The microbial communities of aspen and spruce forest floors are resistant to changes in litter inputs and microclimate

Previous studies have shown that forest floors from stands dominated by trembling aspen (ASPEN; Populus tremuloides Michx.) tend to support a greater microbial biomass with a different microbial community structure than forest floors from stands dominated by white spruce (SPRUCE; Picea glauca (Moench) Voss). A reciprocal transfer experiment, in concert with coarse and fine mesh bags that allowed or excluded fine root in-growth, was used to examine how the composition of these forest floor microbial communities respond to changes in belowground inputs from fine roots, aboveground inputs (e.g. from litter and through-fall) and soil microclimatic conditions over 1 year. Neither the microbial biomass nor the microbial community structure (assessed using phospholipid fatty acid analyses and substrate-induced respiration techniques) of forest floors of ASPEN or SPRUCE origin were altered by reciprocal transfer to SPRUCE or ASPEN stands, with or without fine root inputs. Despite the lack of changes in microbial community structure, the stand type during incubation had a strong effect on forest floor moisture content and concentrations of nitrate, while mesh size had a significant effect on forest floor pH and the abundance of mesofauna. Thus, changes in microbial community structure did not co-occur with changes in other characteristics of these forest floors. The resistance of the forest floor microbial communities to change may be a function of the high C contents of these soils. Further treatment effects may have been detected if the study had been extended beyond 1 year. Reciprocal transfer studies using coarse and fine mesh bags allow transferred soils to respond to fluctuations in microclimate, organic inputs and soil biota and, therefore, hold considerable promise for studies examining the influence of disturbances on soil properties.     

Composition of the microbial communities in the mineral soil under different types of natural forest

Phospholipid fatty acid (PLFA) patterns were used to describe the composition of the soil microbial communities under 12 natural forest stands including oak and beech, spruce-fir-beech, floodplain and pine forests. In addition to the quantification of total PLFAs, soil microbial biomass was measured by substrate-induced respiration and chloroform fumigation-extraction. The forest stands possess natural vegetation, representing an expression of the natural site factors, and we hypothesised that each forest type would support a specific soil microbial community. Principal component analysis (PCA) of PLFA patterns revealed that the microbial communities were compositionally distinct in the floodplain and pine forests, comprising azonal forest types, and were more similar in the oak, beech and spruce-fir-beech forests, which represent the zonal vegetation types of the region. In the nutrient-rich floodplain forests, the fatty acids 16:1ω5, 17:0cy, a15:0 and a17:0 were the most prevalent and soil pH seemed to be responsible for the discrimination of the soil microbial communities against those of the zonal forest types. The pine forest soils were set apart from the other forest soils by a higher abundance of PLFA 18:2ω6,9, which is typical of fungi and may also indicate ectomycorrhizal fungi associated with pine trees, and high amounts of PLFA 10Me18:0, which is common in actinomycetes. These findings suggest that the occurrence of azonal forest types at sites with specific soil conditions is accompanied by the development of specific soil microbial communities. The study provides information on the microbial communities in undisturbed forest soils which may facilitate interpretation of data derived from managed or even damaged or degraded forests.     

Microbial communities in forest floors under four tree species in coastal British Columbia

We compare forest floor microbial communities in pure plots of four tree species (Thuja plicata, Tsuga heterophylla, Pseudotsuga menziesii, and Picea sitchensis) replicated at three sites on Vancouver Island. Microbial communities were characterised through community level physiological profiles (CLPP), and profiling of phospholipid fatty acids (PLFA).Microbial communities from cedar forest floors had higher potential C utilisation than the other species. The F layer of the forest floor under cedar contained significantly higher bacterial biomass (PLFA) than the F layer under the other three tree species. There were differences in microbial communities among the three sites: Upper Klanawa had the highest bacterial biomass and potential C utilisation; this site also had the highest N availability in the forest floors. Forest floor H layers under hemlock and Douglas-fir contained greater biomass of Gram positive, Gram negative bacteria and actinomycetes than F layers based on PLFA, and H layers under spruce contained greater biomass of Gram negative bacteria than F layers. There were no significant differences in bacterial biomass between forest floor layers under cedar. Fungal biomass displayed opposite trends to bacteria and actinomycetes, being lowest in cedar forest floors, and highest in the F layer and at the site with lowest N availability. There were also differences in community composition among species and sites, with cedar forest floors having a much lower fungal:bacterial ratio than spruce, hemlock and Douglas-fir. The least fertile Sarita Lake site had a much greater fungal:bacterial ratio than the more fertile San Juan and Upper Klanawa sites. Forest floor layer had the greatest effect on microbial community structure and potential function, followed by site, and tree species. The similarity in trends among measures of N availability and microbial communities is further evidence that these techniques provide information on microbial communities that is relevant to N cycling processes in the forest floor.     

