^?1, respectively. The phenolic compounds responded differently to fertilization. Fine roots of beech showed a significant decrease of (?)‐epicatechin and piceatannol with increasing nitrogen fertilization. The concentration of protocatechuic acid was increased with fertilization. Roots of fertilized Norway spruce showed significantly decreased concentrations of 4‐hydroxyacetophenone and piceatannol. The mycelial growth of three isolates each of Heterobasidion annosum s.l. and Cylindrocarpon destructans was tested on agar media containing various phenolic compounds in concentrations found in fine roots of Norway spruce (Picea abies) and beech (Fagus sylvatica). All three H. annosum isolates were inhibited by p‐coumaric acid and (?)‐epicatechin. Two isolates were inhibited by another four phenolic compounds (p‐hydroxybenzoic acid, 4‐hydroxyacetophenone, piceatannol and protocatechuic acid), one by (+)‐catechin. Two of three C. destructans isolates were inhibited by all phenolic compounds except for (+)‐catechin which affected only one isolate, one isolate did not respond at all.     

Long‐term effects of liming on the fine‐root standing crop of Picea abies and Pinus sylvestris in relation to chemical changes in the soil

The long‐term effects of lime application on fine roots of Norway spruce, Picea abies (L.) Karst, and Scots pine, Pinus sylvestris (L.), have been studied in five experimental forest stands subjected to different lime applications 5 to 18 years before the present study was undertaken. The effects of liming does not seem to significantly influence fine‐root development in forest stands in the long term. The only response to liming in measured root variables was a tendency to increased specific root length (SRL = fine‐root length/fine‐root dry weight, m/g). A correlation between increased SRL, decreased root biomass and increased stem volume growth was indicated. Changes in water extractable amounts of mineral elements—P, K, Ca, Mg, Mn, S, Al and Fe‐in bulk soil and rhizosphere soil from the mineral soil layers were studied in a control area and an area treated with 3830 kg CaCO₃ ha^‐1. Few significant differences were found between treatments, and then mainly in the case of Ca.     

Compaction of till soils and growth tests with Norway spruce and scots pine

Five of the most common soils in forest areas in Sweden, viz., coarse sand, fine sand, coarse sandy till, medium sandy till and fine sandy till soil, were compressed at different loadings for 10 min. In the soils studied, it was found that the more fine material there was in the soil, the easier it compacted, and that till soils compacted much more than sediments of the same main particle class. The chosen compression time seemed to correspond well with an equilibrium level of static compaction. Influence of wall friction in compression tubes was found to be negligible for till soils.After compaction the soil cores were sown with Norway spruce (Picea abies (L.) Karst.) or Scots pine (Pinus sylvestris L.) and 16–20 days later the root lengths were recorded. Lengths of primary roots in per cent of the control were found to be almost inverse to the compaction of the soils; Norway spruce was more impeded than Scots pine. Thus, the bulk density alone is a poor indicator of the impact of soil compaction. Growth to 26, 33 and 120 days showed that the extension of secondary roots might be more impeded by compaction than the primary roots. Fertilization increased growth but did not decrease the growth difference between seedlings on unloaded and loaded soil cores.     

Laser microprobe mass analysis (LAMMA) of aluminum and lead in fine roots and their ectomycorrhizal mantles of Norway spruce (Picea abies (L.) Karst.)

Fine roots and ectomycorrhizal root tips were sampled in a Norway spruce (Picea abies (L.) Karst.) stand in the eastern part of the Belgian Ardennes. The cellular and partly subcellular localizations of aluminum and lead were identified by the micro-analytical laser microprobe mass analysis (LAMMA) technique. In fine roots with secondary structure, localization of aluminum was limited to the peripheral cell layers. Lead was found in the outer layers, and also in the primary phloem. Aluminum penetrated the mycorrhizal mantle, but lead was seldom detected in ectomycorrhizae.     

