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.     

Contribution of amino compounds to dissolved organic nitrogen in forest soils

Dissolved organic nitrogen (DON) may play an important role in plantnutrition and nitrogen fluxes in forest ecosystems. In spite of the apparentimportance of DON, there is a paucity of information concerning its chemicalcomposition. However, it is exactly this chemical characterization that isrequired to understand the importance of DON in ecosystem processes. Theprimaryobjective of this study was to characterize the distribution of free aminoacidsand hydrolyzable peptides/proteins in the DON fraction of Oa horizon leachatesalong an extreme edaphic gradient in northern California. Insitu soil solutions were extracted by centrifugation from Oahorizonscollected beneath Pinus muricata (Bishop pine) andCupressus pygmaea (pygmy cypress) on slightlyacidic/fertile and highly acidic/infertile sites. DON accounted for 77 to99% of the total dissolved nitrogen in Oa horizon leachates. Nitrogen infree amino acids and alkyl amines ranged from 0.04–0.07 mgN/L on the low fertility site to 0.45–0.49 mg N/L onthe high fertility site, and accounted for 1.5 to 10.6% of the DON fraction.Serine, glutamic acid, leucine, ornithine, alanine, aspartic acid andmethylamine were generally the most abundant free amino compounds. Combinedamino acids released by acid hydrolysis accounted for 48 to 74% of theDON, suggesting that proteins and peptides were the main contributor to DON inOa horizon leachates. Together, nitrogen from free andcombined amino compounds accounted for 59 to 78% of the DON. Most of theDON was found in the hydrophobic fraction, which suggests the presence ofprotein/peptide-polyphenol complexes or amino compounds associated withhumic substances. Because free and combined amino acids can be an importantnitrogen source for some plants, soil DON may play an important role in plantnutrition and ecosystem function.     

Modelling organic carbon turnover in cleared temperate forest soils converted to maize cropping by using 13C natural abundance measurements

In southwest France, thick humic acid loamy soils have developed from Quaternary silty alluvial deposits. On these soils, most forest lands have been converted to continuous intensive maize cropping and the loss of C upon conversion to intensive agriculture has been shown to be significant. The objective of this study was to determine if a study of natural ¹³C abundance in soil organic C makes possible an improved modelling of organic carbon turnover in the cultivated horizons of soils in this landscape in southwest France. A chronosequence study is realized by comparing C pools and C-13 natural abundance of three forest sites and 14 adjacent agricultural sites, whose ages of cultivation ranged from 3 to 32 yr. ¹³C ratio is found to increase with time of cultivation. The fraction of C coming from the maize crop increases during the first decades of cultivation, and reaches a plateau thereafter. This equilibrium level is reached after a few decades of cultivation. The decrease of the initial C pool is fitted by a simple model assuming that about half of this pool is mineralized during the first yr of cultivation whereas the other half decreases at a slower rate. Therefore, a general bi-compartmental model is proposed for describing the soil organic carbon dynamics in these soils after forest clearing and intensive maize cropping.     

Diversity and specificity of ectomycorrhizal fungi retrieved from an old-growth Mediterranean forest dominated by Quercus ilex

We analysed the ectomycorrhizal (ECM) fungal diversity in a Mediterranean old-growth Quercus ilex forest stand from Corsica (France), where Arbutus unedo was the only other ECM host. On a 6400 m2 stand, we investigated whether oak age and host species shaped below-ground ECM diversity. Ectomycorrhizas were collected under Q. ilex individuals of various ages (1 yr seedlings; 3-10 yr saplings; old trees) and A. unedo. They were typed by ITS-RFLP analysis and identified by match to RFLP patterns of fruitbodies, or by sequencing. A diversity of 140 taxa was found among 558 ectomycorrhizas, with many rare taxa. Cenococcum geophilum dominated (35% of ECMs), as well as Russulaceae, Cortinariaceae and Thelephoraceae. Fungal species richness was comparable above and below ground, but the two levels exhibited < 20% overlap in fungal species composition. Quercus ilex age did not strongly shape ECM diversity. The two ECM hosts, A. unedo and Q. ilex, tended to share few ECM species (< 15% of the ECM diversity). Implications for oak forest dynamics are discussed.     