Microbial function in adjacent subtropical forest and agricultural soil

Soil microbial communities and their activities are altered by land use change; however impacts and extent of these alterations are often unclear. We investigated the functional responses of soil microbes in agricultural soil under sugarcane and corresponding native soil under Eucalyptus forest to additions of contrasting plant litter derived from soybean, sugarcane and Eucalyptus in a microcosm system, using a suite of complimentary techniques including enzyme assays and community level physiological profiles (CLPP). Initially agricultural soil had 50% less microbial biomass and lower enzyme activities than forest soil, but significantly higher nitrification rates. In response to litter addition, microbial biomass increased up to 11-fold in agricultural soil, but only 1.8-fold in forest soil, suggesting a prevalence of rapidly proliferating ‘r’ and slower growing ‘K’ strategists in the respective soils. Litter-driven change in microbial biomass and activities were short lived, largely returning to pre-litter addition levels by day 150. Decomposition rates of sugarcane and soybean litter as estimated via CO₂ production were lower in agricultural than in forest soil, but decomposition of more recalcitrant Eucalyptus litter was similar in both soils, contradicting the notion that microbial communities specialise in decomposing litter of the dominant local plant species. Enzyme activities and community level physiological profiles (CLPP) were closely correlated to microbial biomass and overall CO₂ production in the agricultural soil but not the forest soil, suggesting contrasting relationships between microbial population dynamics and activity in the two soils. Activities of enzymes that break down complex biopolymers, such as protease, cellulase and phenol oxidase were similar or higher in the agricultural soil, which suggests that the production of extracellular biopolymer-degrading enzymes was not a factor limiting litter decomposition. Enzyme and CLPP analyses produced contrasting profiles of microbial activity in the two soils; however the combination of both analyses offers additional insights into the changes in microbial function and community dynamics that occur after conversion of forest to agricultural land.     

The relationships among microbial parameters and the rate of organic matter mineralization in forest soils,as influenced by forest type

Vegetation type influences the rate of accumulation and mineralization of organic matter in forest soil, mainly through its effect on soil microorganisms. We investigated the relationships among forest types and microbial biomass C (MBC), basal respiration (R_B), substrate-induced respiration (R_S), N mineralization (N_min), specific growth rate μ, microbial eco-physiology and activities of seven hydrolytic enzymes, in samples taken from 25 stands on acidic soils and one stand on limestone, covering typical types of coniferous and deciduous forests in Central Europe. Soils under deciduous trees were less acidic than soils of coniferous forests, which led to increased mineralizing activities R_B and N_min, and a higher proportion of active microbial biomass (R_S/MBC) in the Of horizon. This resulted in more extractable organic C (0.5 M K₂SO₄) in soils of deciduous forests and a higher accumulation of soil organic matter (SOM) in coniferous forest soil. No effect of forest type on the microbial properties was detected in the Oh horizon and in the 0–10 cm layer. The microbial quotient (MBC/C_org), reflecting the quality of organic matter used for microbial growth, was higher in deciduous forests in all three layers. The metabolic quotient qCO₂ (R_B/MBC) and the specific growth rate μ, estimated using respiration growth curves, did not differ in soils of both forest types. Our results showed that the quality of SOM in coniferous forests supported microorganisms with higher activities of β-glucosidase, cellobiosidase and β-xylosidase, which suggested the key importance of fungi in these soils. Processes mediated by bacteria were probably more important in deciduous forest soils with higher activities of arylsulphatase and urease. The results from the stand on limestone showed that pH had a positive effect on microbial biomass and SOM mineralization.     

