Comparative studies on the mycorrhization of Larix decidua and Picea abies by Suillus grevillei

Summary Mycorrhization of Picea abies has been achieved, for the first time, with six strains of Suillus grevillei by a new culture method, using activated charcoal paper and liquid medium as a substrate. Mycorrhization of P. abies and Larix decidua was compared, and the process was found to be significantly different in the two tree species. S. grevillei is not incompatible with P. abies, but it forms mycorrhizae more readily with L. decidua. Hyphal growth was clearly stimulated on the surface of roots of Larix but retarded on Picea. A well organized Hartig net was formed with both tree species, but wall protuberances were frequently observed on the outer cell walls of Picea cortex cells when the Hartig net was not fully developed. No conspicuous cell wall reactions occurred in Larix roots. Cell wall protuberances may be comparable to those in transfer cells and are interpreted as an alternative to Hartig net development. Anatomical differences between roots of Larix and Picea, and physiologically active substances such as recognition factors on the root surfaces, are discussed with respect to their responsibility for the different reactions of S. grevillei.     

Structure of a pristine Picea abies forest in northeastern Europe

Abstract. The forest structure in a large, relatively homogeneous area of pristine Picea abies taiga in the southern boreal region west of the Ural mountains was studied along four 500-m long transects. P. abies dominated the forest in association with Abies sibirica and Betula spp. The mean volume of living trees was 216 m³/ha. This value varied among the four transects, from 138 - 252 m³/ha. Mean density of trees (DBH > 1 cm) (and variation over the transects) was 2 064/ha (1670 - 2710). Living trees classified as dying or seriously damaged made up 2.9 (2.5 - 3.5) % of the living tree volume. Betula was an important canopy component and made up 16% of the living tree volume. The number of dead standing trees varied from 195 - 325/ha, corresponding to a volume of 10.8 - 70.7 m³/ha. The density of trees with a broken stem was 90 - 170/ha and their estimated volume 7.6 - 41.3 m³/ha. Standing dead trees and trees with broken stems represented 10.4 and 8.9% of the total standing tree volume (living + dead), respectively. The mean volume of decaying logs on the forest floor was 117 (84.4–156.2) m³/ha, corresponding to 54 (35 - 113) % of the living tree volume. The canopy-forming trees were present in the understory as abundant saplings and suppressed individuals. The size distribution of the dominant tree species resembled the inverse J-shape. Generally, the forest was characterized by a high small-scale structural variation and a larger-scale relative homogeneity. This pattern is consistent with forest dynamics where the forest consists of a small-scale mosaic of patches in different stages of recovery following disturbance. Our results suggest that the ecology and dynamics of extensive areas of natural boreal forests can be driven by small-scale disturbance.     

Availability of coarse woody debris in a boreal old‐growth Picea abies forest

Abstract. This study reports temporal (based on cross‐dated dead trees) and spatial patterns of availability of coarse woody debris (CWD) from Picea abies in a Swedish boreal landscape with discrete old‐growth forest patches in a wetland matrix. Data were collected from 29 patches ranging in size from 0.3 to 28 ha. A total of 897 dead trees with a minimum diameter of > 15 cm occurred on the 7.2 ha area analysed. The year of death was established for 50% of these trees. CWD volume ranged from 17 to 65 m³/ha for downed logs and from 0.5 to 13 m³/ha for standing snags. CWD of all decay stages and diameter classes occurred abundantly and the probability of finding logs of all decay stages and sizes was very high at the scale of single hectares. Tree mortality differed among 5 yr periods. However, during the last 50 yr no 5 yr period produced less than 3 logs/ha. Decay rates were highly variable among different logs. Logs with soft wood and some wood pieces lost (decay stage 5) died ca. 34 years ago. This suggests a fairly rapid decay in this northern forest. The data indicate a high and continuous availability of CWD of all types. It is likely, therefore, that selection pressures for efficient dispersal among CWD dependent species may not be very high. Consequently, species with narrow habitat demands and/or low dispersal ability may have evolved and this may contribute to the decrease of certain species in the managed landscape.     

