Nitrogen in soils beneath 18–65 year old stands of subtropical evergreen broad-leaved forests in Laoshan Mountains in Eastern China

Monitoring of soil nitrogen (N) cycling is useful to assess soil quality and to gauge the sustainability of management practices. We studied net N mineralization, nitrification, and soil N availability in the 0 10 cm and 11 30 cm soil horizons in east China during 2006 2007 using an in situ incubation method in four subtropical evergreen broad-leaved forest stands aged 18-, 36-, 48-, and 65-years. The proper- ties of surface soil and forest floor varied between stand age classes. C:N ratios of surface soil and forest floor decreased, whereas soil total N and total organic C, available P, and soil microbial biomass N increased with stand age. The mineral N pool was small for the young stand and large for the older stands. NO 3 - -N was less than 30% in all stands. Net rates of N mineralization and nitrification were higher in old stands than in younger stands, and higher in the 0 10 cm than in the 11 30 cm horizon. The differences were significant between old and young stands (p < 0.031) and between soil horizons (p < 0.005). Relative nitrification was somewhat low in all forest stands and declined with stand age. N trans- formation seemed to be controlled by soil moisture, soil microbial bio- mass N, and forest floor C:N ratio. Our results demonstrate that analyses of N cycling can provide insight into the effects of management distur- bances on forest ecosystems.     

Nitrogen additions inhibit nitrification in acidic soils in a subtropical pine plantation: effects of soil pH and compositional shifts in microbial groups

Plantation forests play a pivotal role in carbon sequestration in terrestrial ecosystems, but enhanced nitrogen(N) deposition in these forests may affect plantation productivity by altering soil N cycling. Hence,understanding how simulated N deposition affects the rate and direction of soil N transformation is critically important in predicting responses of plantation productivity in the context of N loading. This study reports the effects of N addition rate(0, 40, and 120 kg N ha^-1 a^-1) and form(NH_4Cl vs. NaNO_3) on net N mineralization and nitrification estimated by in situ soil core incubation and on-soil microbial biomass determined by the phospholipid fatty acid(PLFA) method in a subtropical pine plantation. N additions had no influences on net N mineralization throughout the year. Net nitrification rate was significantly reduced by additions of both NH_4Cl(71.5) and NaNO_3(47.1%) during the active growing season, with the stronger inhibitory effect at high N rates. Soil pH was markedly decreased by 0.16 units by NH_4Cl additions. N inputs significantly decreased the ratio of fungal-to-bacterial PLFAs on average by 0.28(49.1%) in November. Under NH_4Cl additions, nitrification was positively related with fungal biomass and soil pH. Under NaNO_3 additions,nitrification was positively related with all microbial groups except for bacterial biomass. We conclude that simulated N deposition inhibited net nitrification in the acidic soils of a subtropical plantation forest in China,primarily due to accelerated soil acidification and compositional shifts in microbial functional groups. These findings may facilitate a better mechanistic understanding of soil N cycling in the context of N loading.     

Methods of microbial community profiling and their application to forest soils

Advances in molecular, biochemical, and physiological techniques for studying forest soil microbial communities are making it possible to assess the diversity, composition, and functioning of these complex communities. These new approaches avoid the limitations associated with isolating bacteria and fungi in the lab and are based on DNA, membrane phospholipid fatty acids (PLFA), and carbon source utilization. There are, however, limitations associated with these newer methods that need to be appreciated when applying them and interpreting results. Applications of community profiling approaches have advanced our understanding of the functional role of microbial diversity in forest soils, controls on microbial community composition, variability of communities among and within forest ecosystems, and responses to disturbance and forest management activities. Finally, several research directions are identified with potential for greater insight into the link between the microbial community and processes in forest soils.     

