Ectomycorrhizal root growth in scots pine stands in response to manipulation of litter and humus layers

 The effect on ectomycorrhizal root growth in a nitrogen-enriched planted stand of Scots pine (Pinus sylvestris L.) on podzolic sandy soil to manipulation of litter and humus layers (removal, doubling and control treatments) was examined, and compared to ectomycorrhizal root growth in an untreated naturally established Scots pine stand on nutrient-poor non-podzolic sandy soil. Half a year after manipulation of litter and humus layers in the planted stand, ingrowth-cores to a depth of 60 cm were installed in both stands. Scots pine roots were sampled four times during two growing seasons. Ectomycorrhizal roots were found at all sampled soil depths to 60 cm in all plots. Root growth and ectomycorrhizal development were greater in the naturally established stand than in all plots in the planted stand. Numbers of ectomycorrhizal root tips in the litter and humus removal plots were generally higher than in the control plots in the planted stand until May 1992. Doubling litter and humus did not significantly affect root length or the numbers of ectomycorrhizal root tips. The N _dissolved , NH₄ ⁺ and NO₃ ^– concentrations and the organic matter content in the upper 5 cm of the mineral soil in the planted stand on podzolic sandy soil were generally higher and the pH significantly lower than in the naturally established stand on non-podzolic sandy soil. Root growth and ectomycorrhizal development in the secondary stand may have been negatively affected by the chemical composition of the podzolic sandy soil. Accepted: 19 March 1997     

Bacterial microbiomes of individual ectomycorrhizal Pinus sylvestris roots are shaped by soil horizon and differentially sensitive to nitrogen addition

Plant roots select non‐random communities of fungi and bacteria from the surrounding soil that have effects on their health and growth, but we know little about the factors influencing their composition. We profiled bacterial microbiomes associated with individual ectomycorrhizal Pinus sylvestris roots colonized by different fungi and analyzed differences in microbiome structure related to soils from distinct podzol horizons and effects of short‐term additions of N, a growth‐limiting nutrient commonly applied as a fertilizer, but known to influence patterns of carbon allocation to roots. Ectomycorrhizal roots growing in soil from different horizons harboured distinct bacterial communities. The fungi colonizing individual roots had a strong effect on the associated bacterial communities. Even closely related species within the same ectomycorrhizal genus had distinct bacterial microbiomes in unfertilized soil, but fertilization removed this specificity. Effects of N were rapid and context dependent, being influenced by both soil type and the particular ectomycorrhizal fungi involved. Fungal community composition changed in soil from all horizons, but bacteria only responded strongly to N in soil from the B horizon where community structure was different and bacterial diversity was significantly reduced, possibly reflecting changed carbon allocation patterns.     

Distribution of Cren- and Euryarchaeota in Scots Pine Mycorrhizospheres and Boreal Forest Humus

Archaeal 16S rRNA gene sequences have been found in a variety of moderate-temperature habitats including soil and rhizospheres. In this study, the differences of archaeal communities associated with Scots pine (Pinus sylvestris L.) short roots, different types of mycorrhizospheric compartments, and uncolonized boreal forest humus were tested by direct DNA extraction, polymerase chain reaction–denaturing gradient gel electrophoresis (PCR–DGGE), and sequencing. The results indicated that mycorrhizal colonization of Scots pine roots substantially influence the archaeal community of pine rhizospheres. Colonization of short roots by most mycorrhizal fungi tested increased both archaeal frequency and diversity. Most of the archaeal sequences encountered in mycorrhizas belonged to the phylum Euryarchaeota, order of Halobacteriales. The difference in archaeal diversity between the mycorrhizospheric compartments and humus was profound. Most compartments with fungal components contained euryarchaeotal 16S rRNA gene sequences, whereas a high diversity of crenarchaeotal sequences and no euryarchaeotal sequences were found in forest humus outside mycorrhizospheres.     

