Consistent responses of soil microbial communities to elevated nutrient inputs in grasslands across the globe

Soil microorganisms are critical to ecosystem functioning and the maintenance of soil fertility. However, despite global increases in the inputs of nitrogen (N) and phosphorus (P) to ecosystems due to human activities, we lack a predictive understanding of how microbial communities respond to elevated nutrient inputs across environmental gradients. Here we used high-throughput sequencing of marker genes to elucidate the responses of soil fungal, archaeal, and bacterial communities using an N and P addition experiment replicated at 25 globally distributed grassland sites. We also sequenced metagenomes from a subset of the sites to determine how the functional attributes of bacterial communities change in response to elevated nutrients. Despite strong compositional differences across sites, microbial communities shifted in a consistent manner with N or P additions, and the magnitude of these shifts was related to the magnitude of plant community responses to nutrient inputs. Mycorrhizal fungi and methanogenic archaea decreased in relative abundance with nutrient additions, as did the relative abundances of oligotrophic bacterial taxa. The metagenomic data provided additional evidence for this shift in bacterial life history strategies because nutrient additions decreased the average genome sizes of the bacterial community members and elicited changes in the relative abundances of representative functional genes. Our results suggest that elevated N and P inputs lead to predictable shifts in the taxonomic and functional traits of soil microbial communities, including increases in the relative abundances of faster-growing, copiotrophic bacterial taxa, with these shifts likely to impact belowground ecosystems worldwide.

Response of an invasive liana to simulated herbivory: implications for its biological control

Pre-release evaluation of the efficacy of biological control agents is often not possible in the case of many invasive species targeted for biocontrol. In such circumstances simulating herbivory could yield significant insights into plant response to damage, thereby improving the efficiency of agent prioritisation, increasing the chances of regulating the performance of invasive plants through herbivory and minimising potential risks posed by release of multiple herbivores. We adopted this approach to understand the weaknesses herbivores could exploit, to manage the invasive liana, Macfadyena unguis-cati. We simulated herbivory by damaging the leaves, stem, root and tuber of the plant, in isolation and in combination. We also applied these treatments at multiple frequencies. Plant response in terms of biomass allocation showed that at least two severe defoliation treatments were required to diminish this liana's climbing habit and reduce its allocation to belowground tuber reserves. Belowground damage appears to have negligible effect on the plant's biomass production and tuber damage appears to trigger a compensatory response. Plant response to combinations of different types of damage did not differ significantly to that from leaf damage. This suggests that specialist herbivores in the leaf-feeding guild capable of removing over 50% of the leaf tissue may be desirable in the biological control of this invasive species.

Comparative growth and biomass allocation of two varieties of cat's claw creeper, Dolichandra unguis-cati (Bignoniaceae) in Australia

Introduced as an ornamental vine, cat's claw creeper Dolichandra unguis-cati (syn. Macfadyena unguis-cati) has invaded coastal and subcoastal areas of subtropical eastern Australia. Two varieties have been indentified, one of which ('short-pod') is found throughout south-eastern Australia, while the other ('long-pod') appears to be restricted to several sites in south-eastern Queensland. We compared the growth and biomass allocation patterns of the two varieties in the field over a 22-month period to determine if a higher growth rate and/or more efficient allocation of biomass may contribute to this disparity in distribution. The long-pod variety produced greater aboveground and total biomass than the short-pod variety in both riparian and non-riparian zones. Belowground the two varieties produced a similar number of tubers and overall biomass, though the long-pod variety allocated a smaller portion of its carbon belowground. High growth rates and greater biomass allocation aboveground are characteristic of invasive species, allowing them to outcompete and crowd out existing vegetation. There was no significant site by variety interaction, an indication of consistency in variety performance across riparian and non-riparian sites. Results from our study suggest that differences in growth and biomass allocations are unlikely to have contributed to the disparity in distribution of the two varieties. Despite currently occupying a relatively small range, the long-pod variety may be a more adept invader than the short-pod variety, and could become more prevalent in the future. © 2012 CSIRO.