冰雪灾害对森林土壤性质的影响

冰雪灾害是一种常见的自然灾害,易对森林造成巨大破坏。在全球变化加剧的背景下,冰雪灾害发生的频率和强度呈现上升趋势。文中综述了冰雪灾害后森林土壤物理性质、土壤化学性质、土壤微生物群落和土壤酶活性的变化,以便为受损森林生态系统的修复提供参考。今后的研究热点是加强对灾后森林养分循环、土壤微生物和土壤种子库的长期研究,开展土壤微生物群落、土壤呼吸和土壤理化性质相互关系及作用机理研究,运用3S技术监测不同立地条件下土壤灾后动态变化、建立更科学精准的受灾森林生态系统评估体系,以及建立生态修复模型预测冰雪灾害后的森林恢复过程。     

林窗效应研究综述

森林群落常发生一些小规模的内源干扰,从而形成林窗。林窗的形成对推动森林群落的演替更新和生态系统发展至关重要。林窗面积的大小与树木的倒伏方式和林冠冠幅及大小有关。林窗面积及林窗内位置的不同,导致其小气候和土壤理化性质等环境因子发生改变,进而影响到林窗内树种更新和物种组成、林窗植被的物种多样性及其微生物和土壤动物的种类、数量等方面。未来林窗研究重点应该放在次生林和人工林的林窗效应,林窗对森林生态系统碳储量影响机制,林窗凋落物分解因子间的相互关系、作用机理和养分循环,不同树种的林窗与最适更新面积的关系,林窗的边缘效应,林窗的土壤动物和微生物动态及过程。     

Changes in aboveground and belowground properties during secondary natural succession of a cool-temperate forest in Japan

Forest development in temperate regions is considered to be a global carbon sink. Many studies have examined forest development after harvesting or fire from aboveground (e.g., biomass) or belowground (e.g., soil nutrient) perspectives. However, few studies have explored forest development from both perspectives simultaneously in cool-temperate forests in Japan. In this study, we examined changes over 105 years in both aboveground and belowground components during secondary natural succession. The aboveground biomass increased for 50 years and reached a plateau in a 105-year-old stand. The N mineralization rate increased during succession for 50 years, but showed a decline in the 105-year-old stand due to the decrease in the nitrification rate in late succession. The percent nitrification (i.e., relative contribution of nitrification to N mineralization) decreased significantly with increasing forest stand age. The N mineralization rates had significant relationships with N concentrations of the dominant tree foliage and litter fall and with the amount of litter fall N. Meanwhile, other belowground properties (i.e., soil pH, phenol concentration, soil microbial respiration, and litter mass loss) did not show any significant relationship with forest stand age. This may be because the soil at the study sites was heterogeneous and consisted of Cambisols and Andosols, the latter of which originally has high organic matter content, and thus may have buffered the effect of the aboveground development. These results indicate that belowground N dynamics are more closely associated with aboveground development than other belowground properties in these forests.     

Response of soil microbial biomass and community composition to chronic nitrogen additions at Harvard forest

Soil microbial communities may respond to anthropogenic increases in ecosystem nitrogen (N) availability, and the microbial response may ultimately feed back on ecosystem carbon and N dynamics. We examined the long-term effects of chronic N additions on soil microbes by measuring soil microbial biomass, composition and substrate utilization patterns in pine and hardwood forests at the Harvard Forest Chronic N Amendment Study. Functional and structural genes for important N cycling processes were studied using DNA community profiles. In the O horizon soil of both stands, N additions decreased microbial biomass C as determined by chloroform fumigation-extraction. Utilization of N-containing substrates was lower in N-treated pine soils than in the controls, suggesting that N additions reduced potential microbial activity in the pine stand. Counts of fungi and bacteria as determined by direct microscopy and culture techniques did not show a clear response to N additions. Nitrogen additions, however, strongly influenced microbial community DNA profiles. The ammonia monooxygenase gene (amoA) generally was found in high N-treated soils, but not in control soils. The nifH gene for N₂-fixation was generally found in all soils, but was more difficult to amplify in the pine N-treated soil than the controls, suggesting that the population of N₂-fixers was altered by N additions. The 16S rDNA gene for Nitrobacter was found in all samples, but distinct differences among DNA profiles were observed in the pine B horizon in the control, low N, and high N-treated plots. Our findings indicate that chronic N additions decreased chloroform microbial carbon and altered microbial community profiles. These changes in microbial community structure may be an important component of the response of terrestrial ecosystems to human-accelerated N supply.     

