Soil organic carbon in irrigated agricultural systems: A meta-analysis.

Over the last 200 years, conversion of non-cultivated land for agriculture has substantially reduced global soil organic carbon (SOC) stocks in upper soil layers. Nevertheless, practices such as no- or reduced tillage, application of organic soil amendments, and maintenance of continuous cover can increase SOC in agricultural fields. While these management practices have been well studied, the effects on SOC of cropping systems that incorporate irrigation are poorly understood. Given the large, and expanding, agricultural landbase under irrigation across the globe, this is a critical knowledge gap for climate change mitigation. We undertook a systematic literature review and subsequent meta-analysis of data from studies that examined changes in SOC on irrigated agricultural sites through time. We investigated changes in SOC by climate (aridity), soil texture, and irrigation method with the following objectives: (i) to examine the impact of irrigated agriculture on SOC storage; and (ii) to identify the conditions under which irrigated agriculture is most likely to enhance SOC. Overall, irrigated agriculture increased SOC stocks by 5.9%, with little effect of study length (2-47 years). However, changes in SOC varied by climate and soil depth, with the greatest increase in SOC observed on irrigated semi-arid sites at the 0-10 cm depth (14.8%). Additionally, SOC increased in irrigated fine- and medium-textured soils but not coarse-textured soils. Furthermore, while there was no overall change to SOC in flood/furrow irrigated sites, SOC tended to increase in sprinkler irrigated sites, and decrease in drip irrigated sites, especially at depths below 10 cm. This work sheds light on the nuances of SOC change across irrigated agricultural systems, highlights the importance of studying SOC storage in deeper soils, and will help guide future research on the impacts of irrigated agriculture on SOC.

Dual-mycorrhizal plants: their ecology and relevance.

Dual-mycorrhizal plants are capable of associating with fungi that form characteristic arbuscular mycorrhizal (AM) and ectomycorrhizal (EM) structures. Here, we address the following questions: (1) How many dual-mycorrhizal plant species are there? (2) What are the advantages for a plant to host two, rather than one, mycorrhizal types? (3) Which factors can provoke shifts in mycorrhizal dominance (i.e. mycorrhizal switching)? We identify a large number (89 genera within 32 families) of confirmed dual-mycorrhizal plants based on observing arbuscules or coils for AM status and Hartig net or similar structures for EM status within the same plant species. We then review the possible nutritional benefits and discuss the possible mechanisms leading to net costs and benefits. Cost and benefits of dual-mycorrhizal status appear to be context dependent, particularly with respect to the life stage of the host plant. Mycorrhizal switching occurs under a wide range of abiotic and biotic factors, including soil moisture and nutrient status. The relevance of dual-mycorrhizal plants in the ecological restoration of adverse sites where plants are not carbon limited is discussed. We conclude that dual-mycorrhizal plants are underutilized in ecophysiological-based experiments, yet are powerful model plant-fungal systems to better understand mycorrhizal symbioses without confounding host effects.

Rapid nitrogen loss from ectomycorrhizal pine germinants signaled by their fungal symbiont.

Ectomycorrhizal fungi supply their plant partners with nitrogen but can also retain substantial amounts. The concentration of nitrogen in the soil and the amount of carbon supplied from the host seem to influence the proportion of N retained by the fungus. In an experiment designed to determine whether differential supply of nitrogen to two plants influenced nitrogen transfer from fungus to plant within a mycorrhizal network, we observed rapid, substantial loss of nitrogen from pine seedlings. The loss occurred when the mycorrhizal fungus experienced a sudden increase in nitrogen supply. We grew Pinus contorta seedlings in association with Suillus tomentosus in low-nitrogen microcosms where some nitrogen was accessible only by hyphae. After 70 days, foliage of some seedlings was treated with nitrogen. Three days later, hyphal nutrient media were replaced with water or a solution containing nitrogen. Foliar treatment did not affect nitrogen transfer by the fungus to shoots, but by day 75, seedling nitrogen contents had dropped by 60% in microcosms where nitrogen had been added to the hyphal compartments. Those seedlings retained only 55% of the nitrogen originally present in the seed. Loss of nitrogen did not occur if water was added or the hyphae were severed. Because of the severing effect, we concluded that S. tomentosus triggered the loss of seedling nitrogen. Nitrogen may have been lost through increased root exudation or transfer to the fungus. Access to nitrogen from nutrient-rich germinants would benefit rhizosphere microorganisms, including ectomycorrhizal fungi colonizing pine from spores after wildfire.