The influence of elevated CO2 and O3 on fine roots and mycorrhizas of naturally growing young Scots pine trees during three exposure years

Young Scots pine trees naturally established at a pine heath were exposed to two concentrations of CO₂ (ambient and doubled ambient) and two O₃ regimes (ambient and doubled ambient) and their combination in open-top field chambers during growing seasons 1994, 1995 and 1996 (late May to 15 September). Filtered ozone treatment and chamberless control trees were also included in the treatment comparisons. Root ingrowth cores were inserted to the undisturbed soil below the branch projection of each tree at the beginning of the fumigation period in 1994 and were harvested at the end of the fumigation periods in 1995 and 1996. Root biomasses were determined from different soil layers in the ingrowth cores, and the infection levels of different mycorrhizal types were calculated. Elevated O₃ and CO₂ did not have significant effects on the biomass production of Scots pine coarse (Ø > 2 mm) or fine roots (Ø < 2 mm) and roots of grasses and dwarf shrubs. Elevated O₃ caused a transient stimulation, observable in 1995, in the proportion of tuber-like mycorrhizas, total mycorrhizas and total short roots but this stimulation disappeared during the last study year. Elevated CO₂ did not enhance carbon allocation to root growth or mycorrhiza formation, although a diminishing trend in the mycorrhiza formation was observed. In the combination treatment increased CO₂ inhibited the transient stimulating effect of ozone, and a significant increase of old mycorrhizas was observed. Our conclusion is that doubled CO₂ is not able to increase carbon allocation to growth of fine roots or mycorrhizas in nutrient poor forest sites and realistically elevated ozone does not cause a measurable limitation to roots within a period of three exposure years.     

Carbon balance and allocation of assimilated CO2 in Scots pine, Norway spruce, and Silver birch seedlings determined with gas exchange measurements and 14C pulse labelling

Carbon dioxide is released from the soil to the atmosphere in heterotrophic respiration when the dead organic matter is used for substrates for soil micro-organisms and soil animals. Respiration of roots and mycorrhiza is another major source of carbon dioxide in soil CO₂ efflux. The partitioning of these two fluxes is essential for understanding the carbon balance of forest ecosystems and for modelling the carbon cycle within these ecosystems. In this study, we determined the carbon balance of three common tree species in boreal forest zone, Scots pine, Norway spruce, and Silver birch with gas exchange measurements conducted in laboratory in controlled temperature and light conditions. We also studied the allocation pattern of assimilated carbon with ¹⁴C pulse labelling experiment. The photosynthetic light responses of the tree species were substantially different. The maximum photosynthetic capacity (P _max) was 2.21 μg CO₂ s⁻¹ g⁻¹ in Scots pine, 1.22 μg CO₂ s⁻¹ g⁻¹ in Norway spruce and 3.01 μg CO₂ s⁻¹ g⁻¹ in Silver birch seedlings. According to the pulse labelling experiments, 43–75% of the assimilated carbon remained in the aboveground parts of the seedlings. The amount of carbon allocated to root and rhizosphere respiration was about 9–26%, and the amount of carbon allocated to root and ectomycorrhizal biomass about 13–21% of the total assimilated CO₂. The ¹⁴CO₂ pulse reached the root system within few hours after the labelling and most of the pulse had passed the root system after 48 h. The transport rate of carbon from shoot to roots was fastest in Silver birch seedlings.     

Short-term effects of manipulated increase in acid deposition on soil, soil solution chemistry and fine roots in Scots pine (Pinus sylvestris) stand on a podzol

A manipulated increase in acid deposition (15 kg S ha⁻¹), carried out for three months in a mature Scots pine (Pinus sylvestris) stand on a podzol, acidified the soil and raised dissolved Al at concentrations above the critical level of 5 mg l⁻¹ previously determined in a controlled experiment with Scots pine seedlings. The induced soil acidification reduced tree fine root density and biomass significantly in the top 15 cm of soil in the field. The results suggested that the reduction in fine root growth was a response not simply to high Al in solution but to the depletion of exchangeable Ca and Mg in the organic layer, K deficiency, the increase in NH₄:NO₃ ratio in solution and the high proton input to the soil by the acid manipulation. The results from this study could not justify the hypothesis of Al-induced root damage under field conditions, at least not in the short term. However, the study suggests that a short exposure to soil acidity may affect the fine root growth of mature Scots pine.     