Nitrogen dynamics during ecosystem development in tropical forest restoration

We considered whether ecological restoration using high diversity of native tree species serves to restore nitrogen dynamics in the Brazilian Atlantic Forest. We measured δ¹⁵N and N content in green foliage and soil; vegetation N:P ratio; and soil N mineralization in a preserved natural forest and restored forests of ages 21 and 52 years. Green foliage δ¹⁵N values, N content, N:P ratio, inorganic N and net mineralization and nitrification rates were all higher, the older the forest. Our findings indicate that the recuperation of N cycling has not been achieved yet in the restored forests even after 52 years, but show that they are following a trajectory of development that is characterized by their N cycling intensity becoming similar to a natural mature forest of the same original forest formation. This study demonstrated that some young restored forests are more limited by N compared to mature natural forests. We document that the recuperation of N cycling in tropical forests can be achieved through ecological restoration actions.     

Adaptation to exploit nitrate in surface soils predisposes yellow-cedar to climate-induced decline while enhancing the survival of western redcedar: A new hypothesis

Yellow-cedar (Chamaecyparis nootkatensis (D. Don) Spach) and western redcedar (Thuja plicata Donn), two valuable tree species of Pacific Northwest forests, are competitive in low productivity forests on wet, nearly saturated soils with low nitrogen (N) availability and turnover. We propose a mechanism where cedar trees survive in marginal conditions through exploiting a coupled Ca–NO₃⁻ nutrient cycle where trees assimilate N as nitrate (NO₃⁻), but must accumulate a counter-ion to NO₃⁻ such as calcium (Ca⁺²) to control their internal cell pH and provide electrochemical balance. The availability of NO₃⁻ in cedar forests is favored by increased microbial activity and shifts in microbial community composition that is conducive to N mineralization and nitrification at higher pH. Cedars influence the soils under their canopy by enriching the forest floor with calcium compounds leading to increases in pH. Cedars are also prone to precocious dehardening in the spring when N is released from freeze–thaw events in the soils and conditions appear to favor nitrifying microbial communities. Cedars must concentrate fine-root biomass near the soil surface to access Ca and NO₃⁻, but this beneficial physiological adaptation also creates a vulnerability to periodic root freezing injury that is leading to the decline and mortality of at least one of them—yellow-cedar.     

Soil nitrogen transformations varied with plant community under Nanchang urban forests in mid-subtropical zone of China

Soil N transformations using the polyvinyl chloride (PVC) closed-top tube in situ incubation method were studied in Nanchang urban forests of the mid-subtropical region of China in different months of 2007. Four plots of 20 m × 20 m were established in four different plant communities that represented typical successional stages of forest development including shrubs, coniferous forest, mixed forest and broad- leaved forest. Average concentrations of soil NH 4 + -N from January to December were not different among the four plant communities. The concentrations of soil NO 3 - -N and mineral N, and the annual rates of ammonification, nitrification and net N-mineralization under the early successional shrub community and coniferous forest were generally lower than that of the late successional mixed and broad-leaved forests (p<0.05). Similar differences among the plant communities were also shown in the relative nitrification index (NH 4 + -N/NO 3 - -N) and relative nitrification intensity (nitrification rate/net N-mineralization rate). The annual net N-mineralization rate was increased from younger to older plant communities, from 15.1 and 41.4 kg·ha -1 ·a -1 under the shrubs and coniferous forest communities to 98.0 and 112.9 kg·ha -1 ·a -1 under the mixed and broad-leaved forests, respectively. Moreover, the high annual nitrification rates (50-70 kg·ha -1 ·a -1 ) and its end product, NO 3 - -N (2.4-3.8 mg·kg -1 ), under older plant communities could increase the potential risk of N loss. Additionally, the temporal patterns of the different soil N variables mentioned above varied with different plant community due to the combined affects of natural biological processes associated withforest maturation and urbanization. Our results indicated that urban for- ests are moving towards a state of "N saturation" (extremely nitrification rate and NO 3 - -N content) as they mature.     