Forest inventory-based large-scale forest biomass and carbon budget assessment: new enhanced methods and use of remote sensing for verification

In recent years, concern has arisen over the effects of increasing carbon dioxide (CO2) in the earth's atmosphere due to the burning of fossil fuels. One way to mitigate increase in atmospheric CO2 concentration and climate change is carbon sequestration to forest vegeta-tion through photosynthesis. Comparable regional scale estimates for the carbon balance of forests are therefore needed for scientific and political purposes. The aim of the present dissertation was to improve methods for quantifying and verifying inventory-based carbon pool estimates of the boreal forests in the mineral soils. Ongoing forest inventories provide a data based on statistically sounded sampling for estimating the level of carbon stocks and stock changes, but improved modelling tools and comparison of methods are still needed. In this dissertation, the entire inventory-based large-scale forest carbon stock assessment method was presented together with some separate methods for enhancing and comparing it. The enhancement methods presented here include ways to quantify the biomass of understorey vegetation as well as to estimate the litter production of needles and branches. In addition, the optical remote sensing method illustrated in this dis-sertation can be used to compare with independent data. The forest inventory-based large-scale carbon stock assessment method demonstrated here provided reliable carbon estimates when compared with independent data. Future ac-tivity to improve the accuracy of this method could consist of reducing the uncertainties regarding belowground biomass and litter production as well as the soil compartment. The methods developed will serve the needs for UNFCCC reporting and the reporting under the Kyoto Protocol. This method is principally intended for analysts or planners interested in quantifying carbon over extensive forest areas.

Microbial communities in Pb contaminated boreal forest soil

Lead contamination in the environment is of particular concern, as it is a known toxin. Until recently, however, much less attention has been given to the local contamination caused by activities at shooting ranges compared to large-scale industrial contamination. In Finland, more than 500 tons of Pb is produced each year for shotgun ammunition. The contaminant threatens various organisms, ground water and the health of human populations. However, the forest at shooting ranges usually shows no visible sign of stress compared to nearby clean environments. The aboveground biota normally reflects the belowground ecosystem. Thus, the soil microbial communities appear to bear strong resistance to contamination, despite the influence of lead. The studies forming this thesis investigated a shooting range site at Hälvälä in Southern Finland, which is heavily contaminated by lead pellets. Previously it was experimentally shown that the growth of grasses and degradation of litter are retarded. Measurements of acute toxicity of the contaminated soil or soil extracts gave conflicting results, as enchytraeid worms used as toxicity reporters were strongly affected, while reporter bacteria showed no or very minor decreases in viability. Measurements using sensitive inducible luminescent reporter bacteria suggested that the bioavailability of lead in the soil is indeed low, and this notion was supported by the very low water extractability of the lead. Nevertheless, the frequency of lead-resistant cultivable bacteria was elevated based on the isolation of cultivable strains. The bacterial and fungal diversity in heavily lead contaminated shooting sectors were compared with those of pristine sections of the shooting range area. The bacterial 16S rRNA gene and fungal ITS rRNA gene were amplified, cloned and sequenced using total DNA extracted from the soil humus layer as the template. Altogether, 917 sequenced bacterial clones and 649 sequenced fungal clones revealed a high soil microbial diversity. No effect of lead contamination was found on bacterial richness or diversity, while fungal richness and diversity significantly differed between lead contaminated and clean control areas. However, even in the case of fungi, genera that were deemed sensitive were not totally absent from the contaminated area: only their relative frequency was significantly reduced. Some operational taxonomic units (OTUs) assigned to Basidiomycota were clearly affected, and were much rarer in the lead contaminated areas. The studies of this thesis surveyed EcM sporocarps, analyzed morphotyped EcM root tips by direct sequencing, and 454-pyrosequenced fungal communities in in-growth bags. A total of 32 EcM fungi that formed conspicuous sporocarps, 27 EcM fungal OTUs from 294 root tips, and 116 EcM fungal OTUs from a total of 8 194 ITS2 454 sequences were recorded. The ordination analyses by non-parametric multidimensional scaling (NMS) indicated that Pb enrichment induced a shift in the EcM community composition. This was visible as indicative trends in the sporocarp and root tip datasets, but explicitly clear in the communities observed in the in-growth bags. The compositional shift in the EcM community was mainly attributable to an increase in the frequencies of OTUs assigned to the genus Thelephora, and to a decrease in the OTUs assigned to Pseudotomentella, Suillus and Tylospora in Pb-contaminated areas when compared to the control. The enrichment of Thelephora in contaminated areas was also observed when examining the total fungal communities in soil using DNA cloning and sequencing technology. While the compositional shifts are clear, their functional consequences for the dominant trees or soil ecosystem remain undetermined. The results indicate that at the Hälvälä shooting range, lead influences the fungal communities but not the bacterial communities. The forest ecosystem shows apparent functional redundancy, since no significant effects were seen on forest trees. Recently, by means of 454 pyrosequencing , the amount of sequences in a single analysis run can be up to one million. It has been applied in microbial ecology studies to characterize microbial communities. The handling of sequence data with traditional programs is becoming difficult and exceedingly time consuming, and novel tools are needed to handle the vast amounts of data being generated. The field of microbial ecology has recently benefited from the availability of a number of tools for describing and comparing microbial communities using robust statistical methods. However, although these programs provide methods for rapid calculation, it has become necessary to make them more amenable to larger datasets and numbers of samples from pyrosequencing. As part of this thesis, a new program was developed, MuSSA (Multi-Sample Sequence Analyser), to handle sequence data from novel high-throughput sequencing approaches in microbial community analyses. The greatest advantage of the program is that large volumes of sequence data can be manipulated, and general OTU series with a frequency value can be calculated among a large number of samples.