Forest floor microbial communities in relation to stand composition and timber harvesting in northern Alberta

With the growing interest in silvicultural techniques that more closely emulate natural disturbance regimes, there is a need to better understand how partial harvesting affects the soil microbial community in stands with varying ecological characteristics, e.g., tree species composition. Four and a half and 5.5 years post-harvest, we used phospholipid fatty acid (PLFA) and substrate-induced respiration (SIR) analyses to compare the microbial biomass and microbial community structure of forest floors from stands dominated by white spruce (Picea glauca; SPRUCE) or by trembling aspen (Populus tremuloides; ASPEN) and from mixed-species (MIXED) stands in northern Alberta, Canada, that had been clearcut, partial-cut with 20% retention, partial-cut with 50% retention or left uncut (controls). PLFA and SIR analyses revealed that ASPEN forest floors supported a larger microbial biomass with a very different community structure than MIXED or SPRUCE forest floors. The microbial community structure of these soils appeared to be strongly affected by the presence of white spruce and the composition of the understory vegetation. There were no effects of timber harvesting detected within or across stand types on any of the variables measured, with the exception of the PLFA 16:1ω5, which was relatively more abundant in the clearcuts and 50% retention treatments than in the uncut controls, perhaps in response to an increased forest floor pH and grass cover in the disturbed areas. The resilience to timber harvesting of the forest floors from these stands may be the result of efforts to minimize soil disturbance during harvesting and to allow vegetation to regenerate naturally. From the perspective of the forest floor microbial community, partial harvesting does not appear to have any benefit over clearcut harvesting at these boreal forest sites.     

Soil microbial communities and activities in sand dunes of subtropical coastal forests

To understand the soil microbial activities and community structures in different forests in a sand-dune ecosystem, we conducted a study of 2 topographic conditions, upland and lowland, under a Casuarina forest. As well, in the lowland site, we compared forest soil microbial properties under 3 coastal forests (Casuarina, Hibiscus and mixed stand). The soil microbial biomass did not significantly differ between the upland and lowland Casuarina forest sites. At the lowland site, the soil microbial biomass was higher in the Hibiscus than Casuarina forest soil. Cellulase, xylanase, phosphatase and urease activities did not show a consistent trend by topography or vegetation. Analysis of phospholipid fatty acids (PLFAs) of bacteria and actinomycetes revealed a significant difference in microbial community structure by both topography and vegetation. PLFA content was higher at upland than lowland sites in the Casuarina forest. At the lowland site, the level of PLFAs was higher in Hibiscus than Casuarina forest soil. In addition, we examined the ratios 16:1ω7t/16:1ω7c and, cy17:0/16:1ω7c as indicators of physiological stress; the soil in the Casuarina forest had the highest values, which suggests that the microbial community in the Casuarina forest soil is under physiological stress or starvation conditions. Comparison of soil microbial properties suggest that planting Hibiscus may help to enrich soil fertility and increase microbial activities in coastal sand-dune Casuarina forest.     

Timber harvesting alters soil carbon mineralization and microbial community structure in coniferous forests

Timber harvesting influences both above and belowground ecosystem nutrient dynamics. Impact of timber harvesting on soil organic matter (SOM) mineralization and microbial community structure was evaluated in two coniferous forest species, ponderosa pine (Pinus ponderosa) and lodgepole pine (Pinus contorta). Management of ponderosa pine forests, particularly even-aged stand practices, increased the loss of CO₂-C and hence reduced SOM storage potential. Changes in soil microbial community structure were more pronounced in ponderosa pine uneven-aged and heavy harvest stands and in lodgepole pine even-aged stand as compared to their respective unmanaged stands. Harvesting of trees had a negative impact on SOM mineralization and soil microbial community structure in both coniferous forests, potentially reducing coniferous forest C storage potential.     