Nitrification and denitrification in subalpine coniferous forests of different restoration stages in western Sichuan, China

Nitrification is the biological conversion of organic or inorganic nitrogen compounds from a reduced to a more oxidized state. Denitrification is generally referred to as the microbial reduction of nitrate to nitrite and further gaseous forms of nitric oxide, nitrous oxide and molecular nitrogen. They are functionally interconnected processes in the soil nitrogen cycle that are involved in the control of long-term nitrogen losses in ecosystems through nitrate leaching and gaseous N losses. In order to better understand how nitrification and denitrification change during the process of ecosystem restoration and how they are affected by various controlling factors, gross nitrification rates and denitrification rates were determined using the barometric process separation (BaPS) technique in subalpine coniferous forests of different restoration stages. The results showed that forest restoration stage had no significant effects on gross nitrification rates or denitrification rates (One-way ANOVA (analysis of variance), p < 0.05). There was no significant difference in the temperature coefficient (Q ₁₀) for gross nitrification rate among all the forest sites (One-way ANOVA, p < 0.05). Gross nitrification rates were positively correlated with water content (p < 0.05), but not with soil pH, organic matter, total nitrogen, or C/N ratios. Denitrification rates in all the forest soils were low and not closely correlated with water content, soil pH, organic matter, or total nitrogen. Nevertheless, we found that C/N ratios obviously affected denitrification rates (p < 0.05). Results from this research suggest that gross nitrification is more responsible for the nitrogen loss from soils compared with denitrification. Translated from Journal of Plant Ecology, 2006, 30(1): 90–96 [译自: 植物生态学报]     

An alternative modelling approach to predict emissions of N<Subscript>2</Subscript>O and NO from forest soils

Emissions of N₂O from forest soils in Europe are an important source of global greenhouse gas emissions. However, influencing the emission rates by forest management is difficult because the relations and feedbacks between forest and soils are complex. Process-based models covering both vegetation and soil biogeochemical processes are frequently used to analyse emission patterns. Particularly, the simulation of soil C and N turnover processes driving N₂O production, consumption and emission from forest soils requires highly specific input data which renders their regional application difficult since at this scale, soil conditions are often not well understood. Therefore, a soil C and N model (DecoNit) has been developed which describes biogeochemical processes with a simplified structure compared to existing carbon/nitrogen models that nevertheless follows the basic physical and chemical laws involved and which allows to simulate N trace gas emissions. The DecoNit model was previously calibrated using an extensive dataset on decomposition rates of incubated plant materials, microbial dynamics and nitrification. The DecoNit model has now been embedded in a modular simulation environment (MoBiLE) where it is combined with soil water balance and forest process sub-modules. Here, we present the evaluation of MoBiLE-DecoNit with emission data of N₂O and NO from forest soils of 15 European sites and compare simulation results with a previous study in which a more complex model (PnET-N-DNDC) was used. Evaluation criteria were as follows: (1) precision of modelled annual average emission rates; (2) coherence of modelled and measured annual average and daily emissions; (3) a dynamic representation of emission rates that correspond with the observed variance of fluxes. The results show that MoBiLE-DecoNit captures average annual emission rates more precisely than the more complex model PnET-N-DNDC. Also the structural underestimation of N trace gas fluxes from forest soils was resolved. Moreover, we present evidence that the new modelling approach is also somewhat more adequate for describing inter-daily emission dynamics. The combined MoBiLE-DecoNit is therefore thought to be a promising approach to simulate forest development and greenhouse gas balances on site and regional scales.     

Post-fire vegetative dynamics as drivers of microbial community structure and function in forest soils

Soil microorganisms have numerous functional roles in forest ecosystems, including: serving as sources and sinks of key nutrients and catalysts of nutrient transformations; acting as engineers and maintainers of soil structure; and forming mutualistic relationships with roots that improve plant fitness. Although both prescribed and wildland fires are common in temperate forests of North America, few studies have addressed the long-term influence of such disturbances on the soil microflora in these ecosystems. Fire alters the soil microbial community structure in the short-term primarily through heat-induced microbial mortality. Over the long-term, fire may modify soil communities by altering plant community composition via plant-induced changes in the soil environment. In this review, we summarize and synthesize the various studies that have assessed the effects of fire on forest soil microorganisms, emphasizing the mechanisms by which fire impacts these vital ecosystem engineers. The examples used in this paper are derived primarily from studies of ponderosa pine-dominated forests of the Inland West of the USA; these forests have some of the shortest historical fire-return intervals of any forest type, and thus the evolutionary role of fire in shaping these forests is likely the strongest. We argue that the short-term effects of fire on soil microflora and the processes they catalyze are transient, and suggest that more research be devoted to linking long-term plant community responses with those of the mutually dependent soil microflora.     