Defoliation and patchy nutrient return drive grazing effects on plant and soil properties in a dairy cow pasture

Large herbivores can influence plant and soil properties in grassland ecosystems, but especially for belowground biota and processes, the mechanisms that explain these effects are not fully understood. Here, we examine the capability of three grazing mechanisms-plant defoliation, dung and urine return, and physical presence of animals (causing trampling and excreta return in patches)-to explain grazing effects in Phleum pratense-Festuca pratensis dairy cow pasture in Finland. Comparison of control plots and plots grazed by cows showed that grazing maintained original plant-community structure, decreased shoot mass and root N and P concentrations, increased shoot N and P concentrations, and had an inconsistent effect on root mass. Among soil fauna, grazing increased the abundance of fungivorous nematodes and Aporrectodea earthworms and decreased the abundance of detritivorous enchytraeids and Lumbricus earthworms. Grazing also increased soil density and pH but did not affect average soil inorganic-N concentration. To reveal the mechanisms behind these effects, we analyzed results from mowed plots and plots that were both mowed and treated with a dung and urine mixture. This comparison revealed that grazing effects on plant attributes were almost entirely explained by defoliation, with only one partly explained by excreta return. Among belowground attributes, however, the mechanisms were more mixed, with effects explained by defoliation, patchy excreta return, and cow trampling. Average soil inorganic-N concentration was not affected by grazing because it was simultaneously decreased by defoliation and increased by cow presence. Presence of cows created great spatial heterogeneity in soil N availability and abundance of fungivorous nematodes. A greenhouse trial revealed a grazing-induced soil feedback on plant growth, which was explained by patchiness in N availability rather than changes in soil biota. Our results show that grazing effects on plant attributes can be satisfactorily predicted using the effects of defoliation, whereas those on soil fauna and soil N availability need understanding of other mechanisms as well. The results indicate that defoliation-induced changes in plant ecophysiology and the great spatial variation in N availability created by grazers are the two key mechanisms through which large herbivores can control grassland ecosystems.