The capacity of soils to preserve organic C and N by their association with clay and silt particles

Although it has been recognized that the adsorption of organics to clay and silt particles is an important determinant of the stability of organic matter in soils, no attempts have been made to quantify the amounts of C and N that can be preserved in this way in different soils. Our hypothesis is that the amounts of C and N that can be associated with clay and silt particles is limited. This study quantifies the relationships between soil texture and the maximum amounts of C and N that can be preserved in the soil by their association with clay and silt particles. To estimate the maximum amounts of C and N that can be associated with clay and silt particles we compared the amounts of clay- and silt-associated C and N in Dutch grassland soils with corresponding Dutch arable soils. Secondly, we compared the amounts of clay- and silt-associated C and N in the Dutch soils with clay and silt-associated C and N in uncultivated soils of temperate and tropical regions.We observed that although the Dutch arable soils contained less C and N than the corresponding grassland soils, the amounts of C and N associated with clay and silt particles was the same indicating that the amounts of C and N that can become associated with this fraction had reached a maximum. We also observed close positive relationships between the proportion of primary particles < 20 m in a soil and the amounts of C and N that were associated with this fraction in the top 10 cm of soils from both temperate and tropical regions. The observed relationships were assumed to estimate the capacity of a soil to preserve C and N by their association with clay and silt particles. The observed relationships did not seem to be affected by the dominant type of clay mineral. The only exception were Australian soils, which had on average more than two times lower amounts of C and N associated with clay and silt particles than other soils. This was probably due to the combination of low precipitation and high temperature leading to low inputs of organic C and N.The amount of C and N in the fraction > 20 m was not correlated with soil texture. Cultivation decreased the amount of C and N in the fraction > 20 m to a greater extent than in the fraction < 20 m, indicating that C and N associated with the fraction < 20 m is better protected against decomposition.The finding of a given soil having a maximum capacity to preserve organic C and N will improve our estimations of the amounts of C and N that can become stabilized in soils. It has important consequences for the contribution of different soils to serve as a sink or source for C and N in the long term.     

Effects of forest management on soil N cycling in beech forests stocking on calcareous soils

The effects of forest management (thinning) on gross and net N conversion, the balance of inorganic N production and consumption, inorganic N concentrations and on soil microbial biomass in the Ah layer were studied in situ during eight intensive field measuring campaigns in the years 2002–2004 at three beech (Fagus sylvatica L.) forest sites. At all sites adjacent thinning plots (“T”) and untreated control plots (“C”) were established. Since the sites are characterized either by cool-moist microclimate (NE site and NW site) or by warm-dry microclimate (SW site) and thinning took place in the year 1999 at the NE and SW sites and in the year 2003 at the NW site the experimental design allowed to evaluate (1) short-term effects (years 1–2) of thinning at the NW site and (2) medium-term effects (years 4–6) of thinning under different microclimate at the SW and NE site. Microbial biomass N was consistently higher at the thinning plots of all sites during most of the field campaigns and was overall significantly higher at the SWT and NWT plots as compared to the corresponding untreated control plots. The size of the microbial biomass N pool was found to correlate positively with both gross ammonification and gross nitrification as well as with extractable soil NO₃⁻ concentrations. At the SW site neither gross ammonification, gross nitrification, gross ammonium (NH₄⁺) immobilization and gross nitrate (NO₃⁻) immobilization nor net ammonification, net nitrification and extractable NH₄⁺ and NO₃⁻ contents were significantly different between control and thinning plot. At the NET plot lower gross ammonification and gross NH₄⁺ immobilization in conjunction with constant nitrification rates coincided with higher net nitrification and significantly higher extractable NO₃⁻ concentrations. Thus, the medium-term effects of thinning varied with different microclimate. The most striking thinning effects were found at the newly thinned NW site, where gross ammonification and gross NH₄⁺ immobilization were dramatically higher immediately after thinning. However, they subsequently tended to decrease in favor of gross nitrification, which was significantly higher at the NWT plot as compared to␣the␣NWC plot during all field campaigns after␣thinning except for April 2004. This increase␣in␣gross nitrification at the NWT plot (1.73 mg N kg⁻¹ sdw day⁻¹ versus 0.48 mg N kg⁻¹ sdw day⁻¹ at the NWC plot) coincided with significantly higher extractable NO₃⁻ concentrations (4.59 mg N kg⁻¹ sdw at the NWT plot versus 0.96 mg N kg⁻¹ sdw at the NWC plot). Pronounced differences in relative N retention (the ratio of gross NH₄⁺ immobilization + gross NO₃⁻ immobilization to gross ammonification + gross nitrification) were found across the six research plots investigated and could be positively correlated to the soil C/N ratio (R = 0.94; p = 0.005). In sum, the results obtained in this study show that (1) thinning can lead to a shift in the balance of microbial inorganic N production and consumption causing a clear decrease in the N retention capacity in the monitored forest soils especially in the first two years after thinning, (2)␣the resistance of the investigated forest ecosystems to disturbances of N cycling by thinning may vary with different soil C contents and C/N ratios, e. g. caused by differences in microclimate, (3) thinning effects tend to decline with the growth of understorey vegetation in the years 4–6 after thinning.     