Mineral cycling processes and system stability in the eucalypt forest

The effects of disturbance on the stability of the eucalypt forest are considered in terms of Bormann and Likens' (1979) schema of reorganisation, aggradation, transition and steady state phases of secondary succession.The difficulties of making reliable estimates of ecosystem nutrient reserves are discussed against a background of anisotropic nutrient distribution, inadequate knowledge of the volume of soil effectively exploited by roots, and the limitations of existing techniques for soil chemical analysis. The relative significance of the atmosphere and parent material for the accumulation of nutrient reserves is outlined and the overriding importance of the latter on soil phosphorus capital in the eucalypt forest is emphasised.The capacity of forest ecosystems to restrict changes in the ionic concentration of percolating soil water is indicated and some important facets of nutrient dynamics are described. Spatial and temporal patterns of substrate availability and microbial activity greatly complicate the measurement of organic matter decomposition and nutrient flux on the forest floor. Adaptations of eucalypts potentially significant in mineral cycling are mentioned.The role of natural agents of disturbance as determinants of pattern in plant communities is indicated, and the effects of man-induced disturbances such as clearcutting, burning of logging debris, and hazard reduction burning are discussed in the context of biogeochemical cycling. The importance of close operational control of management practices is stressed, together with the need for appropriate measures of functional stability.     

物种多样性与生产力研究进展

物种多样性作为生物多样性重要组成之一,与生产力密切相关。文中围绕目前物种多样性与森林生产力存在的4种相关关系展开论述。在大量草地生态系统中物种均匀度和丰富度的增加更有利于促进生产力,而在群落结构和组成起决定作用的自然群落中,生产力不受物种多样性的影响,在人工草地群落中物种多样性甚至会降低生产力,随着森林演替过程物种多样性和生产力呈现非线性关系。样地丰富度、林分结构、环境异质性及尺度被认为是造成不同生态系统物种多样性和生产力相关关系差异的主要因素。然而,目前相关研究主要集中于小尺度、均质生境,忽略了尺度、空间异质性以及优势树种对物种多样性和生产力关系的影响,因此亟待加强大尺度、复杂条件自然群落中物种多样性和生产力之间相关关系的研究。     

Changes in soil N mineralization and nitrification pathways along a mixed forest chronosequence

Changes in soil N mineralization pathways occurring along a full rotation cycle have received little attention to date, while tree uptake for N may change during forest ageing. The aims of this study were (i) to characterize changes in potential net N mineralization and potential net nitrification within organic layers and the topsoil (organo-mineral horizon) along a 100-year chronosequence for a temperate oak–hornbeam forest and (ii) to reveal covariances between potential net N mineralization pathways and the properties of the humic epipedon (defined as the sum of organic layers and topsoil). For that purpose, a space-for-time substitution procedure and aerobic laboratory incubation method for 28 days at 28 °C in the dark were used. In addition, acetylene and captan were used to discriminate between autotrophic and heterotrophic (bacterial and/or fungal) nitrification. Several humic epipedon properties were determined, e.g. pH, exchangeable cation concentrations, effective cation exchange capacity, total C and N, dissolved organic C and N, fungal and microbial biomass N. Potential net N mineralization and nitrification pathways changed greatly along the mixed forest chronosequence. Potential net N mineralization in the organic layers increased with stand maturation whereas potential net nitrification in the topsoil decreased significantly. Selective inhibitors revealed changes in nitrification pathways along the chronosequence, i.e. potential net nitrification was autotrophic in the topsoil while it was mainly heterotrophic within the organic layers. In the organic layer, potential net nitrification was autotrophic at the onset of the chronosequence while it appeared heterotrophic during the aggradation phase and finally fungal in mature stands. A Co-Inertia Analysis was used to reveal covariances between N mineralization pathways and humic epipedon properties. The analysis showed two functional temporal shifts within N cycling along the chronosequence, one probably controlled by organic matter quality and high competition for available N resulting in the autotrophic versus heterotrophic nitrification shift in the organic layers and one mainly controlled by (i) fine organic matter abundance, allowing high N mineralization in the organic layers and (ii) acidity inhibited autotrophic nitrification in the topsoil.     