Variations of bioavailable Sr concentration and 87Sr/86Sr ratio in boreal forest ecosystems

The mean depth of Sr and water uptake in mixed Norway spruce (Picea abies) and Scots pine (Pinus sylvestris) stands was investigated, using natural variations of ⁸⁷Sr/⁸⁶Sr and ¹⁸O/¹⁶O in soils in relation to depth. Three spruce-pine pairs were studied on a podzol and a peat site in Northern Sweden. Tree leaf and wood, as well as soils, soil solutions and roots below each tree were analysed for Sr and Ca concentrations and ⁸⁷Sr/⁸⁶Sr ratio. The ¹⁸O/¹⁶O ratio was also determined in xylem sap and soil solutions in relation to depth. Soil solution ¹⁸O/¹⁶O decreased in relation to depth. Comparing with xylem sap ¹⁸O/¹⁶O data indicated a deeper uptake of soil water by pine than spruce on the podzol site and a superficial uptake by both species on the peat. The ⁸⁷Sr/⁸⁶Sr ratio of bioavailable Sr generally increased in soils in relation to depth. Contrastingly, the ⁸⁷Sr/⁸⁶Sr ratio in spruce wood was generally higher than in pine wood suggesting a deeper uptake of Sr by spruce. But the ⁸⁷Sr/⁸⁶Sr ratio and concentrations of bioavailable Sr were systematically higher below spruce than below pine. In order to explain these unexpected results, we built a simple flux model to investigate the possible effects of interspecific variations in Sr cycling, soil mineral weathering and depth of Sr uptake on soil and tree ⁸⁷Sr/⁸⁶Sr ratio. At the study sites, spruce cycled in litterfall up to 12 times more strontium than pine. The use of the model showed that this difference in Sr cycling could alone explain higher isotopic signatures of trees and topsoils below spruce. Besides, high isotopic signatures of roots in the A/E horizons below spruce led us to hypothesise a species-specific weathering process. Finally, the comparison between the ⁸⁷Sr/⁸⁶Sr ratios in wood and root or soil solutions below each species suggested that the average depth of Sr and water uptake were close, but irregular variations of the Sr isotopic ratio with depth reduce the accuracy of the results. Tree species strongly influence Sr isotopic ratios in boreal forest soils through differences in Sr cycling, and possibly through specific mineral weathering.     

Ectomycorrhizal-Dominated Boreal and Tropical Forests Have Distinct Fungal Communities,but Analogous Spatial Patterns across Soil Horizons

Fungi regulate key nutrient cycling processes in many forest ecosystems, but their diversity and distribution within and across ecosystems are poorly understood. Here, we examine the spatial distribution of fungi across a boreal and tropical ecosystem, focusing on ectomycorrhizal fungi. We analyzed fungal community composition across litter (organic horizons) and underlying soil horizons (0–20 cm) using 454 pyrosequencing and clone library sequencing. In both forests, we found significant clustering of fungal communities by site and soil horizons with analogous patterns detected by both sequencing technologies. Free-living saprotrophic fungi dominated the recently-shed leaf litter and ectomycorrhizal fungi dominated the underlying soil horizons. This vertical pattern of fungal segregation has also been found in temperate and European boreal forests, suggesting that these results apply broadly to ectomycorrhizal-dominated systems, including tropical rain forests. Since ectomycorrhizal and free-living saprotrophic fungi have different influences on soil carbon and nitrogen dynamics, information on the spatial distribution of these functional groups will improve our understanding of forest nutrient cycling.     

Ectomycorrhizal colonization slows root decomposition: the post-mortem fungal legacy

The amount of carbon plants allocate to mycorrhizal symbionts exceeds that emitted by human activity annually. Senescent ectomycorrhizal roots represent a large input of carbon into soils, but their fate remains unknown. Here, we present the surprising result that, despite much higher nitrogen concentrations, roots colonized by ectomycorrhizal (EM) fungi lost only one-third as much carbon as non-mycorrhizal roots after 2 years of decomposition in a piñon pine ( Pinus edulis ) woodland. Experimentally excluding live mycorrhizal hyphae from litter, we found that live mycorrhizal hyphae may alter nitrogen dynamics, but the afterlife (litter-mediated) effects of EM fungi outweigh the influences of live fungi on root decomposition. Our findings indicate that a shift in plant allocation to mycorrhizal fungi could promote carbon accumulation in soil by this pathway. Furthermore, EM litters could directly contribute to the process of stable soil organic matter formation, a mechanism that has eluded soil scientists.     

