Alf Ekblad

Large-scale forest girdling shows that current photosynthesis drives soil respiration

The respiratory activities of plant roots, of their mycorrhizal fungi and of the free-living microbial heterotrophs (decomposers) in soils are significant components of the global carbon balance, but their relative contributions remain uncertain. To separate mycorrhizal root respiration from heterotrophic respiration in a boreal pine forest, we conducted a large-scale tree-girdling experiment, comprising 9 plots each containing about 120 trees. Tree-girdling involves stripping the stem bark to the depth of the current xylem at breast height terminating the supply of current photosynthates to roots and their mycorrhizal fungi without physically disturbing the delicate root–microbe–soil system. Here we report that girdling reduced soil respiration within 1–2 months by about 54% relative to respiration on ungirdled control plots, and that decreases of up to 37% were detected within 5 days. These values clearly show that the flux of current assimilates to roots is a key driver of soil respiration; they are conservative estimates of root respiration, however, because girdling increased the use of starch reserves in the roots. Our results indicate that models of soil respiration should incorporate measures of photosynthesis and of seasonal patterns of photosynthate allocation to roots.

Boreal forest plants take up organic nitrogen

Plant growth in the boreal forest, the largest terrestrial biome, is generally limited by the availability of nitrogen. The presumed cause of this limitation is slow mineralization of soil organic nitrogen,. Here we demonstrate, to our knowledge for the first time, the uptake of organic nitrogen in the field by the trees Pinus sylvestris and Picea abies, the dwarf shrub Vaccinium myrtillus and the grass Deschampsia flexuosa. These results show that these plants, irrespective of their different types of root–fungal associations (mycorrhiza), bypass nitrogen mineralization. A trace of the amino acid glycine, labelled with the stable isotopes 13C and 15N, was injected into the organic (mor) layer of an old successional boreal coniferous forest. Ratios of 13C:15N in the roots showed that at least 91, 64 and 42% of the nitrogen from the absorbed glycine was taken up in intact glycine by the dwarf shrub, the grass and the trees, respectively. Rates of glycine uptake were similar to those of 15N-ammonium. Our data indicate that organic nitrogen is important for these different plants, even when they are competing with each other and with non-symbiotic microorganisms. This has major implications for our understanding of the effects of nitrogen deposition, global warming and intensified forestry.

Roots and Associated Fungi Drive Long-Term Carbon Sequestration in Boreal Forest

Forest Fungi Boreal forest is one of the worlds major biomes, dominating the subarctic northern latitudes of Europe, Asia, and America. The soils of boreal forest function as a net sink in the global carbon cycle and, hitherto, it has been thought that organic matter in this sink primarily accumulates in the form of plant remains. Clemmensen et al. (p. 1615; see the Perspective by Treseder and Holden) now show that most of the stored carbon in boreal forested islands in Sweden is in fact derived from mycorrhizal mycelium rather than from plant litter. Biochemical and sequencing studies show that carbon sequestration is regulated by functional and phylogenetic shifts in the mycorrhizal fungal community. The results will need to be explicitly considered in models of the role of the boreal forest in the global carbon cycle. Reservoirs of carbon in boreal forest soils are revisited in an island chronosequence, using modeling and molecular approaches. [Also see Perspective by Treseder and Holden] Boreal forest soils function as a terrestrial net sink in the global carbon cycle. The prevailing dogma has focused on aboveground plant litter as a principal source of soil organic matter. Using 14C bomb-carbon modeling, we show that 50 to 70% of stored carbon in a chronosequence of boreal forested islands derives from roots and root-associated microorganisms. Fungal biomarkers indicate impaired degradation and preservation of fungal residues in late successional forests. Furthermore, 454 pyrosequencing of molecular barcodes, in conjunction with stable isotope analyses, highlights root-associated fungi as important regulators of ecosystem carbon dynamics. Our results suggest an alternative mechanism for the accumulation of organic matter in boreal forests during succession in the long-term absence of disturbance.