Early-successional ectomycorrhizal fungi effectively support extracellular enzyme activities and seedling nitrogen accumulation in mature forests.

After stand-replacing disturbance, regenerating conifer seedlings become colonized by different ectomycorrhizal fungi (EMF) than the locally adapted EMF communities present on seedlings in mature forests. We studied whether EMF species that colonized subalpine fir (Abies lasiocarpa) seedlings in clearcuts differed from those that colonized seedlings in adjacent mature forests with respect to mycorrhizoplane extracellular enzyme activities (EEAs) and N status of the seedlings. We tested two alternate hypotheses: (1) that EEAs would differ between the two EMF communities, with higher activities associated with forest-origin communities, and (2) that acclimation to soil environment was considerable enough that EEAs would be determined primarily by the soil type in which the ectomycorrhizas were growing. Naturally colonized fir seedlings were reciprocally transplanted between clearcuts and forests, carrying different EMF communities with them. EEAs were influenced more by destination environment than by EMF community. EEAs were as high in early-successional as in late-successional communities in both destination environments. Buds of clearcut-origin seedlings had the same or higher N contents as forest seedlings after a growing season in either environment. These results indicate that (i) symbiotic EMF and/or their associated microbial communities demonstrate substantial ability to acclimate to new field environments; (ii) the ability to produce organic matter-degrading enzymes is not a trait that necessarily distinguishes early- and late-successional EMF communities in symbiosis; (iii) early-successional EMF are as capable of supporting seedling N accumulation in forest soils as late-successional EMF; and (iv) disturbed ecosystems where early-successional EMF are present should have high resilience for organic matter degradation.

Ectomycorrhizal communities of ponderosa pine and lodgepole pine in the south-central Oregon pumice zone.

Forest ecosystems of the Pacific Northwest of the USA are changing as a result of climate change. Specifically, rise of global temperatures, decline of winter precipitation, earlier loss of snowpack, and increased summer drought are altering the range of Pinus contorta. Simultaneously, flux in environmental conditions within the historic P. contorta range may facilitate the encroachment of P. ponderosa into P. contorta territory. Furthermore, successful pine species migration may be constrained by the distribution or co-migration of ectomycorrhizal fungi (EMF). Knowledge of the linkages among soil fungal diversity, community structure, and environmental factors is critical to understanding the organization and stability of pine ecosystems. The objectives of this study were to establish a foundational knowledge of the EMF communities of P. ponderosa and P. contorta in the Deschutes National Forest, OR, USA, and to examine soil characteristics associated with community composition. We examined EMF root tips of P. ponderosa and P. contorta in soil cores and conducted soil chemistry analysis for P. ponderosa cores. Results indicate that Cenococcum geophilum, Rhizopogon salebrosus, and Inocybe flocculosa were dominant in both P. contorta and P. ponderosa soil cores. Rhizopogon spp. were ubiquitous in P. ponderosa cores. There was no significant difference in the species composition of EMF communities of P. ponderosa and P. contorta. Ordination analysis of P. ponderosa soils suggested that soil pH, plant-available phosphorus (Bray), total phosphorus (P), carbon (C), mineralizable nitrogen (N), ammonium (NH4), and nitrate (NO3) are driving EMF community composition in P. ponderosa stands. We found a significant linear relationship between EMF species richness and mineralizable N. In conclusion, P. ponderosa and P. contorta, within the Deschutes National Forest, share the same dominant EMF species, which implies that P. ponderosa may be able to successfully establish within the historic P. contorta range and dominant EMF assemblages may be conserved.

Little evidence for niche partitioning among ectomycorrhizal fungi on spruce seedlings planted in decayed wood versus mineral soil microsites.