Effects of far-red light on the growth,mycorrhizas and mineral nutrition of Scots pine seedlings

The effects of supplementary far-red sidelight on the formation of mycorrhizas and on the accumulation and allocation of dry weight and mineral nutrients were studied in Scots pine (Pinus sylvestris L.) seedlings. Starting one week after germination the seedlings were subjected to two different light quality regimes: control and simulated sparse-canopy conditions (FR+). In the FR+ regime, light reflected by neighbouring plants was simulated by means of supplementary far-red light sources, which reduced the horizontal red/far-red photon ratio (R:FR) without affecting PAR. Seedlings were harvested after three months of treatment. FR+ increased stem height and decreased the total dry weight of seedlings. Dry weight allocation to needles was not affected, whereas dry weight allocation to roots was reduced and that to stems was increased in FR+ treated seedlings. The total number of short root tips and developing mycorrhizas per seedling were lower in FR+ than in control plants. Most short roots were developing mycorrhizas, while non-mycorrhizal short roots and mycorrhizas with mantle or external mycelium were very scarce. Changes in the allocation of nutrients in general followed the changes in dry weight allocation, and changes in nutrient content followed those in total dry weight. However, mismatches among these changes resulted in significant changes in nutrient concentrations in some organs: the concentrations of nitrogen and potassium in needles and the concentration of nitrogen in stems were higher in FR+ than in control seedlings. Changes in biomass and nutrient allocation under low R:FR may promote rapid height growth during early development in stands of Scots pine seedlings, but concomitant reductions in growth of the root system and mycorrhizas may negatively affect tree performance over the long term.     

Assessing fine-root biomass and production in a Scots pine stand – comparison of soil core and root ingrowth core methods

Soil core and root ingrowth core methods for assessing fine-root (< 2 mm) biomass and production were compared in a 38-year-old Scots pine (Pinus sylvestris L) stand in eastern Finland. 140 soil cores and 114 ingrowth cores were taken from two mineral soil layers (0–10 cm and 10–30 cm) during 1985–1988. Seasonal changes in root biomass (including both Scots pine and understorey roots) and necromass were used for calculating fine-root production. The Scots pine fine-root biomass averaged annually 143 g/m² and 217 g/m² in the upper mineral soil layer, and 118 g/m² and 66 g/m² in the lower layer of soil cores and ingrowth cores, respectively. The fine-root necromass averaged annually 601 g/m² and 311 g/m² in the upper mineral soil layer, and 196 g/m² and 159 g/m² in the lower layer of soil cores and ingrowth cores, respectively. The annual fine-root production in a Scots pine stand in the 30 cm thick mineral soil layer, varied between 370–1630 g/m² in soil cores and between 210 – 490 g/m² in ingrowth cores during three years. The annual production calculated for Scots pine fine roots, varied between 330–950 g/m² in soil cores and between 110 – 610 g/m² in ingrowth cores. The horizontal and vertical variation in fine-root biomass was smaller in soil cores than in ingrowth cores. Roots in soil cores were in the natural dynamic state, while the roots in the ingrowth cores were still expanding both horizontally and vertically. The annual production of fine-root biomass in the Scots pine stand was less in root ingrowth cores than in soil cores. During the third year, the fine-root biomass production of Scots pine, when calculated by the ingrowth core method, was similar to that calculated by the soil core method. Both techniques have sources of error. In this research the sampling interval in the soil core method was 6–8 weeks, and thus root growth and death between sampling dates could not be accurately estimated. In the ingrowth core method, fine roots were still growing into the mesh bags. In Finnish conditions, after more than three growing seasons, roots in the ingrowth cores can be compared with those in the surrounding soil. The soil core method can be used for studying both the annual and seasonal biomass variations. For estimation of production, sampling should be done at short intervals. The ingrowth core method is more suitable for estimating the potential of annual fine-root production between different site types.     