Gap structure,disturbance and regeneration in a primeval Picea abies forest

We investigated patterns of disturbance and recovery in Fiby urskog, a primeval spruce (Picea abies) forest, situated south of the border between the Boreo-nemoral and Boreal regions in East-central Sweden. The main types of disturbances are storm damage, fungal infection and insect attacks. The response of the different tree species varied and the mode of tree-fall depended on the different combinations of disturbance agents. The DBH distributions of gap creators and gap-border trees were almost the same. There was a high age diversity (100–240 yr) among the fallen trees. We concluded that all canopy trees (DBH > 20 cm) had the same probability of being felled by storms, irrespective of their age and DBH. According to an estimate along transect lines, gaps made up 31% of the spruce forest area. Individual gap sizes ranged from 9 m² to 360 m², but 83% of the gaps were < 150 m². The varied age structure of logs in individual gaps indicated that gap enlargements were common. 96 tree-falls were observed on four days with an hourly mean wind speed > 12.0 m/s; all trees fell in the direction of the wind. However, when we consult the 30-yrrecord(l 959–1989)ofthemeanhourly wind speed >12.0 m/s, it is clear that the pattern of storm-directions does not match the pattern of orientation of fallen logs. The present disturbance regime and the predominance of small gaps were more favourable for the regeneration oí Picea abies than of light-demanding tree species. In one large, 2900 m² gap, not crossed by the transects, all the major tree species had established within 7 yr, suggesting that classical succession in the sense of complete species replacement or ‘relay floristics’ didnot occur. Our observations seem rather to fit the ‘initial floristic’ model. Estimates of turnover time ranged from 170 to 228 yr, depending on the method used.     

Revegetation following experimental disturbance in a boreal old-growth Picea abies forest

Abstract. We studied revegetation patterns after experimental fine-scale disturbance (e.g. uprooting) in an old-growth Picea abies forest in southeastern Norway. An experimental severity gradient was established by manipulation of the depth of soil disturbance; two types of disturbed areas were used. Species recovery was recorded in the disturbed patches in three successive years after disturbance. The cover of vascular plants and, even more so the cover of bryophytes and lichens, recovered slowly after disturbance. The least severe treatments (removal of vegetation and removal of vegetation and the litter layer) was followed by the fastest recovery. The mean number of vascular plant species was usually higher three years after disturbance than before disturbance, while the opposite was true for bryophytes. Several vascular plant species that were abundant in intact forest floor vegetation (Vaccinium myrtillus, V. vitis-idaea and Deschampsia flexuosa) recovered during a three-year period primarily by resprouting from intact rhizomes and clonal in-growth. Other important recovery mechanisms included germination from a soil-buried propagule bank (e.g. Luzula pilosa, Plagiothecium laetum agg., Pohlia nutans and Polytrichum spp.) and dispersal of propagules into the disturbed patches (e.g. Betula pubescens and Picea abies). Microsite limitation seemed to occur in several species that were abundant in the soil propagule bank (e.g. the ferns Athyrium filix-femina, Gymnocarpium dryopteris and Phegopteris connectilis) but which did not appear in disturbed patches. Disturbance severity influenced revegetation patterns, recorded both as trajectories of vegetation composition in a DCA ordination space and as change in floristic dissimilarity. The length of the successional path (compositional change measured in β-diversity units) increased with increasing disturbance severity, and was also influenced by the area of the disturbed patch and the distance to intact vegetation. The rate of succession depended on the method by which it was measured; decreasing year by year in floristic space, while first decreasing and then increasing in ordination space. The reason for this difference is explained.     