Soil physicochemical and microbial drivers of temperature sensitivity of soil organic matter decomposition under boreal forests

Soil organic matter(SOM)in boreal forests is an important carbon sink.The aim of this study was to assess and to detect factors controlling the temperature sensitivity of SOM decomposition.Soils were collected from Scots pine,Norway spruce,silver birch,and mixed forests(O horizon)in northern Finland,and their basal respiration rates at five different temperatures(from 4 to 28℃)were measured.The Q_(10) values,showing the respiration rate changes with a 10℃ increase,were calculated using a Gaussian function and were based on temperature-dependent changes.Several soil physicochemical parameters were measured,and the functional diversity of the soil microbial communities was assessed using the MicroResp?method.The temperature sensitivity of SOM decomposition differed under the studied forest stands.Pine forests had the highest temperature sensitivity for SOM decomposition at the low temperature range(0–12℃).Within this temperature range,the Q_(10) values were positively correlated with the microbial functional diversity index(H'_(mic))and the soil C-to-P ratio.This suggested that the metabolic abilities of the soil microbial communities and the soil nutrient content were important controls of temperature sensitivity in taiga soils.     

Forest floor development and biochemical properties in reconstructed boreal forest soils

Following resource extraction by surface mining in the oil sands region of northeastern Alberta, sites are reclaimed by reconstructing soils using a variety of salvaged organic and mineral materials, and planted to native tree species. This study assessed the influence of three distinct stand types (Populus tremuloides Michx., Pinus banksiana Lamb., and Picea glauca (Moench) Voss) on forest floor development (thickness, morphology, total carbon and nitrogen contents), soil organic matter composition, and associated soil microbial communities. Forest floor and top mineral soil (0–5 cm) samples were collected from 32 sites reclaimed 16–33 years ago. Soil organic matter composition was measured using ramped-cross-polarization ¹³C nuclear magnetic resonance, and microbial communities were characterized using phospholipid fatty acid analysis. Morphological characteristics indicated little mesofaunal or fungal activities within the forest floors. Stands dominated by P. tremuloides fostered more rapid forest floor development than the coniferous (P. banksiana and P. glauca) stands, and showed a significant increase in forest floor thickness with time since reclamation. Within the P. tremuloides stands, forest floor development was accompanied by temporal changes in soil organic matter composition that reflected inputs from the canopy. Soil microbial community composition differed among reclamation treatments of the reconstructed soils, specifically as a function of their subsoil mineral textures, when canopy cover was below 30%. Above 30%, significant differences became apparent among stand types. Taken together, our results document how canopy cover and stand type were both important factors for the reestablishment of plant–soil relationships at these sites. Furthermore, achieving a canopy cover of 30% emerged as a critical threshold point during soil reclamation.     

不同植被群落下喀斯特土壤养分及生物化学性质特征

为了探讨植被群落对喀斯特土壤养分和生物化学性质的影响,对贵州茂兰国家级自然保护区不同植被群落下土壤养分、基础呼吸、微生物量碳及酶活性进行比较研究。结果表明:不同植被群落之间土壤养分含量差异显著,土壤总有机碳、全氮、全磷、碱解氮和有效磷表现为裸露地<草丛<灌丛≤乔林;土壤基础呼吸表现为裸露地<灌丛<乔林<草丛,诱导微生物量碳为裸露地<灌丛<草丛=乔林,而熏蒸微生物量碳为裸露地<草丛<灌丛<乔林;不同植被群落下土壤蔗糖酶活性高于裸露地,但不同植被群落间差异不显著,土壤脲酶活性有着裸露地<草丛=灌丛<乔林的趋势,而土壤碱性磷酸酶活性表现为裸露地<灌丛<乔林<草丛;土壤呼吸商和诱导微生物量与熏蒸微生物量比值均表现为乔林=灌丛<草丛=裸露地。结果分析表明,不同植被群落下喀斯特土壤养分水平及其凋落物差异影响着土壤微生物群落结构及活性,而土壤微生物执行着土壤营养元素生物化学过程,其群落结构和活性又影响着土壤养分水平及养分有效性。     