Laccaria bicolor S238N improves Scots pine mineral nutrition by increasing root nutrient uptake from soil minerals but does not increase mineral weathering

The role of ectomycorrhizal fungi on mineral nutrient mobilization and uptake is crucial for tree nutrition and growth in temperate forest ecosystems. By using a “mineral weathering budget” approach, this study aims to quantify the effect of the symbiosis with the ectomycorrhizal model strain Laccaria bicolor S238N on mineral weathering and tree nutrition, carrying out a column experiment with a quartz/biotite substrate. Each column was planted with one Scots pine (Pinus sylvestris L.) non-mycorrhizal or mycorrhizal with L. bicolor, with exception of the abiotic control treatment. The columns were continuously supplied with a nutrient-poor solution. A mineral weathering budget was calculated for K and Mg. The pine shoot growth was significantly increased (73%) when plants were mycorrhizal with L. bicolor. Whatever their mycorrhizal status, pines increased mineral weathering by factors 1.5 to 2.1. No difference between non-mycorrhizal and mycorrhizal pine treatments was revealed, however, mycorrhizal pines assimilated significantly more K and Mg. This suggests that in our experimental conditions, L. bicolor S238N improved shoot growth and K and Mg assimilation in Scots pine mainly by increasing the uptake of dissolved nutrients, linked to a better exploration and exploitation of the soil by the mycorrhizal roots.     

Frost hardiness of mycorrhizal (Hebeloma sp.) and non-mycorrhizal Scots pine roots

The frost hardiness (FH) of mycorrhizal [ectomycorrhizal (ECM)] and non-mycorrhizal (NM) Scots pine (Pinus sylvestris) seedlings was studied to assess whether mycorrhizal symbiosis affected the roots’ tolerance of below-zero temperatures. ECM (Hebeloma sp.) and NM seedlings were cultivated in a growth chamber for 18 weeks. After 13 weeks’ growth in long-day and high-temperature (LDHT) conditions, a half of the ECM and NM seedlings were moved into a chamber with short-day and low-temperature (SDLT) conditions to cold acclimate. After exposures to a range of below-zero temperatures, the FH of the roots was assessed by means of the relative electrolyte leakage test. The FH was determined as the inflection point of the temperature-response curve. No significant difference was found between the FH of mycorrhizal and non-mycorrhizal roots in LDHT (?8.9 and ?9.8 °C) or SDLT (?7.5 and ?6.8 °C). The mycorrhizal treatment had no significant effect on the total dry mass, the allocation of dry mass among the roots and needles or nutrient accumulation. The mycorrhizal treatment with Hebeloma sp. did not affect the FH of Scots pine in this experimental setup. More information is needed on the extent to which mycorrhizas tolerate low temperatures, especially with different nutrient contents and different mycorrhiza fungi.     

Fine root distribution of trees and understory in mature stands of maritime pine (Pinus pinaster) on dry and humid sites

Maritime pine (Pinus pinaster) is the main tree cropping species in the Landes of Gascogne forest range in south western France. Soils are nutrient poor, sandy podzosols and site fertility is determined essentially by organic matter content and depth of water table, which is known to limit root growth. We hypothesised, with an ultimate goal of constructing a nutrient uptake model applicable to this region, that the organic top horizons together with the depth of the water table should be the most important parameters related to fine root distribution and presence of associated mycorrhiza. To test this hypothesis, we compared two adult Pinus pinaster stands, contrasting in depth of water table and soil fertility and evaluated fine roots (diameter ≤2 mm) of understory species and fine roots and ectomycorrhizal morphotypes of Pinus pinaster down to 1.2 m, using a soil corer approach. Total fine root biomass of Pinus pinaster was not significantly different between both sites (3.6 and 4.5 t ha⁻¹ for the humid, respectively, dry site), but root distribution was significantly shallower and root diameter increased more with depth at the humid site, presumably due to more adverse soil conditions as related to the presence of a hardpan, higher amount of aluminium oxides and / or anoxia. Fine roots of Pinus pinaster represented only about 30% of total fine root biomass and 15% of total fine root length, suggesting that the understory species cannot be ignored with regards to competition for mineral nutrients and water. A comparison of the ectomycorrhizal morphotypes showed that the humid site could be characterised by a very large proportion of contact exploration types, thought to be more relevant in accessing organic nutrient sources, whereas the dry site had a significantly higher proportion of both long-distance and short-distance exploration types, the latter of which was thought to be more resistant to short-term drought periods. These results partly confirm our hypothesis on root distribution as related to the presence of soil mineral nutrients (i.e. in organic matter), point out the potential role of understory plant species and ectomycorrhizal symbiosis and are a valuable step in building a site-specific nutrient uptake model.     