Assessment of the C/N ratio as an indicator of the decomposability of organic matter in forest soils

The usefulness of the C/N ratio as an indicator of the decomposability of organic matter in forest soil was assessed. The assessment was based on the relationship between the C/N ratio and the contents of soil organic carbon (SOC), soil nitrogen (total N), dissolved total organic carbon (DTOC) and dissolved inorganic nitrogen (DIN). SOC, total N, DTOC and DIN were determined in soils sampled in coniferous and coniferous–deciduous forest sites from genetic horizons of 48 soil profiles. The variability of the above soil parameters was determined and the correlation between these parameters and the C/N values were calculated. It was found that the C/N ratio in soil was shaped by the difference in the mobility of both elements, whereas the decrease in the C content in subsequent horizons was mostly higher than the decrease in the N content, which means that the C/N value decreased with the depth of a soil profile. When the loss of SOC and total N contents occurs at a similar rate, the C/N ratio is maintained at a more or less stable level despite the advancing SOM mineralization. When the rate of the carbon release from SOM differs from that of nitrogen or when there is an N input from external sources, the C/N ratio does not adequately describe the process of SOM mineralization as well. The correlation coefficients between the C/N ratio and other parameters indicate that the relationships between them are not significant or that there is no correlation at all. It was found that the percentage of DTOC in SOC seemed to be a better indicator of SOM mineralization than the C/N ratio.     

Forms and cycling of phosphorus in prairie and boreal forest soils

The distribution of soil P among inorganic and organic forms was examined in prairie and boreal forest soil profiles from Saskatchewan, Canada. A sequential extraction procedure was employed to separate P into labile and stable inorganic (P_i) and organic (P_o) fractions. Profile depth, climate, vegetation, and cultivation all had a major influence on the distribution of P which is attributed to differing intensities of pedogenic processes such as weathering and leaching, and their relationship to P transformations in the soil environment.     

有机物料中有机碳和有机氮的分解进程及分解残留率

在中国科学院沈阳生态站(埋袋法)和海伦站(砂滤管法)研究了不同有机物料中有机碳和有机氮的矿化进程、分解残留率和C/N的动态变化.结果表明:有机物料中有机碳和有机氮的分解进程可分为快、慢2个阶段;各有机物料中有机氮的矿化速率均明显低于有机碳,因而其残留率高于有机碳;有机物料C/N的下降速率也可分为快、慢2个阶段.经3～5年分解后,各种有机物料残留物的C/N趋于稳定:低C/N有机物料(猪粪)残留物的C/N值接近土壤腐殖质的C/N,约为10,已完成其腐殖化过程;高C/N有机物料残留物的C/N值处于适宜微生物活动的25左右,利于增加土壤有机质、培肥土壤.     