Pasture and forest soil microbial communities show distinct patterns in their catabolic respiration responses at a landscape scale

Catabolic responses to specific substrates can be used to differentiate soil microbial communities. We hypothesized that the catabolic respiration responses of microbial communities from pastures would differ from those of forest soils, and that the differences would be consistent at a landscape scale, due to inherent differences in litter quality and management regimes. We analysed respiration responses to 25 different substrates of 20 pasture soils (dominated by rye grass and white clover) and 20 forest soils (indigenous forest species or the plantation species Pinus radiata) over a wide geographical range in New Zealand.Within each pasture or forest category, the catabolic responses showed a similar pattern, suggesting similarities in functional catabolic capability and microbial community Indigenous forests and pine forests microbial communities did not differ in their responses. Pasture soil communities had significantly higher relative responses to carbohydrate and amino-acid substrates and significantly lower relative response to carboxylic acid substrates, than microbial communities from forest soils. Forest soils had relatively greater responses to carboxylic acids as a group, as well as citric acid, α-ketobutyric acid, α-ketoglutaric acid, and α-ketovaleric acid, than did the pasture soils. A subset of 6 substrates was equally as effective at differentiating the microbial catabolic response of pasture soils from forest soils as the entire set of 25 substrates. The results demonstrated distinct differences in the respiration responses of the soil microbial communities of pastures and forests, but showed strong similarities within each vegetation class, despite the wide geographical spread, different soils and plant species.     

Spatial variability of enzyme activities and microbial biomass in the upper layers of Quercus petraea forest soil

Extracellular lignocellulose-degrading enzymes are responsible for the transformation of organic matter in hardwood forest soils. The spatial variability on a 12 × 12 m plot and vertical distribution (0–8 cm) of the ligninolytic enzymes laccase and Mn-peroxidase, the polysaccharide-specific hydrolytic enzymes endoglucanase, endoxylanase, cellobiohydrolase, 1,4-β-glucosidase, 1,4-β-xylosidase and 1,4-β-N-acetylglucosaminidase and the phosphorus-mineralizing acid phosphatase were studied in a Quercus petraea forest soil profile. Activities of all tested enzymes exhibited high spatial variability in the L and H horizons. Acid phosphatase and 1,4-β-N-acetylglucosaminidase exhibited low variability in both horizons, while the variability of Mn-peroxidase activity in the L horizon, and endoxylanase and cellobiohydrolase activities in the H horizon were very high. The L horizon contained 4× more microbial biomass (based on PLFA) and 7× fungal biomass (based on ergosterol content) than the H horizon. The L horizon also contained relatively more fungi-specific and less actinomycete-specific PLFA. There were no significant correlations between enzyme activities and total microbial biomass. In the L horizon cellulose and hemicellulose-degrading enzymes correlated with each other and also with 1,4-β-N-acetylglucosaminidase and acid phosphatase activities. Laccase, Mn-peroxidase and acid phosphatase activities correlated in the H horizon. The soil profile showed a gradient of pH, organic carbon and humic compound content, microbial biomass and enzyme activities, all decreasing with soil depth. Ligninolytic enzymes showed preferential localization in the upper part of the H horizon. Differences in enzyme activities were accompanied by differences in the microbial community composition where the relative amount of fungal biomass decreased and actinomycete biomass increased with soil depth. The results also showed that the vertical gradients occur at a small scale: the upper and lower parts of the H horizon only 1 cm apart were significantly different with respect to seven out of nine activities, microbial biomass content and community composition.     