Estimating “autotrophic” belowground respiration in spruce and beech forests: decreases following girdling

Seasonal fluxes of CO₂ from soil and the contribution of autotrophic (root + mycorrhizal) to total soil respiration (SR) were estimated for a mixed stand of European beech (Fagus sylvatica) and Norway spruce (Picea abies) in Central Europe. Mature trees of each species were girdled in August 2002 to eliminate carbohydrate allocation to roots. SR was measured at distances of 0.5, 1.0, and 1.5/2.0 m from the bole of each tree at 1–2 weeks intervals throughout the fall of 2002 and monthly during the spring and summer of 2003. The contribution of roots and mycorrhizae to total SR was estimated by the decrease in SR compared to ungirdled control trees to account for seasonal patterns evident in controls. SR decreased with soil temperature in the fall 2002 and increased again in 2003 as soil warmed. During most of the study period, SR was strongly related to soil temperature. During the dry summer of 2003, however, SR appeared to be uncoupled from temperature and was strongly related to soil water content (SWC). Mean rates of SR in beech and spruce control plots as well as root densities did not show a clear pattern with distance from the bole. SR decreased to levels below controls in beech within a few days after girdling, whereas spruce did not show a significant decrease until October 2002, 6 weeks after girdling. In both beech and spruce, decreased SR in response to girdling was greatest closest to the bole, possibly reflecting increased mycorrhizal activity close to the bole. Autotrophic respiration was estimated in beech to be as much as 50% of the total SR in the stand. The contribution of autotrophic respiration was less certain for spruce, although close to the bole, the autotrophic fraction may contribute to total SR as much as in beech. The large fraction of autotrophic respiration in total SR requires better understanding of tree level stresses that affect carbon allocation below ground.     

Changes in fine root production and longevity in relation to water and nutrient availability in a Norway spruce stand in northern Sweden

Effects of irrigation and liquid fertilization on fine root (< 1 mm) production and longevity, and fine root (< 0.5-2 mm) biomass were studied in a Norway spruce (Picea abies (L.) Karst.) stand in northern Sweden. Fine root length production and longevity were measured by the minirhizotron technique at 0-10 cm depth in the following treatments: irrigation (I), liquid fertilization (IL) and control (C). Standing root biomass and root length density (RLD) were studied in the litter-fermented humus (LFH) layer and at depths of 0-10, 10-20 and 20-30 cm using soil cores in solid fertilized (F) and C plots. Minirhizotrons were installed in October 1994 and measurements recorded monthly from July to September 1995 and during the growing season in 1996. Soil cores were sampled in 1996. Fine root production increased significantly in IL plots compared with C plots, but the I treatment did not increase root production. Root mortality increased significantly in IL plots compared with C plots. Fine root longevity in IL plots was significantly lower compared with C and I plots. No significant difference was found between longevity of fine roots in I and C plots. Compared with C, F treatment increased fine root biomass in the LFH and mineral soil layers, and increased the amount of fine roots in mineral soil layers relative to the LFH layer. Furthermore, F increased RLD and the number of mycorrhizal root tips significantly.     

Effect of Thinning and Phlebiopsis gigantea Stump Treatment on the Growth of Heterobasidion parviporum Inoculated in Picea abies

The spread of Heterobasidion parviporum Niemelä & Korhonen in roots of Norway spruce was studied in three unthinned first rotation stands of Norway spruce [Picea abies (L.) Karst.] on former agricultural land in south-western Sweden. Heterobasidion parviporum was inoculated at stump height into the trunk of 135 standing trees in a randomized block design. One year after inoculation, two-thirds of the trees were thinned out and one-third was left standing. Half of the stumps left by thinning were treated with spores of Phlebiopsis gigantea (Fr.) Jül and half were left untreated. The spread of H. parviporum was examined both 3 and 5 yrs after inoculation. The rate of spread of H. parviporum and the proportion of infected roots were found to be significantly higher in the root systems of the stumps than in those of the standing trees. Treatment with P. gigantea had no significant effect on the development of H. parviporum in the stumps. There was a tendency 5 yrs after inoculation, however, for a lower proportion of H. parviporum-infected roots in the stumps treated with P. gigantea than in the untreated stumps. In conclusion, thinning of infected Norway spruce was found to increase the rate of spread of H. parviporum in the root systems of the infected trees, which could increase the risk of a rapid build-up of infection in the remaining stand.     

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.