Ectomycorrhizal fungal (EMF) communities vary among microhabitats, supporting a dominant role for deterministic processes in EMF community assemblage. EMF communities also differ between forest and clearcut environments, responding to this disturbance in a directional manner over time by returning to the species composition of the original forest. Accordingly, we examined EMF community composition on roots of spruce seedlings planted in three different microhabitats in forest and clearcut plots: decayed wood, mineral soil adjacent to downed wood, or control mineral soil, to determine the effect of retained downed wood on EMF communities over the medium and long term. If downed and decayed wood provide refuge habitat distinct from that of mineral soil, we would expect EMF communities on seedlings in woody habitats in clearcuts to be similar to those on seedlings planted in the adjacent forest. As expected, we found EMF species richness to be higher in forests than clearcuts (P ≤ 0.01), even though soil nutrient status did not differ greatly between the two plot types (P ≥ 0.05). Communities on forest seedlings were dominated by Tylospora spp., whereas those in clearcuts were dominated by Amphinema byssoides and Thelephora terrestris. Surprisingly, while substrate conditions varied among microsites (P ≤ 0.03), especially between decayed wood and mineral soil, EMF communities were not distinctly different among microhabitats. Our data suggest that niche partitioning by substrate does not occur among EMF species on very young seedlings in high elevation spruce-fir forests. Further, dispersal limitations shape EMF community assembly in clearcuts in these forests.

Ectomycorrhizal fungal community assembly on regenerating Douglas-fir after wildfire and clearcut harvesting.

Wildfire severity in forests is projected to increase with warming and drying conditions associated with climate change. Our objective was to determine the impact of wildfire and clearcutting severity on the ectomycorrhizal fungal (EMF) community of Douglas-fir seedlings in the dry forests of interior British Columbia, Canada. We located our study within and surrounding the area of the McLure fire (August 2003). We hypothesized that disturbance would affect EMF community assembly due to reductions in fungal inoculum. Five treatments representing a range of disturbance severities were compared: high severity burn, low severity burn, screefed clearcut (manual removal of forest floor), clearcut, and undisturbed forest. EMF communities in the undisturbed forest were more complex than those in all disturbance treatments. However, aspects of community assembly varied with disturbance type, where the burn treatments had the simplest communities. After 4 months, regenerating seedlings in the burn treatments had the lowest colonization, but seedlings in all treatments were fully colonized within 1 year. EMF communities were similar among the four disturbance types, largely due to dominance of Wilcoxina throughout the study period. However, forest floor retention influenced community assembly as the EMF in the clearcut treatment, where forest floor was retained, had levels of diversity and richness comparable to the undisturbed forest. Overall, the results suggest that increasing forest floor disturbance can alter EMF community assembly in the first year of regeneration. A correlation between poorly colonized seedlings and seedling productivity also suggests a role for productivity in influencing community assembly.

Positive citation bias and overinterpreted results lead to misinformation on common mycorrhizal networks in forests.

A common mycorrhizal network (CMN) is formed when mycorrhizal fungal hyphae connect the roots of multiple plants of the same or different species belowground. Recently, CMNs have captured the interest of broad audiences, especially with respect to forest function and management. We are concerned, however, that recent claims in the popular media about CMNs in forests are disconnected from evidence, and that bias towards citing positive effects of CMNs has developed in the scientific literature. We first evaluated the evidence supporting three common claims. The claims that CMNs are widespread in forests and that resources are transferred through CMNs to increase seedling performance are insufficiently supported because results from field studies vary too widely, have alternative explanations or are too limited to support generalizations. The claim that mature trees preferentially send resources and defence signals to offspring through CMNs has no peer-reviewed, published evidence. We next examined how the results from CMN research are cited and found that unsupported claims have doubled in the past 25 years; a bias towards citing positive effects may obscure our understanding of the structure and function of CMNs in forests. We conclude that knowledge on CMNs is presently too sparse and unsettled to inform forest management.

Phosphorus deficiencies invoke optimal allocation of exoenzymes by ectomycorrhizas.