Tree root and soil heterotrophic respiration as revealed by girdling of boreal Scots pine forest: extending observations beyond the first year

Limitations in available techniques to separate autotrophic (root) and soil heterotrophic respiration have hampered the understanding of forest C cycling. The former is here defined as respiration by roots, their associated mycorrhizal fungi and other micro‐organisms in the rhizosphere directly dependent on labile C compounds leaked from roots. In order to separate the autotrophic and heterotrophic components of soil respiration, all Scots pine trees in 900 m² plots were girdled to instantaneously terminate the supply of current photosynthates from the tree canopy to roots. Högberg et al. (Nature 411, 789–792, 2001) reported that autotrophic activity contributed up to 56% of total soil respiration during the first summer of this experiment. They also found that mobilization of stored starch (and likely also sugars) in roots after girdling caused an increased apparent heterotrophic respiration on girdled plots. Herein a transient increase in the δ¹³C of soil CO₂ efflux after girdling, thought to be due to decomposition of ¹³C‐enriched ectomycorrhizal mycelium and root starch and sugar reserves, is reported. In the second year after girdling, when starch reserves of girdled tree roots were exhausted, calculated root respiration increased up to 65% of total soil CO₂ efflux. It is suggested that this estimate of its contribution to soil respiration is more precise than the previous based on one year of observation. Heterotrophic respiration declined in response to a 20‐day‐long 6 °C decline in soil temperature during the second summer, whereas root respiration did not decline. This did not support the idea that root respiration should be more sensitive to variations in soil temperature. It is suggested that above‐ground photosynthetic activity and allocation patterns of recent photosynthates to roots should be considered in models of responses of forest C balances to global climate change.     

Respiration activity of ectomycorrhizas from Cenococcum geophilum and Lactarius sp. in relation to soil water potential in five beech forests

Forest trees are involved in root symbioses with hundreds of species of ectomycorrhizal fungi which constitute functional guilds able to improve the water and mineral nutrition of host trees. In temperate ecosystems, water shortage is a main factor limiting tree vitality. To assess how soil water conditions affected the physiological state of beech (Fagus silvatica L.) ectomycorrhizal roots, we monitored glucose respiration of two ectomycorrhizal types (Lactarius sp. and Cenococcum geophilum) during two complete growing seasons. Five stands of contrasting soil conditions were chosen in north-eastern France. The top soil horizons were equipped with micropsychrometers for measuring water potential and temperature. Glucose respiration on individual ectomycorrhizas was measured in vitro by trapping [¹⁴C]-CO₂ from radiolabelled glucose. For soil water potential <-0.2 MPa, the potential respiration activity of C. geophilumectomycorrhizas was significantly less altered than that of Lactariussp. ectomycorrhizas, indicating that C. geophilumis more likely than Lactariussp. to maintain the physiological integrity of beech roots facing drought stress.     

The survival and development of inoculant ectomycorrhizal fungi on roots of outplanted Eucalyptus globulus Labill

The survival and development of two inoculant ectomycorrhizal fungi (Hebeloma westraliense Bough. Tom. and Mal. and Setchelliogaster sp. nov.) on roots of outplanted Eucalyptus globulus Labill. was examined at two expasture field sites in the south-west of Western Australia. Site 1 was a gravelly yellow duplex soil, and Site 2 was a yellow sandy earth. Plants were grown in steamed or unsteamed soil, in root bags designed as field containers for young growing trees. Three, 6 and 12 months after outplanting, plants were removed from these bags and assessed for dry weights of shoots and ectomycorrhizal colonization of roots.The inoculant ectomycorrhizal fungi (identified on the basis of the colour and morphology of their mycorrhizas) survived on roots of E. globulus for at least 12 months after outplanting at both field sites. At Site 1, however, colonization of new fine roots by the inoculant fungi was low (less than 20% of fine root length). Inoculation had no effect on the growth of E. globulus at this site. In contrast, at Site 2 the inoculant ectomycorrhizal fungi colonized up to 30–50% of new fine root length during the first 6 months after outplanting. There was a corresponding growth response to ectomycorrhizal inoculation at this site, with a close relationship (r²=0.82^**) between plant growth at 12 months and root colonization at 3 months. Plant growth at 12 months was related less closely with root colonization at 6 or 12 months. Root colonization by resident ectomycorrhizal fungi increased with time at both field sites. At Site 2, this increase appeared to be at the expense of colonization by the inoculant fungi, which was reduced to less than 10% of fine root length at 12 months. Steaming the soil had little effect on colonization by the inoculant ectomycorrhizal fungi at either field site, but decreased colonization by the resident ectomycorrhizal fungi.     