The effect of harvesting intensity on the fate of applied 15N-ammonium to the organic horizons of a coniferous forest in N. Wales

¹⁵N-ammonium sulphate equivalent to 0.5 kg N/ha was added as a tracer to lysimeters containing the organic horizons of an acid forest soil. The effect of logging debris (brash), vegetation and second rotationPicea sitchensis seedlings on the amount of the¹⁵N found in various soil, vegetation and leachate pools was followed over a period of 60 days. Transformation of¹⁵N-ammonium to nitrate occurred within 24 hours. Although total nitrate leachate losses were high, tracer-derived nitrate represented only 0.4%–4.2% of the applied¹⁵N-ammonium. The atom % excess of the KCI-extractable organic-N pool was initially lower than for the inorganic species but due to the large pool size, consistently represented 3–6% of the applied¹⁵N-ammonium. The similarity of the atom % excess of the ammonium and nitrate pools indicated an autotrophic nitrification pathway.A significant proportion of the¹⁵N-ammonium passed through the microbial biomass which contained between 16 and 48% of the¹⁵N-ammonium 2 days after addition of the¹⁵N-ammonium. This nitrogen was in a readily available form or short-term pool for the first two weeks (with no change in the overall biomass pool), after which the nitrogen appeared to become transformed into more stable compounds representing a long-term pool. Total recovery of the¹⁵N was between 68% and 99% for the different treatments. The presence of brash reduced microbial immobilisation of the¹⁵N-ammonium and total retention in the organic matter. This is suggested to be a consequence of greater nitrification and denitrificatiion rate in organic horizons beneath a brash covering due to different microclimatic conditions.     

Differential effects of sugar maple, red oak, and hemlock tannins on carbon and nitrogen cycling in temperate forest soils

Tannins are abundant secondary chemicals in leaf litter that are hypothesized to slow the rate of soil-N cycling by binding protein into recalcitrant polyphenol–protein complexes (PPCs). We studied the effects of tannins purified from sugar maple, red oak, and eastern hemlock leaf litter on microbial activity and N cycling in soils from northern hardwood–conifer forests of the northeastern US. To create ecologically relevant conditions, we applied tannins to soil at a concentration (up to 2 mg g⁻¹ soil) typical of mineral soil horizons. Sugar maple tannins increased microbial respiration significantly more than red oak or hemlock tannins. The addition of sugar maple tannins also decreased gross N mineralization by 130% and, depending upon the rate of application, decreased net rates of N mineralization by 50–290%. At low concentrations, the decrease in mineralization appeared to be driven by greater microbial-N immobilization, while at higher concentrations the decrease in mineralization was consistent with the formation of recalcitrant PPCs. Low concentrations of red oak and hemlock tannins stimulated microbial respiration only slightly, and did not significantly affect fluxes of inorganic N in the soil. When applied to soils containing elevated levels of protein, red oak and hemlock tannins decreased N mineralization without affecting rates of microbial respiration, suggesting that PPC formation decreased substrate availability for microbial immobilization. Our results indicate that tannins from all three species form recalcitrant PPCs, but that the degree of PPC formation and its attendant effect on soil-N cycling depends on tannin concentration and the pool size of available protein in the soil.     