Long-term effects of river regulation on river margin vegetation

(1) The effects of river regulation on river margin vegetation were evaluated by comparing two parallel seventh order rivers, one natural and the other strongly regulated, in northern Sweden. Prior to regulation, both rivers had similar vegetation. (2) No difference between the natural and the regulated river was found in width and height (relative to the summer low-water level) of the river margin, number of substrates, and mean annual discharge. (3) Frequency distributions of species differed in that the regulated river had fewer frequent and more infrequent species. Species-richness and the percentage cover of vegetation were both lower per site in the regulated river. The proportion of annual plus biennial species-richness was higher and that of perennial species-richness lower along the regulated river. (4) Reservoirs retaining pre-regulation river margins and remnants of their former vegetation, and stretches with a modest flow regulation, were most floristically similar to the natural river. (5) Regression equations including eight independent variables explained 10-77% of the variation in species-richness in thirteen groups of plants and in plant cover for two vegetation layers. Presence of pre-regulation river margin vegetation, water-level regime, height of the river margin, and mean annual discharge were the most important variables for species-richness, while water-level regime, mean annual discharge and substrate fineness were most important for plant cover. (6) In most cases, values of species-richness were higher in natural sites and in regulated sites with remnants of pre-regulation river margin vegetation, whereas they decreased with increasing height of the river margin. Percentage cover of ground vegetation was highest in natural sites with a fine-grade substrate.

The production and turnover of extramatrical mycelium of ectomycorrhizal fungi in forest soils: role in carbon cycling

There is growing evidence of the importance of extramatrical mycelium (EMM) of mycorrhizal fungi in carbon (C) cycling in ecosystems. However, our understanding has until recently been mainly based on laboratory experiments, and knowledge of such basic parameters as variations in mycelial production, standing biomass and turnover as well as the regulatory mechanisms behind such variations in forest soils is limited. Presently, the production of EMM by ectomycorrhizal (EM) fungi has been estimated at ~140 different forest sites to be up to several hundreds of kg per ha per year, but the published data are biased towards Picea abies in Scandinavia. Little is known about the standing biomass and turnover of EMM in other systems, and its influence on the C stored or lost from soils. Here, focussing on ectomycorrhizas, we discuss the factors that regulate the production and turnover of EMM and its role in soil C dynamics, identifying important gaps in this knowledge. C availability seems to be the key factor determining EMM production and possibly its standing biomass in forests but direct effects of mineral nutrient availability on the EMM can be important. There is great uncertainty about the rate of turnover of EMM. There is increasing evidence that residues of EM fungi play a major role in the formation of stable N and C in SOM, which highlights the need to include mycorrhizal effects in models of global soil C stores.

A Comparison of Species Richness and Traits of Riparian Plants between a Main River Channel and Its Tributaries

Summary1 We examined differences in species richness and frequencies of vascular plants in the riverbank vegetation between the main channel of the Vindel River system and seven of its tributaries ...

Pine Forest Floor Carbon Accumulation in Response to N and PK Additions: Bomb 14C Modelling and Respiration Studies

The addition of nitrogen via deposition alters the carbon balance of temperate forest ecosystems by affecting both production and decomposition rates. The effects of 20 years of nitrogen (N) and phosphorus and potassium (PK) additions were studied in a 40-year-old pine stand in northern Sweden. Carbon fluxes of the forest floor were reconstructed using a combination of data on soil 14C, tree growth, and litter decomposition. N-only additions caused an increase in needle litterfall, whereas both N and PK additions reduced long-term decomposition rates. Soil respiration measurements showed a 40% reduction in soil respiration for treated compared to control plots. The average age of forest floor carbon was 17 years. Predictions of future soil carbon storage indicate an increase of around 100% in the next 100 years for the N plots and 200% for the NPK plots. As much as 70% of the increase in soil carbon was attributed to the decreased decomposition rate, whereas only 20% was attributable to increased litter production. A reduction in decomposition was observed at a rate of N addition of 30 kg C ha−1 y−1, which is not an uncommon rate of N deposition in central Europe. A model based on the continuous-quality decomposition theory was applied to interpret decomposer and substrate parameters. The most likely explanations for the decreased decomposition rate were a fertilizer-induced increase in decomposer efficiency (production-to-assimilation ratio), a more rapid rate of decrease in litter quality, and a decrease in decomposer basic growth rate.