Site preparation alters soil distribution of roots and ectomycorrhizae on outplanted western white pine and Douglas-fir

This report documents root and ectomycorrhizal development on container-produced (1-0), outplanted, western white pine and Douglas-fir seedlings growing in site-prepared forest soils typical of the Inland Northwestern US. The following site preparations were used: 1) mounding organic and surface mineral horizons; 2) mounding with subsequent physical removal or chemical control of competing vegetation; 3) scalping to reduce competing vegetation; and, 4) a control or no post-harvest disturbance. Treatments were applied on relatively harsh and moderate sites in northern Idaho. Most ectomycorrhizae on the seedling population were found in the mineral substrates that dominated planting sites. However, compared to mineral substrates, highest seedling ectomycorrhizal tip counts were recorded in organic matter, particularly decayed wood or mixtures containing decayed wood. Strong ectomycorrhizal development was characteristic of western white pine. It supported highest ectomycorrhizal activity in organic substrates on the harshest treatments (scalps). Douglas-fir showed even stronger relative increases of ectomycorrhizae in organic substrates on harsh treatments. Three of the four common ectomycorrhizal morphological types were concentrated in mineral substrates with all treatments. A treatment-induced change of behavior was shown by the principal pine type. It occurred at highest numbers in organic substrates of the mound with competing vegetation treatment and in mineral substrates with the control. If relative availability to seedling roots was considered, organics (especially decomposed wood) were generally equal or superior to mineral substrates for supporting ectomycorrhizal activity on planted seedlings.     

Survival of black alder (Alnus glutinosa L.), silver birch (Betula pendula Roth.) and Scots pine (Pinus sylvestris L.) seedlings in a reclaimed oil shale mining area

Early survival and growth of black alder, silver birch and Scots pine were investigated on reclaimed extremely stony and heterogeneous calcareous (pH 8) opencast oil shale mining areas (OOSMAs). Biomass allocation, production, leaf and root adaptations, and mineral nutrition in relation to tree species and soil heterogeneity were analysed. The adaptive strategies of tree species in first-year plantations on OOSMA were different. Scots pine allocated 1.5–2 times more biomass into leaves and fine roots than deciduous trees. The lower leaf/fine root biomass ratio was in proportion to the better survival (%) of seedlings, decreasing in the following order: black alder (93%) ≥ Scots pine (83%) > silver birch (64%). Deciduous trees improved mineral nutrition more by fine-root morphological adaptations than Scots pine; e.g. the mean specific root length (SRL, m g^?1) of short roots increased in the following order: Scots pine (62) < black alder (172) < silver birch (314). The effect of soil heterogeneity on growth and adaptations was minor. All studied species suffered from P and N, and deciduous species also from K deficiency. In the first year after planting, black alder was best adapted to the harsh conditions of the post-mining substrate. The approaches of this study can be used for other regions where wastelands require reclamation.     

Variation of carbon age of fine roots in boreal forests determined from 14C measurements

Background and aims

The main objectives of this study were to determine how the carbon age of fine root cellulose varies between stands, tree species, root diameter and soil depth. In addition, we also compared the carbon age of fine roots from soil cores of this study with reported values from the roots of the same diameter classes of ingrowth cores on the same sites.

Methods

We used natural abundance of ¹⁴C to estimate root carbon age in four boreal Norway spruce and Scots pine stands in Finland and Estonia.

Results

Age of fine root carbon was older in 1.5–2 mm diameter fine roots than in fine roots with <0.5 mm diameter, and tended to be older in mineral soil than in organic soil. Fine root carbon was older in the less fertile Finnish spruce stands (11–12 years) than in the more fertile Estonian stand (3 and 8 years), implying that roots may live longer in less fertile soil. We further observed that on one of our sites carbon in live fine roots with the 1.5–2 mm diameter was of similar C age (7–12 years) than in the ingrowth core roots despite the reported root age in the ingrowth cores – being not older than 2 years.

Conclusions

From this result, we conclude that new live roots may in some cases use old carbon reserves for their cellulose formation. Future research should be oriented towards improving our understanding of possible internal redistribution and uptake of C in trees.