Ectomycorrhizal (EM) fungi can acquire phosphorus (P) through the production of extracellular hydrolytic enzymes (exoenzymes), but it is unclear as to the manner and extent native EM fungal communities respond to declining soil P availability. We examined the activity of six exoenzymes (xylosidase, N-acetyl glucosaminidase, ß-glucosidase, acid phosphomonoesterase, acid phosphodiesterase [APD], laccase) from EM roots of Pseudotsuga menzesii across a soil podzolization gradient of coastal British Columbia. We found that APD activity increased fourfold in a curvilinear association with declining inorganic P. Exoenzyme activity was not related to organic P content, but at a finer resolution using 31P-NMR, there was a strong positive relationship between APD activity and the ratio of phosphodiesters to orthophosphate of surface organic horizons (forest floors). Substantial increases (two- to fivefold) in most exoenzymes were aligned with declining foliar P concentrations of P. menzesii, but responses were statistically better in relation to foliar nitrogen (N):P ratios. EM fungal species with consistently high production of key exoenzymes were exclusive to Podzol plots. Phosphorus deficiencies in relation to N limitations may provide the best predictor of exoenzyme investment, reflecting an optimal allocation strategy for EM fungi. Resource constraints contribute to species turnover and the assembly of distinct, well-adapted EM fungal communities.

Access to mycorrhizal networks and roots of trees: importance for seedling survival and resource transfer.

Mycorrhizal networks (MNs) are fungal hyphae that connect roots of at least two plants. It has been suggested that these networks are ecologically relevant because they may facilitate interplant resource transfer and improve regeneration dynamics. This study investigated the effects of MNs on seedling survival, growth and physiological responses, interplant resource (carbon and nitrogen) transfer, and ectomycorrhizal (EM) fungal colonization of seedlings by trees in dry interior Douglas-fir (Pseudotsuga menziesii var. glauca) forests. On a large, recently harvested site that retained some older trees, we established 160 isolated plots containing pairs of older Douglas-fir "donor" trees and either manually sown seed or planted Douglas-fir "receiver" seedlings. Seed- and greenhouse-grown seedlings were sown and planted into four mesh treatments that served to restrict MN access (i.e., planted into mesh bags with 0.5-, 35-, 250-microm pores, or without mesh). Older trees were pulse labeled with carbon (13CO2) and nitrogen (15NH4(15)NO3) to quantify resource transfer. After two years, seedlings grown from seed in the field had the greatest survival and received the greatest amounts of transferred carbon (0.0063% of donor photo-assimilates) and nitrogen (0.0018%) where they were grown without mesh; however, planted seedlings were not affected by access to tree roots and hyphae. Size of "donor" trees was inversely related to the amount of carbon transferred to seedlings. The potential for MNs to form was high (based on high similarity of EM communities between hosts), and MN-mediated colonization appeared only to be important for seedlings grown from seed in the field. These results demonstrate that MNs and mycorrhizal roots of trees may be ecologically important for natural regeneration in dry forests, but it is still uncertain whether resource transfer is an important mechanism underlying seedling establishment.

Effects of NaCl on responses of ectomycorrhizal black spruce (Picea mariana), white spruce (Picea glauca) and jack pine (Pinus banksiana) to fluoride.

Black spruce (Picea mariana), white spruce (Picea glauca) and jack pine (Pinus banksiana) were inoculated with Suillus tomentosus and subjected to potassium fluoride (1 mM KF and 5 mM KF) in the presence and absence of 60 mM NaCl. The NaCl and KF treatments reduced total dry weights in jack pine and black spruce seedlings, but they did not affect total dry weights in white spruce seedlings. The addition of 60 mM NaCl to KF treatment solutions alleviated fluoride-induced needle injury in ectomycorrhizal (ECM) black spruce and white spruce, but had little effect in jack pine seedlings. Both KF and 60 mM NaCl treatments reduced E values compared with non-treated control seedlings. However, with the exception of small reductions of K(r) by NaCl treatments in black spruce, the applied KF and NaCl treatments had little effect on K(r) in ECM plants. Chloride tissue concentrations in NaCl-treated plants were not affected by the presence of KF in treatment solutions. However, shoot F concentrations in ECM black spruce and white spruce treated with 5 mM KF + 60 mM NaCl were significantly reduced compared with the 5 mM KF treatment. The results point to a possible competitive inhibition of F transport by Cl. We also suggest that the possibility that aquaporins may be involved in the transmembrane transport of F should be further investigated.