城市与郊区森林土壤微生物群落特征差异研究

森林是具有重要服务功能的生态系统之一,在城区和郊区都普遍存在,但对于森林土壤微生物群落特征的城郊差异研究却很少。本研究在北京城市与郊区选择三种相同树种(油松Pinus tabulaeformis、侧柏Platycladus orientalis和毛白杨Populus tomentosa)的森林,主要研究比较了土壤微生物量碳、微生物群落结构和功能多样性的城郊差异。结果表明,郊区森林的土壤微生物量碳含量显著高于城区(郊区微生物量碳为367.29 mg kg-1,城区为294.23 mg kg-1)。同一树种下郊区微生物活性高于城区,差异未达到显著。微生物多样性和均匀度城郊差异不明显;但树种间存在一定的差异,城区毛白杨林下土壤微生物群落的均匀度显著低于柏树与油松,多样性指数显著低于柏树。结果表明城市化过程对城市土壤微生物量以及微生物群落的结构和功能有一定影响,且不同树种对于这种影响存在差异。随着城市化加快,城乡梯度上土壤微生物量碳的城乡差异明显,而土壤微生物群落结构、功能的差异有减小的趋势。     

Measurements of microbial community activities in individual soil macroaggregates

The functional potential of single soil macroaggregates may provide insights into the localized distribution of microbial activities better than traditional assays conducted on bulk quantities of soil. Thus, we scaled down enzyme assays for β-glucosidase, N-acetyl-β-d-glucosaminidase, lipase, and leucine aminopeptidase to measure of the enzyme potential of individual macroaggregates (250–1000 μm diameter). Across all enzymes, the smallest macroaggregates had the greatest activity and the range of enzyme activities observed in all macroaggregates supports the hypothesis that functional potential in soil may be distributed in a patchy fashion. Paired analyses of ATP as a surrogate for active microbial biomass and β-glucosidase on the same macroaggregates suggest the presence of both extracellular β-glucosidase functioning in macroaggregates with no detectable ATP and also of relatively active microbial communities (high ATP) that have low β-glucosidase potentials. Studying function at a scale more consistent with microbial habitat presents greater opportunity to link microbial community structure to microbial community function.     

Temporal dynamics and variation with forest type of phospholipid fatty acids in litter and soil of temperate forests across regions

Microorganisms form the basis of soil food webs and represent key control points of carbon cycling and sequestration. Virtually all central European forests are managed and land-use regimes likely impact microbial abundance and community composition. Consequently, knowledge on how land-use intensity and abiotic variables, such as pH, C-to-N ratios, moisture regimes and concomitantly different stress levels, affect microbial communities is needed. We investigated phospholipid fatty acid (PLFA) profiles of leaf litter and soil from four forest types differing in foliage, age and management intensity, replicated in three regions across Germany. To account for temporal variation, samples were taken twice in the same season, but with an interval of three years. Total microbial biomass and microbial community composition differed between years, presumably due to between year variations in weather conditions. The litter layer was more prone to effects of drying, with a reduction of almost 30% of total PLFAs in the drier year. In soil effects of weather conditions depended on soil type and therefore differed between regions, with microorganisms in the sandy soils of the Schorfheide being more susceptible to water-stress, as evidenced by a ten-fold increase of the stress indicator cy/pre ratio in the drier year. Despite temporal variations in microbial biomass and community composition, the balance between the fungal and bacterial energy channel, as measured by fungal-to-bacterial ratios, remained rather constant in particular in soil. While total microbial biomass did not differ between forest types, microbial community composition differed significantly between beech and coniferous forests. Despite more acidic conditions, the fungal energy channel was less pronounced in leaf litter of coniferous forests than in broad-leaved forests, whereas the proportion of bacterial fatty acids was the highest in coniferous forests. Increasing management intensity presumably fosters the bacterial energy channel in the exposed litter layer. Supporting this assumption coniferous forests featured significantly higher values of the stress indicators cy/pre and SAT/MONO ratio. Bacterial community structure and biomass closely correlated with pH, with particular PLFAs dominating at high and low pH, respectively, indicating pH-specific microbial communities. In contrast, fungal abundance in leaf litter was correlated with C-to-N ratio. The results suggest that leaf litter and soil need to be considered separately when investigating changes in microbial community composition, since susceptibility of microorganisms to environmental stressors differs markedly between these layers. This, and repeated sampling events, may be particularly important when investigating subtle effects such as those related to climate change.     