Urban ecosystem services at the plant-soil interface

Continuing urbanization is a crucial driver of land transformation, having widespread impacts on virtually all ecosystems. Terrestrial ecosystems, including disturbed ones, are dependent on soils, which provide a multitude of ecosystem services. As soils are always directly and/or indirectly impacted through land transformation, land cover change causes soil change. Knowledge of ecosystem properties and functions in soils is increasing in importance as humans continue to concentrate into already densely-populated areas. Urban soils often have hampered functioning due to various disturbances resulting from human activity. Innovative solutions are needed to bring the lacking ecosystem services and quality of life to these urban environments. For instance, the ecosystem services of the urban green infrastructure may be substantially improved through knowledge of their functional properties. In the research forming this thesis, the impacts of four plant species (Picea abies, Calluna vulgaris, Lotus corniculatus and Holcus lanatus) on belowground biota and regulatory ecosystem services were investigated in two different urban soil types. The retention of inorganic nitrogen and phosphorus in the plant-soil system, decomposition of plant litter, primary production, and the degradation of polycyclic aromatic hydrocarbons (PAHs) were examined in the field and under laboratory conditions. The main objective of the research was to determine whether the different plant species (representing traits with varying litter decomposability) will give rise to dissimilar urban belowground communities with differing ecological functions. Microbial activity as well as the abundance of nematodes and enchytraeid worm biomass was highest below the legume L. corniculatus. L. corniculatus and the grass H. lanatus, producing labile or intermediate quality litter, enhanced the proportion of bacteria in the soil rhizosphere, while the recalcitrant litter-producing shrub C. vulgaris and the conifer P. abies stimulated the growth of fungi. The loss of nitrogen from the plant-soil system was small for H. lanatus and the combination of C. vulgaris + P. abies, irrespective of their energy channel composition. These presumably nitrogen-conservative plant species effectively diminished the leaching losses from the plant-soil systems with all the plant traits present. The laboratory experiment revealed a difference in N allocation between the plant traits: C. vulgaris and P. abies sequestered significantly more N in aboveground shoots in comparison to L. corniculatus and H. Lanatus. Plant rhizosphere effects were less clear for phosphorus retention, litter decomposition and the degradation of PAH compounds. This may be due to the relatively short experimental durations, as the maturation of the plant-soil system is likely to take a considerably longer time. The empirical studies of this thesis demonstrated that the soil communities rapidly reflect changes in plant coverage, and this has consequences for the functionality of soils. The energy channel composition of soils can be manipulated through plants, which was also supported by the results of the separate meta-analysis conducted in this thesis. However, further research is needed to understand the linkages between the biological community properties and ecosystem services in strongly human-modified systems.

Root system traits of Norway spruce, Scots pine, and silver birch in mixed boreal forests: an analysis of root architecture, morphology, and anatomy

The aim of this thesis was to unravel the functional-structural characteristics of root systems of Betula pendula Roth., Picea abies (L.) Karst., and Pinus sylvestris L. in mixed boreal forest stands differing in their developmental stage and site fertility. The root systems of these species had similar structural regularities: horizontally-oriented shallow roots defined the horizontal area of influence, and within this area, each species placed fine roots in the uppermost soil layers, while sinker roots defined the maximum rooting depth. Large radial spread and high ramification of coarse roots, and the high specific root length (SRL) and root length density (RLD) of fine roots indicated the high belowground competitiveness and root plasticity of B. pendula. Smaller radial root spread and sparser branching of coarse roots, and low SRL and RLD of fine roots of the conifers could indicate their more conservative resource use and high association with and dependence on ectomycorrhiza-forming fungi. The vertical fine root distributions of the species were mostly overlapping, implying the possibility for intense belowground competition for nutrients. In each species, conduits tapered and their frequency increased from distal roots to the stem, from the stem to the branches, and to leaf petioles in B. pendula. Conduit tapering was organ-specific in each species violating the assumptions of the general vascular scaling model (WBE). This reflects the hierarchical organization of a tree and differences between organs in the relative importance of transport, safety, and mechanical demands. The applied root model was capable of depicting the mass, length and spread of coarse roots of B. pendula and P. abies, and to the lesser extent in P. sylvestris. The roots did not follow self-similar fractal branching, because the parameter values varied within the root systems. Model parameters indicate differences in rooting behavior, and therefore different ecophysiological adaptations between species.

Potential for increasing carbon sink in Himachal Pradesh, India

In this study, the potential for increasing the tree cover and thereby the biomass and carbon as a mitigation option of three categories of wastelands, irrespective of their tenure, are considered. The area under wastelands in Himachal Pradesh, according to NRSA (2005), is estimated to be 2.83 Mha. Among the 28 categories of wastelands reported by NRSA, only 15 categories exist in Himachal Pradesh. In the present study, three land categories are considered for estimating the mitigation potential. They include: (i) Degraded forestland, (ii) Degraded community land and (iii) Degraded and abandoned private land. Choice of species or the mix of species to be planted on the three land categories considered for reforestation is discussed. Carbon pools considered in the present study are those, which account only for aboveground biomass, belowground biomass and soil organic carbon. This study estimates the mitigation potential at the state level considering land available under more than one category. It also provides a roadmap for future work in support of mitigation analysis and implementation.