50 years of change in a Swedish boreal old-growth Picea abies forest

Abstract. Changes in the tree layer (> 1.3m) and sapling layer (< 1.3m, including seedlings) of a Swedish boreal old-growth Picea abies (Norway spruce) forest from the 1930s to the 1980s were studied in permanent plots. The plots were established in 1938–1939 and re-analysed in 1983–1988. Regeneration, mortality, turnover rate in the tree layer and amount of decomposing logs as well as the time required for complete decomposition of logs were investigated using the detailed data from the 1930s. Ca. 25 % of the trees present during the first analysis were no longer alive. This mortality was balanced by recruitment from the sapling layer. The rate of mortality suggests a turnover time for the tree layer of ca. 200 yr. The number of spruces in the sapling layer has increased by ca. 85 %, hypothetically in response to an increase in amount of decomposing wood that can serve as nurse logs and stumps. The mean time for total decomposition was calculated as ca. 200 yr. Spruce regeneration on logs does not occur until the log is at least ca. 50 yr old. The survival pattern in the sapling layer suggests a high mortality rate at the seedling stage (≤ 1 yr) and a low mortality rate at the sapling stage. In conclusion, it is suggested that the amount of coarse woody debris available for regeneration, the occurrence of seedlings, and seedling mortality constitute concurrent factors through which climatic fluctuations, in a long-term perspective, direct stand recruitment and density. As a consequence, these boreal forests will be kept in a dynamical equilibrium.     

Structure and regeneration patterns in a virgin Picea abies forest in northern Sweden

Abstract. A high-altitude boreal Picea abies forest, with tree ages of up to 410 yr, was studied with respect to age structure, spatial regeneration patterns, and substrate. The results suggest that recruitment is primarily dependent on germination substrate but also negatively correlated with the density of the tree layer. 60 % of all spruces < 1.3 m high grew on substrates connected with tree-fall; ca. 40 % were found on decomposing logs and stumps, covering only ca. 6 % of the forest floor. Individual logs remain important as a regeneration substrate for ca. 150 yr. Continuous presence of decomposing coarse wood is a condition for the maintenance of the population structure under the prevailing climatic conditions. Peaks in the age distribution (the 1870's and the 1940's - 1950's) are probably climatically induced. The results challenge the previous assumptions that this kind of forest undergoes cyclic development. Long-term structural stability with climatically induced minor variations may be an alternative model.     

Effects of aluminium on growth and nutrient uptake of small Picea abies and Pinus sylvestris plants

Summary The effects of aluminium concentrations between 0.2 and 30 mM at pH 3.8 ±0.2 on small plants of Norway spruce [(Picea abies (L.) Karst], Scots pine (Pinus sylvestris L.), and Scots pine infected with the ectomycorrhizal fungus Suillus bovinus (L. ex Fr.) O. Kuntze were investigated. The plants were grown at maximum relative growth rate (R_G % day^–1) with free access but very low external concentrations of nutrients. Steady-state conditions with respect to relative growth rate (R_G) and internal nutrient concentrations were achieved before addition of aluminium, which was added as AlCl₃ and/or Al(NO₃)₃. There were reductions in rg at aluminium concentrations of 0.3 mM in spruce, 6 mM in pine and 10 mM in ectomycorrhizal pine, i. e. at aluminium concentrations considerably higher than those normally occurring in the top layer of the mineral soil where most fine roots are found. Nutrient uptake rate per unit root growth rate was calculated for different nutrient elements. The uptake rate of calcium and magnesium was reduced at aluminium concentrations of 0.2 mM (spruce), 1 mM (pine) and 3 mM (ectomycorrhizal pine), without influencing R_g. The results question the validity of the hypothesis of aluminium toxicity to forest tree species at low external concentrations.     