Leaf and litter nitrogen and phosphorus in three forests with low P supply

We compared the N and P contents of the main labile components of nutrient cycles in three different forest ecosystems [a tropical evergreen forest (TEF); a tropical dry forest (TDF); and a Mediterranean temperate forest (MTF)] with low P supply. A mass-balance approach was used to estimate mean residence times for organic matter, N and P in the forest floor, and to examine the flexibility of N and P intra-system cycling in the three forest ecosystems. For this purpose, we combined published values of N and P in foliage, litterfall, forest floor litter and mineral soils in these three forest ecosystems. The results of our analysis were consistent with the widely held belief that the N content of leaves (both green and senescent) and litter increases with increasing temperatures. In contrast, the data did not support the hypothesis that leaf P content decreases with increasing temperatures and precipitation: leaf and litterfall P contents were higher in both tropical forests than they were in the temperate forest. The TEF had the highest P content of the three forests studied. The mass-balance analysis indicated that although P mineralization in the TDF can run ahead of litter decomposition stoichiometry when P is in short supply, flexibility is much reduced or absent in the TEF and the MTF. Our analysis provides additional evidence of the importance of climatic factors in forest ecosystem processes and highlights the role of flexibility in ecosystem nutrient cycling, especially for P in ecosystems with a limited P supply.     

小兴安岭红松土壤微生物群落结构综合评价模型

土壤微生物对森林生态系统的养分循环利用起着重要作用,其多样性的变化对土壤结构、土壤有机质转换、土壤肥力、土壤碳截获和植物健康等方面有重要影响,而外界干扰与环境因素又影响土壤微生物多样性。利用烘干法、凯氏定氮法和钼锑抗比色法等获得相应林地土壤容重、全氮、全磷、pH和有机质,通过平板培养法获取微生物区系组成等数据,采用SPSS19．0软件应用多元线性回归分析小兴安岭林区地带性植被阔叶红松林下土壤理化性质与微生物群落结构的变化关系,引入外界干扰与环境因子,构建出适合于不同立地条件和不同林型下的土壤微生物群落分析模型,该模型可根据具体林型立地条件及气候因素对模型进行参数修正,可对森林的不同生长发育阶段的微生物群落动态变化研究提供新思路。     

扬州茱萸湾风景区森林土壤细菌多样性研究

森林土壤微生物群落作为分解者、共生体或病原体,在调节生物地球化学循环方面起到关键作用。土壤细菌能够释放土壤无机物、矿石或有机质中的营养元素,在森林生态系统的养分循环中起着重要作用。该研究以扬州市茱萸湾风景区内6种不同植被类型的城市森林作为研究对象,对不同植被类型下土壤细菌群落结构及多样性指数进行比较研究。结果表明,水杉林土壤细菌的物种总数、菌群丰度和复杂度最高,其次是竹林和山茱萸林。水杉林的土壤细菌群落的均匀度最高,其次是竹林和杂阔林。山茱萸林独有的土壤细菌的OTUs数目最多,与其他森林群落土壤细菌的OTUs分布差异较大;在门分类水平上,该地区相对丰度平均值大于1%的优势菌门有11个。     

Shifts in the bacterial community structure and function along a vegetation gradient in the Great Xing’an Mountains

The aim of this study was to compare soil bacterial communities in the Great Xing’an Mountains that represent three dominant vegetation types (Quercus mongolica forest, shrub mixed with herb and grassland). Soil bacterial communities were analyzed by both culture-dependent physiological profiling (Biolog) and culture-independent DNA-based approaches. The Q. mongolica forest and shrub mixed with herb had higher average well color development than the grassland, and the Q. mongolica forest and shrub mixed with herb soil bacterial communities easily utilized miscellaneous and amines/amides. The bacterial community structure was distinct across the three sites. Most of Acidobacteria, Proteobacteria and Bacteroidetes were found in grassland soil, while Firmicutes was present at a higher percentage in the Q. mongolica soil. Extracellular enzyme assays indicated that the soil ecosystem in the grassland experienced altered N and P nutrient cycling dynamics. pH, available phosphorus, potassium and nitrogen were important in shaping bacterial community structure. These results suggest that vegetation type was a strong determinant of the structure and function of bacterial communities, which may subsequently lead to significant changes in ecosystem functioning.