Elevated amino sugar nitrogen concentrations in soils: a potential method for assessing N fertility enhancement by actinorhizal plants

No single soil nitrogen (N) assay seems to be adequate to predict N fertility. An amino sugar-nitrogen assay has been proposed as a method to estimate soil N fertility and determine N fertilization requirements of agronomic crops. Such chemical methods of estimating potentially-available soil N are appealing because of their rapidity and simplicity. In this case study an amino sugar-N assay was used in established plantings of actinorhizal woody plants to determine whether they increased soil amino sugar N concentrations in soil and by how much. Amino sugar N is derived primarily from bacterial and fungal cell walls, but not plant cells, and probably reflects N-stimulation of microbial activity in soils. It was hypothesized that actinorhizal plants would increase soil amino sugar N. Mine spoil Entisols supporting Frankia-nodulated autumn olive shrubs (Elaeagnus umbellata) had 104 mg kg^-1 of amino sugar-N while European alder (Alnus glutinosa) trees on the same soil type had amino sugar N concentrations of 98 mg kg^-1 beneath their canopies. Soil adjacent to, and between, the two plantations and lacking N-fixing plants had 54 mg kg^-1 N of soil amino sugars. At another location a cluster of European alder trees increased soil amino sugar concentrations in a fertile Mollisol by a similar amount, 45 mg kg^-1, but by a lesser proportion. This Mollisol had a high amino sugar N background level of 336.5 mg kg^-1 indicating high N fertility. Results indicated that there was no correspondence between amino sugar N concentrations and total N concentrations, nor was there correspondence between amino sugar N concentrations and total soil organic matter concentration on mine spoils. Actinorhizal plants apparently increase the amino sugar N concentration in soils and an amino sugar N assay has potential for development as an additional means of indexing N fertility enhancement by N-fixing trees.     

Formation and mineralization of organic sulfur in forest soils

The incorporation of sulfur from inorganic sulfate into organic matter was examined using³⁵S for O1, O2 and A-horizon samples from two hardwood forests located at the Coweeta hydrologic laboratory, near Franklin, NC. This temperature-dependent transformation was stimulated by increased availability of sulfate or energy and the process was inhibited by sodium azide, erythromycin and candicidin. These data suggest that bacteria and fungi mediate sulfur incorporation via the formation of the covalent linkages. The latter possibility was confirmed by characterization of the organic sulfur fraction after isolation and partial purification from O2 layer material. Evidence is presented which indicates that the sulfur of this fraction is subject to mineralization after depolymerization of the carbon matrix and methods are given for the direct and indirect assessment of potential turnover rates. The availability of sulfate from mineralization appears to depend upon the rate of incorporation of sulfur into organic matter. Thus, substantially higher levels of extractable sulfate were detected when turnover of the isolated organosulfur fraction was assayed for in the presence of azide, an inhibitor of sulfate incoporation. However, the reverse was true when turnover was monitored in the presence of glucose and succinate which stimulate sulfate incorporation.Contribution from a symposium on the role of sulfur in ecosystem processes held August 10, 1983, at the annual meeting of the A.I.B.S., Grand Forks, ND; Myron Mitchell, convenor.     

Root turnover as determinant of the cycling of C,N, and P in a dry heathland ecosystem

Root production and turnover were studied using sequential core sampling and observations in permanent minirhizotrons in the field in three dry heathland stands dominated by the evergreen dwarfshrub Calluna vulgaris and the grasses Deschampsia flexuosa and Molinia caerulea, respectively. Root biomass production, estimated by core sampling, amounted to 160 (Calluna), 180 (Deschampsia) and 1380 (Molinia) g m^-2 yr^-1, respectively. Root biomass turnover rate in Calluna (0.64 yr^-1) was lower compared with the grasses (Deschampsia: 0.96 yr^-1; Molinia 1.68yr^-1)). Root length turnover rate was 0.75–0.77 yr^-1 (Deschampsia) and 1.17–1.49 yr^-1 (Molinia), respectively. No resorption of N and P from senescing roots was observed in either species. Input of organic N into the soil due to root turnover, estimated using the core sampling data, amounted to 1.8 g N m^-2 yr^-1(^Calluna), 1.7 g N m^-2 yr^-1 (Deschampsia) and 19.7 g N m^-2 yr^-1 (Molinia), respectively. The organic P input was 0.05, 0.07 and 0.55 g P M^-2 yr^-1, respectively. Using the minirhizotron turnover estimates these values were20–22% (Deschampsia) and 11–30% (Molinia) lower.When the biomass turnover data were used, it appeared that in the Molinia stand root turnover contributed 67% to total litter production, 87% to total litter nitrogen loss and 84% to total litter phosphorus loss. For Calluna and Deschampsia these percentages were about three and two times lower, respectively.This study shows that (1) Root turnover is a key factor in ecosystem C, N, and P cycling; and that (2) The relative importance of root turnover differs between species.     