Regeneration of Juvenile Woody Plants after Fire in a Seasonally Dry Tropical Forest of Southern India

Woody tree species in seasonally dry tropical forests are known to have traits that help them to recover from recurring disturbances such as fire. Two such traits are resprouting and rapid post-fire growth. We compared survival and growth rates of regenerating small-sized individuals (juveniles) of woody tree species after dry season fire (February-March) at eight adjacent pairs of burnt and unburnt transects in a seasonally dry tropical forest in southern India. Juveniles were monitored at 3-mo intervals between August 2009 and August 2010. High juvenile survivorship (>95%) was observed in both burnt and unburnt areas. Growth rates of juveniles, analyzed at the community level as well as for a few species individually (especially fast-growing ones), were distinctly higher in burnt areas compared to unburnt areas after a fire event, particularly during the pre-monsoon season immediately after a fire. Rapid growth by juveniles soon after a fire may be due to lowered competition from other vegetative forms such as grasses, possibly aided by the availability of resources stored belowground. Such an adaptation would allow a juvenile bank to be retained in the understory of a dry forest, from where individuals can grow to a possible fire-tolerant size during favorable conditions.

农牧交错区不同植物群落的根冠比特征及土壤呼吸日动态的观测研究

本论文对草地群落地上/地下生物量构成、根冠比特征及其影响因子以及土壤呼吸测定方法比较等的国内外研究进展和主要成果进行了综述，在此基础之上对地处我国北方农牧交错带中段的内蒙古多伦县境内的18种草地群落（包括天然草地和人工草地群落）进行了相关内容的研究工作。 在2002年生长季期间，对这18个植物群落中选取的16个进行了群落学调查，测定了其地上、地下生物量，同时测定了土壤含水量、土壤容重、土壤全氮含量和土壤有机质含量。分别分析了地上、地下生物量以及根冠比（root to shoot ratio）与这些立地因子间的相关关系。同期，从18个群落中选定10个代表性群落测定其土壤呼吸速率，测定方法选用了动态红外气体分析法 (Infra red gas analysis， abbreviated as IRGA)和碱液吸收法（Alkali absorption, abbreviated as AA）。对这两种方法的测定结果进行了比较分析，同时分析了不同群落间土壤呼吸变化与土壤水分和养分状况等的相关关系。主要结论如下： ①16种植物群落的地上和地下生物量差异明显，地上生物量变化范围在80~500 g•m-2之间;相比之下，地下生物量的变化范围要大得多，16个群落中地下生物量最小的为猪毛菜群落，最大的为拂子茅群落，分别为533 g•m-2和2590 g•m-2。群落的根冠比在1.5～11.21之间，平均根冠比为 5.69。 ②土壤含水量对地上和地下生物量有着重要的影响，土壤含水量高的样地（羊草样地）较含水量低（小米蒿样地）的样地地上生物量高，反之亦然。但含水量与地下生物量之间的这种关系却不明显，即土壤含水量高的样地其地下生物量并不一定比含水量低的样地地下生物量高;根冠比与土壤含水量之间基本上呈负相关。土壤全氮含量和有机质含量与地上、地下生物量也存在着一定的正相关关系，而土壤容重却与生物量存有负相关关系;根冠比与土壤全氮、有机质和容重的关系正好与此相反，即根冠比与全氮和有机质含量呈负相关，与容重为正相关。 ③10种植物群落土壤呼吸的昼夜变化比较明显，均为单峰型曲线，主要受土壤温度的驱动，但同时也受到当日降水情况和云量、风速等气象因子的较大影响。因此，影响到这些群落土壤呼吸日动态的一致性，使得规律性并不明显。 ④用碱液吸收法和动态密闭气室法测定的10个群落的土壤呼吸速率变化范围分别为394~894mg C•m-2•d-1和313~2043 mg C•m-2•d-1，其中碱液吸收法测定结果平均为动态气室法的67.5%，明显低于动态密闭气室法。 ⑤两种测定方法具有很好的相关性，R2为0.8739。本研究中发现，在土壤呼吸速率低的情况下，两种方法的测定结果十分接近，甚至碱液吸收法的测定结果稍大于动态密闭气室法;而在土壤呼吸速率较高的情况下，动态密闭气室法测定结果则显著高于碱液吸收法。上述结果与国内外同类研究的结果高度一致，从而为校正我们以往采用碱液吸收法在该区域的测定结果提供了可靠依据。 ⑥各个群落间的土壤呼吸变化与立地土壤水分和土壤养分之间存有一定的相关关系，但并不显著，可能与这些群落土壤呼吸测定不是在同一天进行有关。