高黎贡山不同森林类型土壤肥力状况研究

以高黎贡山7种不同森林类型（亚高山草甸、灌丛、铁杉林、中山湿性常绿阔叶林、季风常绿阔叶林、热带沟谷雨林、干热河谷）土壤为研究对象,通过对林地土壤不同层次pH、有机质及全氮、全磷、全钾及主要速效养分含量的测定,分析不同森林类型下土壤养分的差异状况。结果表明,除干热河谷土壤pH值呈碱性（平均为7.10）,其余6个森林类型土壤pH值均介于4.09～5.26,呈酸性。且随深度增加,pH呈现出稍微增大的趋势（除干热河谷外）,土壤有机质、全氮、全磷、速效氮、速效磷含量均呈下降趋势。研究区总体上呈现出土壤有机质、全氮、全磷和速效性养分含量较高、土壤肥力较好,但不同森林类型土壤的肥力特征以及同一森林类型不同层次土壤的肥力特征均存在着差异。通过主成分分析计算各森林类型综合得分并进行排序,土壤肥力状况结果表明:亚高山草甸 > 灌丛 > 中山湿性常绿阔叶林 > 铁杉林 > 季风常绿阔叶林 > 热带沟谷雨林 > 干热河谷。     

Rhododendron thickets alter N cycling and soil extracellular enzyme activities in southern Appalachian hardwood forests

Rhododendron maximum L., a spreading understory shrub, inhibits overstory regeneration and alters forest community structure in southern Appalachian hardwood forests. Using paired plots and reciprocal litter transplants in forests with and without R. maximum cover, we examined the influence of R. maximum on litter mass and quality, N cycling and soil extracellular enzymes. Standing stocks of soil organic matter, soil N, leaf litter mass and fine root biomass were greater in forests with R. maximum than those without. Tannin extracts from R. maximum foliage, and leaf litter and fine roots collected under R. maximum had a relatively high capacity to precipitate protein compared to extracts from trees. Across the growing season, soil inorganic N availability was generally lower under R. maximum, mostly due to reduced NO₃⁻ availability. Our data suggest that R. maximum litter alters N cycling through the formation of recalcitrant polyphenol–organic N complexes. Soil extracellular enzymes indicate the potential processing rates of organic substrates. Between forest types, polyphenol oxidase activity was greatest in R. maximum O horizons, regardless of litter type, suggesting that the local microbial community can better degrade and access protein–tannin-complexed N. Protease activity did not differ between forest types, but was greater on R. maximum leaf litter than hardwood leaf litter. The alteration of the N cycle via the formation of polyphenol–organic N complexes may contribute to hardwood seedling suppression, while the enzymatic release of these complexes by ericoid mycorrhizal fungi may increase N acquisition for R. maximum and contribute to its expansion in southern Appalachian forests.     

Effects of tree species and N additions on forest floor microbial communities and extracellular enzyme activities