Differential energization of the tonoplast in suspension cells and seedlings from Picea abies

 Vacuolar ATPase (EC 3.6.1.3) and PPase (EC 3.6.1.1) were studied in suspension cells and seedlings from spruce [Picea abies (L.) Karst. Proton transport activity and uncoupler (1 μM nigericin) stimulated substrate hydrolysis were measured in tonoplast enriched membrane vesicles. In suspension cells the vacuolar PPase exhibited 1.8-fold activity of the ATPase. In roots and needles from 12-week-old spruce seedlings the vacuolar PPase was inactive, whereas the ATPase was active. Therefore, we investigated whether the preparation of spruce tonoplast vesicles from roots and needles inactivates the vacuolar PPase but not the ATPase. For this purpose, maize (Zea mays L.) tonoplast membranes exhibiting vacuolar PPase as well as ATPase activity were used as a probe and added to the homogenization medium prior to the preparation of spruce vesicles. The preparation of spruce vesicles was more inhibitory to the vacuolar ATPase than to the PPase. The comparison of vacuolar PPases from spruce suspension cells and maize roots revealed similar enzymatic properties. After isopycnic centrifugation on continuous sucrose gradients the vacuolar PPase from spruce suspension cells co-purified with the vacuolar ATPase. Together, these data show: (1) vacuolar PPases from spruce suspension cells and maize roots are similar, (2) the preparation of tonoplast vesicles from spruce roots and needles does not inactivate the vacuolar PPase, (3) tonoplasts of suspension cultured cells and seedlings from spruce are differentially energized by the vacuolar pyrophosphatase that may indicate a difference in pyrophosphate metabolism between embryogenic and differentiated spruce cells, and (4) tonoplast vesicles from spruce seedlings may allow investigations of the effect of pyrophosphate on the vacuolar ATPase in the absence of vacuolar PPase activity. Received: 2 July 1998 / Accepted: 14 September 1998     

Effects of liming and boron fertilization on mycorrhizas of Picea abies

Effects of liming and B fertilization on Norway spruce [Picea abies (L.) Karst.] mycorrhizas were studied in factorial field experiments. The lime was applied twice, about 30 years and 12 years before sampling (2000 and 4000 kg ha^-1 dolomite). B was applied at the rate of 1.5 kg B ha^-1 two years before sampling.Boron fertilization doubled the number of root tips in the top 10 mm of the humus layer. The proportion of dead short root tips was increased from 10 % in control plots to 29 % in the limed plots. Numbers of dead root tips were increased when both lime and B were applied. The % of mycorrhizas with external mycelium was slightly increased and the % of Piloderma croceum Erikss. and Hjortst. was decreased by lime. In conclusion, adverse effects of lime on mycorrhizas were found, which were ameliorated by B fertilization, but lime-induced B deficiency alone was not the only reason for the effects of lime on root mortality.     

Effects of soil pH and calcium on mycorrhizas of Picea abies

The effects of lime, increased soil pH and increased soil Ca concentration on the mycorrhizas of Norway spruce. [Picea abies (L.) Karst.] were studied independently of each other to elucidate the different mechanisms through which lime may influence mycorrhizas in acidic soil. In a field experiment (mature Norway spruce in podzol), lime was applied as CaCO₃; increased Ca concentration without an increase in pH was achieved with CaSO₄; and soil pH was increased without calcium by means of Na₂CO₃ and K₂CO₃ (Na+K treatment). Treatments were done in October, and mycorrhizas were counted from samples collected in the following June and September. All treatments increased the percentage of dead short root tips compared to controls in September, and Na+K already in June. Cenococcum geophilum Fr. increased in proportion in plots treated with Na+K.In a sand culture experiment, Norway spruce seedlings were grown from seed and inoculated with Cenococcum geophilum, or root inoculum, or left uninoculated. When mycorrhizas were beginning to form, CaCO₃ and CaSO₄ treatments were applied. Six weeks later, the percent of dead short root tips in both salt treatments was significantly increased from control, but formation of mycorrhizas was not inhibited by treatments.As all the treatments increased the proportion of dead short root tips, it is concluded that lime directly and adversely affected mycorrhizas of Norway spruce in sand culture and in mor humus. Both increased ionic strength and increased pH may be reasons for this rather than Ca²⁺ specifically.     