放牧对内蒙古草原群落和土壤种子库的影响及两者关系的研究

选取内蒙古草原三种主要草原类型（草甸草原、典型草原和荒漠草原）代表性群落羊草杂类草群落、羊草群落和大针茅群落、小针茅群落，应用样线法沿水分梯度研究放牧对内蒙古草原不同植物群落功能群组成、多样性、生产力以及多样性与生产力关系的影响和放牧对土壤种子库组成、大小以及多样性的影响，在此基础上，研究土壤种子库与地上植被间的关系。主要结论如下： 1 放牧对植物群落的影响 荒漠草原的放牧演替规律为小针茅群落→猪毛菜 + 小针茅群落→猪毛菜群落;典型草原为羊草或大针茅群落→糙隐子草 + 大针茅群落或克氏针茅群落→星毛委陵菜 + 糙隐子草群落;草甸草原为羊草杂类草群落→羊草 + 贝加尔针茅群落，这是不同物种对牧压的不同适应结果造成的。 放牧使4种草原群落生活型功能群组分间发生强烈的生态替代作用，但不同的群落生态替代模式不同：放牧使小针茅群落多年生丛生禾草作用减弱，一二年生草本作用增强;羊草群落和大针茅群落多年生丛生禾草、多年生根茎禾草作用减弱，多年生杂类草作用增强;羊草杂类草群落多年生根茎禾草作用减弱，多年生丛生禾草作用增强。放牧使非旱生和C3植物作用减弱，而旱生、C4植物作用增强。 放牧对4种群落物种和功能群多样性的影响随不同的群落而表现不同：物种丰富度、物种多样性、生活型多样性 和水分生态类型多样性除羊草杂类草群落外随放牧强度的加大而降低，但适度放牧增加了羊草杂类草群落的上述多样性指标。 群落地上现存量一般随放牧强度的增大而下降，但小针茅群落反之，主要与一年生植物猪毛菜的生物量迅速增加有关。除羊草群落外，0~10 cm 地下生物量随放牧强度的变化不显著;除大针茅群落外，放牧显著降低0~30 cm 地下生物量。 放牧影响下内蒙古草原植物群落生物量随水分生态类型多样性的升高而升高，其回归方程为：Y = 809 + 774x （R2=0.84, P<0.001），其中Y代表群落地上现存量和地下生物量之和，x代表群落水分生态类型多样性。 2 放牧对土壤种子库的影响 小针茅群落、大针茅群落、羊草群落和羊草杂类草群落土壤种子库组成中均以多年生杂类草为主，分别占各自群落种子库总物种数的40%、52%、54%和67%。 生活型功能群种子库密度除羊草杂类草群落外，均以一二年生草本占优势。中度放牧升高了除小针茅群落外多年生禾草种子库密度;放牧增大了小针茅群落和羊草杂类草群落一二年生草本种子库密度;除羊草杂类草群落外，放牧对多年生杂类草种子库密度影响不大;总种子库组成中，灌木半灌木和小半灌木种子库密度不大，不随取样时间和牧压而变化。 中度放牧种子库总密度最大，小针茅群落在重度放牧最大，主要是由于猪毛菜种子库密度在重度放牧突增所致。总体上，内蒙古草原4种群落在不同取样时间不同牧压下种子库总密度波动在20.8~3819.2粒/m2。 土壤种子库物种丰富度最大值一般出现在10月份，除羊草杂类草群落外，不放牧群落较放牧群落为高，中度放牧使羊草杂类草群落土壤种子库物种丰富度增加。中度放牧增加了小针茅群落、大针茅群落7月份和羊草杂类草群落各取样时间土壤种子库物种多样性。 3 地上植被与土壤种子库的关系 土壤种子库的优势种在特定时间特定放牧强度下与地上现有植被优势种一致，但一致率仅为三次取样时间不同放牧强度下总体的23.3%。 地上植被与土壤种子库物种组成相似性指数受不同取样时间的影响，一般的10月取样最大。不同放牧强度对二者间的相似性亦有影响，中度放牧提高了小针茅群落、羊草杂类草群落各取样时间和大针茅群落、羊草群落4月份的相似性指数。隔年二次萌发法提高了二者间的相似性水平。总体上相似性指数变动在0.1~0.75之间。 地上植被现存量、总密度与各取样时间土壤种子库总密度之间不存在显著的相关性。 4 对于估计土壤种子库密度、物种组成和确定种子库与地上植被间的关系，隔年二次萌发法对于弥补直接萌发法本身所具有的不足不失为一种有益的尝试。