Forest nitrogen (N) retention and soil carbon (C) storage are influenced by tree species and their associated soil microbial communities. As global change factors alter forest composition, predicting long-term C and N dynamics will require understanding microbial community structure and function at the tree species level. Because atmospheric N deposition is increasing N inputs to forested ecosystems across the globe, including the northeastern US, it is also important to understand how microbial communities respond to added N. While prior studies have examined these topics in mixed-species stands, we focused on the responses of different tree species and their associated microbial communities within a single forest type - a northern hardwood forest in the Catskills Mountains, NY. Based on prior studies, we hypothesized that N additions would stimulate extracellular enzyme activities in relatively labile litters, but suppress oxidative enzyme activities in recalcitrant litters, and tested for independent tree species effects within this context. During the 2007 growing season (May-June), we measured enzyme activities and microbial community composition (using phospholipid fatty acid analysis - PLFA) of the forest floor in single-species plots dominated by sugar maple (Acer saccharum), yellow birch (Betula alleghaniensis), red oak (Quercus rubra), American beech (Fagus grandifolia) and eastern hemlock (Tsuga canadensis), species whose litters range from relatively labile to recalcitrant. Half the plots were fertilized with N by adding NH₄NO₃ (50 kg ha⁻¹ y⁻¹) from 1997 to 2009. Non-metric multidimensional scaling (NMS) and multi-response permutation procedures (MRPP) were used to examine microbial community structure and relationship to enzyme activities.We found that in response to N additions, both microbial community composition and enzyme activities changed; however the strength of the changes were tree species-specific and the direction of these changes was and not readily predictable from prior studies conducted in mixed-species stands. For example, in contrast to other studies, we found that N additions caused a significant overall increase in fungal biomass that was strongest for yellow birch (24% increase) and weakest for sugar maple (1% increase). Contrary to our initial hypotheses and current conceptual models, N additions reduced hydrolytic enzyme activities in hemlock plots and reduced oxidative enzyme activity in birch plots, a species with relatively labile litter. These responses suggest that our understanding of the interactions between microbial community composition, enzyme activity, substrate chemistry, and nutrient availability as influenced by tree species composition is incomplete. NMS ordination showed that patterns in microbial community structure (PLFA) and function (enzyme activity) were more strongly influenced by tree species than by fertilization, and only partially agreed with the structure-function relationships found in other studies. This finding suggests that tree species-specific responses are likely to be important in determining the structure and function of northeastern hardwood forests in the future. Enhanced understanding of microbial responses to added N in single and mixed-species substrates with varying amounts of lignin and phenols may be needed for accurate predictions of future soil C and N dynamics.     

Parent material and vegetation influence bacterial community structure and nitrogen functional genes along deep tropical soil profiles at the Luquillo Critical Zone Observatory

Microbial communities mediate every step of the soil nitrogen cycle, yet the structure and associated nitrogen cycle functions of soil microbial communities remain poorly studied in tropical forests. Moreover, tropical forest soils are often many meters deep, but most studies of microbial nitrogen cycling have focused exclusively on surface soils. The objective of our study was to evaluate changes in bacterial community structure and nitrogen functional genes with depth in soils developed on two contrasting geological parent materials and two forest types that occur at different elevations at the Luquillo Critical Zone Observatory in northeast Puerto Rico. We excavated three soil pits to 140 cm at four different sites representing the four soil × forest combinations (n = 12), and collected samples at ten-centimeter increments from the surface to 140 cm. We used bacterial 16S rRNA gene-DGGE (denaturant gradient gel electrophoresis) to fingerprint microbial community structures, and quantitative PCR to measure the abundance of five functional genes involved in various soil nitrogen transformations: nifH (nitrogen fixation), chiA (organic nitrogen decomposition), amoA (ammonia oxidation), nirS (nitrite reduction) and nosZ (nitrous oxide reduction). Multivariate analyses of DGGE fingerprinting patterns revealed differences in bacterial community structure across the four soil × forest types that were strongly correlated with soil pH (r = 0.69, P < 0.01) and nutrient stoichiometry (r² ≥ 0.36, P < 0.05). Across all soil and forest types, nitrogen functional genes declined significantly with soil depth (P < 0.001). Denitrification genes (nirS and nosZ) accounted for the largest proportion of measured nitrogen functional genes. Measured nitrogen functional genes were positively correlated with soil carbon, nitrogen and phosphorus concentrations (P < 0.001) and all genes except amoA were significantly more abundant in the Inceptisol soil type compared with the Oxisol soil type (P < 0.03). Greater abundances and a stronger vertical zonation of nitrogen functional genes in Inceptisols suggest more dynamic nitrogen transformation processes in this soil type. As the first study to examine bacterial nitrogen functional gene abundances below the surface 20 cm in tropical forest soils, our work provides insight into how pedogenically-driven vertical gradients control the nitrogen-cycling capacity of soil microbial communities. While previous studies have shown evidence for redox-driven hotspots in tropical nitrogen cycling on a watershed scale, our study corroborates this finding on a molecular scale.