Thiobacilli and sulphate production from inorganic sulphur compounds in upper horizons of spruce forest soils

The species representation of Thiobacilli was investigated in horizons F, H and A of spruce forest at ten localities. Concentrations of Thiobacilli in the selected localities and ability of the soils to oxidize sulphur compounds to sulphate were determined. Horizons F exhibited a high oxidative activity, a lower activity was found in horizon H and the lowest one was detected in horizon A. The activities showed spring and autumn maxima. Horizons F, H and A contained 10⁴-10⁵, 10²-10³ and 10^l-10³, respectively, Thiobacilli in 1 g dry soil.Thiobacillus thioparus was detected in all three horizons from all localities,T.thiooxidans was found in all horizons F, only in some horizons H and was not detected in horizons A.T.novellus was found only in some samples without any relation to the horizons,T. denitrificans was not detected at all.     

Relating microbial community structure to functioning in forest soil organic carbon transformation and turnover

Forest soils store vast amounts of terrestrial carbon, but we are still limited in mechanistic understanding on how soil organic carbon (SOC) stabilization or turnover is controlled by biotic and abiotic factors in forest ecosystems. We used phospholipid fatty acids (PLFAs) as biomarker to study soil microbial community structure and measured activities of five extracellular enzymes involved in the degradation of cellulose (i.e., β‐1,4‐glucosidase and cellobiohydrolase), chitin (i.e., β‐1,4‐N‐acetylglucosaminidase), and lignin (i.e., phenol oxidase and peroxidase) as indicators of soil microbial functioning in carbon transformation or turnover across varying biotic and abiotic conditions in a typical temperate forest ecosystem in central China. Redundancy analysis (RDA) was performed to determine the interrelationship between individual PFLAs and biotic and abiotic site factors as well as the linkage between soil microbial structure and function. Path analysis was further conducted to examine the controls of site factors on soil microbial community structure and the regulatory pathway of changes in SOC relating to microbial community structure and function. We found that soil microbial community structure is strongly influenced by water, temperature, SOC, fine root mass, clay content, and C/N ratio in soils and that the relative abundance of Gram‐negative bacteria, saprophytic fungi, and actinomycetes explained most of the variations in the specific activities of soil enzymes involved in SOC transformation or turnover. The abundance of soil bacterial communities is strongly linked with the extracellular enzymes involved in carbon transformation, whereas the abundance of saprophytic fungi is associated with activities of extracellular enzymes driving carbon oxidation. Findings in this study demonstrate the complex interactions and linkage among plant traits, microenvironment, and soil physiochemical properties in affecting SOC via microbial regulations.     

土地利用方式转变后灰色森林土有机碳矿化的温度响应特征

通过室内培育试验,分析了土地利用方式转变后灰色森林土有机碳矿化过程及其对温度变化的响应特征.结果表明：原始林转变为农田后,0～10 cm、10～20 cm的土壤有机碳和全氮含量分别下降了68.5%、76.8%和40.5%、44.4%;而农田土壤有机碳的平均矿化速率和累积矿化量仅分别为原始林的24.4%～43.2%和9.20％～13.7％.低温条件下（<25 ℃）土壤有机碳矿化的温度敏感性显著高于高温条件下（>25 ℃）.低温条件下（<25 ℃）两种利用方式的土壤有机碳矿化对温度变化的敏感性没有显著差异;但高温条件下（>25 ℃）,农田0～10 cm土壤有机碳矿化的温度敏感性高于原始林,而农田10～20 cm土壤有机碳矿化的温度敏感性明显较低.