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.

The mutualism-parasitism continuum in ectomycorrhizas: a quantitative assessment using meta-analysis.

Context dependency is deemed to position the outcomes of species interactions along a continuum of mutualism to parasitism. Thus, it is imperative to understand which factors determine where a particular interspecific interaction falls along the continuum. Over the past 20 years research on the ectomycorrhizal symbiosis has resulted in sufficient independent studies to now generalize about the factors and mechanisms that affect host response to ectomycorrhizas. Using meta-analysis we quantitatively evaluated the role of biotic (partner identity and colonization levels of ectomycorrhizal fungi) and abiotic (phosphorus levels) factors in determining host biomass, height, and shoot:root responses to ectomycorrhizal associations. On average, seedlings across multiple host genera increased in total biomass when inoculated with ectomycorrhizal fungi regardless of the identity of the fungal associate; host genera differed in the magnitude of response for both total biomass and shoot:root ratio. Association with different fungal genera modified only host allocation of biomass to shoots and roots. Neither level of colonization on inoculated seedlings nor the level of contamination on control seedlings relative to colonization levels by target fungi on inoculated seedlings was important in explaining variation in effect sizes for any growth response. None of our proposed factors (identity of partners, colonization level, magnitude of contamination, or duration of association) explained variation in effect sizes for shoot height, although in general seedlings were taller when inoculated with ectomycorrhizal fungi. Phosphorus additions did not influence effect sizes. Although the general trend across studies was for a positive response of hosts to ectomycorrhizal inoculation, publication bias and methodological issues effectively reduce and distort the spectrum on which we evaluate host responses to ectomycorrhizal inoculation. Our results indicate that the variation in ectomycorrhizal fungi perceived by the host may be of a discrete (presence/absence of ectomycorrhizal fungi) rather than continuous nature (variation in identity or abundance of ectomycorrhizal fungi).

Fluxes of carbon and phosphorus between symbionts in willow ectomycorrhizas and their changes with time.

One way of viewing a mycorrhizal symbiosis is as a balance between the nutritional 'benefits' and carbon 'costs' to the phytobiont. Phosphorus acquisition efficiency (the amount of phosphorus taken up per unit of carbon allocated belowground) can be used as an indicator of this balance. In this study, phosphorus uptake and belowground carbon allocation were measured using ectomycorrhizal (M) (Thelephora terrestris (Ehrh.) Fr.) and non-mycorrhizal (NM) Salix viminalis L. cv. Bowles Hybrid. Following 50, 60, 85 or 98 d of growth in a gamma-irradiated soil/sand mixture containing 4 mg bicarbunate-extractable P kg-1 , seven randomly-selected cuttings of each treatment were harvested and their P contents determined. Nine d prior to each harvest, the three median plants from the group of seven were pulse labelled with 14 C to determine the relative allocation of C aboveground and belowground. Mycorrhizal colonization of willow caused a two-fold increase in growth owing to substantially higher P uptake. Phosphorus inflow rates were almost three times as high for M root systems as for NM root systems over the interval up to the first harvest (3.2 × 10 -12 and 1.2 × 1012 mol m-1 s-1, respectively). Over the interval from 50 to 98 d, inflows into M plants were 50% higher than into NM plants (1.4 × 1012 and 0.9 × 10-13 mol m-1 s-1 respectively). The M plants allocated about 25 times as much carbon belowground as the NM plants for both periods. The P acquisition efficiency was higher in M than in NM plants during the first interval (16% and 40% higher using two different calculation methods), whereas during the second interval it was higher in NM than in M plants (33% and 44% higher using the two different methods). Thus, ectomycorrhizas can be very effective in supplying P to their hosts even at an early stage of infection. Furthermore, it is suggested that a temporal separation exists in the maximal fluxes of P and C between the fungus and the host of the mycorrhizal association. The results are discussed in the context of the nutrient requirements and carbon economies of field-grown woody plants.