Is growth of soil microorganisms in boreal forests limited by carbon or nitrogen availability

To study whether the biomass of soil microorganisms in a boreal Pinus sylvestris-Vaccinium vitis-idaea forest was limited by the availability of carbon or nitrogen, we applied sucrose from sugar cane, a C4 plant, to the organic mor-layer of the C3–C dominated soil. We can distinguish between microbial mineralization of the added sucrose and respiration of endogenous carbon (root and microbial) by using the C4-sucrose as a tracer, exploiting the difference in natural abundance of 13C between the added C4-sucrose (δ13C −10.8‰) and the endogenous C3–carbon (δ13C −26.6 ‰). In addition to sucrose, NH4Cl (340 kg N ha−1) was added factorially to the mor-layer. We followed the microbial activity for nine days after the treatments, by in situ sampling of CO2 evolved from the soil and mass spectrometric analyses of δ13C in the CO2. We found that microbial biomass was limited by the availability of carbon, rather than nitrogen availability, since there was a 50% increase in soil respiration in situ between 1 h and 5 days after adding the sucrose. However, no further increase was observed unless nitrogen was also added. Analyses of the δ13C ratios of the evolved CO2 showed that increases in respiration observed between 1 h and 9 days after the additions could be accounted for by an increase in mineralization of the added C4–C.

Determination of chitin in fungi and mycorrhizal roots by an improved HPLC analysis of glucosamine

A method to measure chitin content in fungi and ectomycorrhizal roots with high-performance liquid chromatography (HPLC) was developed. Measurements of fluorescence of 9-fluorenylmethylchloroformate (FMOC-CI) derivatives of glucosamine were made on acid hydrolysates of pure chitin, chitin-root mixtures and fungal-root mixtures. The method was applied on 5 isolates of ectomycorrhizal fungi, and ectomycorrhizal and non-mycorrhizal Pinus sylvestris roots. Interference from amino acids was removed by pre-treatment of samples with 0.2 N NaOH. This pre-treatment did not reduce the recovery of chitin, nor did plant material affect the recovery of chitin. The HPLC method was compared with a colorimetric chitin-method by measurements on root-fungal mixtures, with known fungal content. The HPLC method gave estimates of fungal biomass which were equal to the expected while the colorimetric method showed values significantly (p<0.001) lower than the expected. The present chitin method offers a sensitive and specific tool for the quantification of chitin in fungi and in ectomycorrhizal roots.

Analysis of δ13C of CO2 distinguishes between microbial respiration of added C4-sucrose and other soil respiration in a C3-ecosystem.

The main aim of this study was to test various hypotheses regarding the changes in δ13C of emitted CO2 that follow the addition of C4-sucrose to the soil of a C3-ecosystem. It forms part of an experimental series designed to assess whether or not the contributions from C3-respiration (root and microbial) and C4-respiration (microbial) to total soil respiration can be calculated from such changes. A series of five experiments, three on sieved (root-free) mor-layer material, and two in the field with intact mor-layer (and consequently with active roots), were performed. Both in the experiments on sieved mor-layer and the field experiments, we found a C4-sucrose-induced increase in C3-respiration that accounted for between 30% and 40% of the respiration increase 1 h after sucrose addition. When the course of C3-, C4- and total respiration was followed in sieved material over four days following addition of C4-sucrose, the initially increased respiration of C3-C was transient, passing within less than 24 h. In a separate pot experiment, neither ectomycorrhizal Pinus sylvestrisL. roots nor non-mycorrhizal roots of this species showed respiratory changes in response to exogenous sucrose. No shift in the δ13C of the evolved CO2 after adding C3-sucrose to sieved mor-layer material was found, confirming that the sucrose-induced increase in respiration of endogenous C was not an artefact of discrimination against 13C during respiration. Furthermore, we conclude that the C4-sucrose induced transient increase in C3-respiration is most likely the result of accelerated turnover of C in the microbial biomass. Thus, neither respiration of mycorrhizal roots, nor processes discriminating against δ13C were likely sources of error in the field. The estimated δ13C of evolved soil CO2 in three field experiments lay between −25.2‰ and −23.6‰. The study shows that we can distinguish between CO2 evolved from microbial mineralisation of added C4-sucrose, and CO2 evolved from endogenous carbon sources (roots and microbial respiration).