Ethylene effects on cambial activity and cell wall formation in hypocotyls of Picea abies seedlings

Ethylene regulation of cell division in the vascular cambium and cell wall formation was studied in hypocotyls of Norway spruce ( Picea abies [L.] Karst.) seedlings. Cuttings from 6-week-old seedlings were placed in water culture to which compounds affecting the synthesis and action of ethylene were added. After a 3-week treatment period, growth, ethylene production, morphology and cell wall composition of the hypocotyls were determined. Addition of high concentrations of the potent ethylene releasing agent 2-chloroethylphosphonic acid (ethrel), which increased ethylene emission by more than twice compared to control plants, inhibited the expansion of xylem cells while stimulating the incorporation of cell wall material, especially cellulose. Addition of small amounts of ethrel, which slightly stimulated ethylene emission, led to increases in the size of xylem cells, the amount of phloem tissue and the number of intercellular spaces in the cortex, and thus to increased hypocotyl diameter. However, no significant change in cell wall composition was detected. When ethylene production was decreased by adding Co²⁺ to the nutrient solution, differentiation of new xylem was disturbed, but the rate of cell division was not affected. Although the incorporation of cell wall material was inhibited, the proportions of lignin and cellulose in the wall appeared to remain unchanged. Silver ions stimulated the expansion of both xylem and cortex cells, but had no significant effect on cell wall formation. We conclude that ethylene has a role in regulating the incorporation of wall carbohydrates.     

Response of fine roots to an experimental gap in a boreal Picea abies forest

We examined the initial response of the quantity and distribution of fine roots to the creation of an experimental canopy gap with a diameter of 50 m in a mature managed Norway spruce forest. Under the canopy, the fine root length densities of trees, shrubs, and grasses and herbs were 3207, 707 and 2738 m m^–2, respectively. The fine root biomass of trees, shrubs, and grasses and herbs were 182, 47 and 52 g m^–2, respectively. Two growing seasons after gap creation hardly any fine tree roots were found in the middle part of the gap. The living tree roots in the gap edge zone were mainly located within a 5-m distance from the standing edge trees. The indices developed here to show the influence of trees on fine root lenght density clearly revealed the effect of the vicinity of living trees on fine root lenght density. The root densities of grasses, herbs and dwarf shrubs did not show a clear response to gap creation despite the increase of their foliage. Our results suggest that in boreal spruce forests a gap disturbance creates a distinct tree root gap and that the gap edge trees do not extend their root systems rapidly into the formed root gap.     

Establishment of Picea abies seedlings in a central European mountain grassland: an experimental study

Abstract. Secondary meadows in the Kru?né hory mountains, central Europe, are not colonized by woody species, although they have been abandoned for half a century and are surrounded by Picea abies (Norway spruce) forests. The causes of inhibition of establishment of Norway spruce seedlings in the meadows were tested experimentally. The experiment was started in a masting year to ensure sufficient seed input. Four treatments (vegetation cut; vegetation cut and litter removed; all above-ground biomass and topsoil removed; control) were combined in a factorial design. The effect of browsing, mainly by deer, was assessed by fencing half of the experimental plots. Seedling establishment differed among treatments (all pair-wise differences were significant) but was independent of fencing. In contrast, seedling survival was influenced both by the treatment and fencing. At the end of the second year, only the plots with all biomass and organic topsoil removed supported viable populations of spruce seedlings. Dense herb cover and a thick layer of slowly decaying litter are considered the main factors inhibiting the establishment of woody species.     

Chlorophyll formation and the development of photosynthesis in dark grown seedlings of Picea abies

Abstract Seeds of Picea abies were germinated and grown in either darkness or constant light. The chlorophyll content and photosynthetic carbon dioxide uptake of developing seedlings of different ages was determined. Ten-day-old dark grown seedlings showed an instant ability for photosynthetic carbon dioxide uptake and also formed further chlorophyll most rapidly upon subsequent illumination. These activities progressively diminished when the dark growth period was extended. Light grown seedlings reached a maximum chlorophyll level after 15 days growth, and this preceded maximal photosynthetic development.