Phosphorus relationships and production of extrametrical hyphae by two types of willow ectomycorrhizas at different soil phosphorus levels.

There is much circumstantial evidence for a role of increased P uptake in the growth response of plants to ectomycorrhizas. Full response curves with and without mycorrhizal infection along a P gradient in soil are, however, required to test this hypothesis fully. In this experiment, rooted cuttings of Salix viminalis L. cv. Bowles Hybrid were grown in a 1:2 mixture by volume of gamma-irradiated soil and sterile sand, with bicarbonate-extractable P concentrations of 4, 6, 10, 21, 60 or 90 mg P kg-1 . The cuttings were inoculated by mixing peat/vermiculite spawn of Laccaria proxima (Boud.) Pat., or Thelephora terrestris (Ehrh.) Fr., or autoclaved spawn 1: 5 by volume with the soil: sand mixture. The plants showed a positive growth response to mycorrhizal infection by either fungus at the two lower P levels, and to L. proxima only at 10 mg P kg-1 . At 21 mg kg-1 and above, infection was reduced and neither mycorrhizal inoculation nor further P additions caused significant growth increases. These results imply that the growth responses to ectomycorrhizas in this experiment were solely due to increases in P uptake. Cuttings infected with L. proxima tended to be larger than those infected by T. terrestris. Estimates of percent mycorrhizal infection did not differ between the fungi at the lower P levels. However, overall production of extramatrical hyphae per gram of soil was highest in soil inoculated with L. proxima. At 10 mg P kg-1 the length of L. proxima hyphae per unit length of mycorrhizal root, P uptake per unit root weight, and total P content in plants infected with L. proxima were significantly higher than for T. terrestris. This study confirms that increased P uptake can be an important component of growth stimulation by ectomycorrhizas. It also presents the first quantification of extramatrical ectomycorrhizal hyphae in soil and suggests a role for them in the growth response.

Allocation of 14 C-carbon in ectomycorrhizal willow.

The flow of carbon from plant to fungus in ectomycorrhizal associations has not been well quantified. The objective of this study was to use 14 C to quantify the increase in fixed carbon translocated below ground in ectomycorrhizal relative to non-mycorrhizal willow (Salix viminalis L, Bowles hybrid). Rooted cuttings were inoculated with Thelephora terrestris (Ehrh). Fr. or left non-mycorrhizal. Non-mycorrhizal plants were grown at the same (4 mg kg-l bicarbonate-extractable P) (NM-P) or at a higher (21 mg kg-1 ) soil P concentration (NM +P), one at which the non-mycorrhizal plants were similar in size to the mycorrhizal (M-P) plants. At 41, 51, 76 and 89 days after planting, the shoots were exposed to a pulse of 14 CO2 . Plants were harvested after a 202 h chase period. The 14 Cactivity was quantified in live fractions: shoot tissue, shoot respiration, 'root' tissue (= roots plus fungi), 'root' respiration (= CO2 released below ground) and soil. Of the total 14 C detected in these five fractions, M-P plants allocated from 3.9% (harvest 1) to 11.5% (harvest 4) more to the below-ground fractions ('root' tissue, soil and 'root' respiration), than did the NM-P plants. Differences between NM+P and M-P plants were only half of those above (3.1 % and 4.4% at harvests 2 and 3, respectively, compared to 6.4 % and 7.4%, respectively for the difference between NM-P and M-P). Correction for differences in root/shoot ratio between M-P and NM-P plants eliminated the observed differences in carbon distribution only at the first three harvests. There was no evidence for increased 'root' respiration fates or rhizodeposition being responsible for the increased carbon diverted below ground by M-P plants.

Ectomycorrhizal fungal communities in young forest stands regenerating after clearcut logging.

The effects on the ectomycorrhizal fungal community of clearcut logging, which is used to harvest millions of hectares of ectomycorrhizal forest annually, has been studied for a number of years. Here, we review current knowledge of inoculum sources for ectomycorrhizal fungi in forests and then re-examine earlier studies of ectomycorrhizas on young trees in regenerating stands. We conclude that, taken separately from the effects of site preparation, the major impact of clearcut logging is to change the species composition of the ectomycorrhizal fungal community rather than to reduce the percentage of roots colonized. A thorough examination of site preparation treatments suggests that the changes in fungal species composition are driven by changes in the biology and chemistry of the soil environment after clearcutting as much as they are by loss or change in fungal inoculum. This is an important conclusion because it implies that these new ectomycorrhizal fungal communities are better adapted to the new conditions than the ones in the forest would have been. The shift in fungal species composition and diversity will have implications for seedling establishment and competition. The effects of individual fungi or diverse assemblages of fungi on seedling growth, and effects of changes in the ability of young trees to associate with a common mycelium are discussed. Contents Summary 399 I. Introduction 400 II. Population biology and inoculum potential of ectomycorrhizal fungi 401 III. Ectomycorrhiza development on seedlings regenerating after clearcut logging 402 IV. Which is the most important factor driving changes in the ECM fungal community after clearcut logging: inoculum loss or change in the below-ground environment? 406 V. Possible consequences for regenerating stands of species shifts in ectomycorrhizal fungi 414 VI. Conclusions 416.

Reciprocal transfer of carbon isotopes between ectomycorrhizal Betula papyrifera and Pseudotsuga menziesii.

Interspecific C transfer was studied in laboratory microcosms containing pairs of 6-month-old Betula papyrifera Marsh, and Pseudotsuga menziesii (Mirb.) Franca seedlings growing in individual, root-restrictive (28µm pore size) pouches filled with field soil. Interspecific transfer was examined by reciprocal labelling of seedlings with 13 CO2(gas) and 14 CO2(gas) . At the time of labelling, the root zones of ectomycorrhizal (EM) B. papyrifera and P. menziesii were interconnected by an extensive network of EM mycelium. Carbon transferred through EM connections was distinguished from that through soil pathways by comparing microcosms where interconnecting hyphae were left intact vs. those where they were severed immediately before labelling. Transfer was bidirectional, and represented 5 % of total isotope uptake by both B. papyrifera and P. menziesii together. P. menziesii received on average 50% more 14 C and 66% more 13 C from paper birch than vice versa, however, differences between species were not statistically significant. Neither net nor bidirectional transfer differed between severing treatments, leaving in question the relative importance of EM hyphae versus soil transfer pathways. The tendency for P. menziesii to receive more isotope than B. papyrifera corresponded with a 10-fold greater net photosynthetic rate per seedling and two-fold greater foliar N concentration of B. papyrifera than P. menziesii.

Ectomycorrhizal fungi contribute to soil organic matter cycling in sub-boreal forests.

Soils of northern temperate and boreal forests represent a large terrestrial carbon (C) sink. The fate of this C under elevated atmospheric CO2 and climate change is still uncertain. A fundamental knowledge gap is the extent to which ectomycorrhizal fungi (EMF) and saprotrophic fungi contribute to C cycling in the systems by soil organic matter (SOM) decomposition. In this study, we used a novel approach to generate and compare enzymatically active EMF hyphae-dominated and saprotrophic hyphae-enriched communities under field conditions. Fermentation-humus (FH)-filled mesh bags, surrounded by a sand barrier, effectively trapped EMF hyphae with a community structure comparable to that found in the surrounding FH layer, at both trophic and taxonomic levels. In contrast, over half the sequences from mesh bags with no sand barrier were identified as belonging to saprotrophic fungi. The EMF hyphae-dominated systems exhibited levels of hydrolytic and oxidative enzyme activities that were comparable to or higher than saprotroph-enriched systems. The enzymes assayed included those associated with both labile and recalcitrant SOM degradation. Our study shows that EMF hyphae are likely important contributors to current SOM turnover in sub-boreal systems. Our results also suggest that any increased EMF biomass that might result from higher below-ground C allocation by trees would not suppress C fluxes from sub-boreal soils.