克氏针茅群落地上/地下生物量分配及其对水热因子响应研究

碳循环研究已经成为目前生态学中的研究热点之一。草地生态系统在全球碳循环中占有重要地位，而且大部分草地处于对全球变化敏感的区域，也是研究生态系统对全球变化响应的关键地区。我国草地面积十分辽阔，占我国陆地面积的40%以上，具有极其重要的生态地位和作用。 在碳蓄积研究中，对地上碳库的估计比较容易，而对地下碳库的估计却由于方法上的局限造成估计的精度不高。因此，研究陆地生态系统中植物群落的地上/地下生物量分配模式，尤其是地下生物量的分配，对于全球碳蓄积的 研究具有重要意义。植物生长与环境因子密切相关，植物的生物量分配与环境因子也必然存在一定的相关关系，了 解植物的生物量分配与环境因子之间的关系，并建立相关模式，不但有利于减少生物量调查的工作量，而且能使碳蓄积量估计的精度大大提高。 (1) 本文通过对锡林郭勒草原上的克氏针茅植物群落进行了为期两年的地上和地下生物量观测，初步得到了克氏针茅群落地上/地下生物量分配规律： 2004年地上生物量表现为逐渐增加的趋势，6、7、8月地上生物量较低，9月显著增加，为8月的4.8倍。2005年地上生物量表现出明显的单峰曲线，8月达到峰值，为150.48 g•m-2。2004年地下生物量6月到7月明显增长，7月、8、9月逐渐减少。2005年地下生物量出现两次增加和两次减小的过程。地下生物量在不同层次的分配数量及所占比例存在差异。0~10cm分配的地下生物量最多，所占比例最大，往下各层基本表现出逐层递减的趋势。其中，0~30cm地下生物量占0~80cm总地下生物量的70％。 地下生物量与地上生物量的比值即为根冠比，根冠比也具有明显的季节变化。2004年从6月到9月根冠比表现为逐渐减小的趋势，最小根冠比值出现在9月，为9.22。2005年为明显的反抛物线形状，最小值出现在8月，为19.93。 (2) 分析克氏针茅群落地上生物量、地下生物量、根冠比三者关系的结果表明：克氏针茅草原群落地上生物量和根冠比的相关关系较为明显，而地下生物量与根冠比的相关关系较差。地上生物量与地下生物量的相关关系较好，呈正相关关系。地上生物量对根冠比变化的贡献较大，而地下生物量对根冠比变化的贡献较小。 （3）在观测生物量的同时，用小气候自动观测系统记录了同时期的降水和气温资料，通过分析克氏针茅群落地上、地下生物量及根冠比与降水、气温的关系，得出以下结论： 2004年和2005年地上生物量均与上月降水量有显著相关关系，而两年的地下生物量与降水量的相关关系较差。两年的根冠比变化与上月降水量也具有较好的相关关系。2004年地下生物量与上月平均气温的相关关系较好。2005年地下生物量与月平均气温的相关关系较差。2004和2005年地上生物量与月平均气温有很好的线性关系，但两年的相关关系正好相反。两年的根冠比与月平均气温的相关关系也表现的恰恰相反。 (4) 最后，分析了温室控制试验羊草种群的生物量分配对水热因子响应的阈值。得出以下结论： 土壤相对含水量范围为6065%、温度为26℃时的地上、地下干重都最大，表明这是羊草地上、地上部分生长最适宜的温度和土壤水分。土壤为相对含水量2530%、温度为20℃时根冠比值最大。 温度、土壤水分以及二者的交互作用对羊草根冠比的影响均达到极显著水平。但根冠比对不同温度和土壤水分的敏感度不同。根冠比在一定的温度范围和土壤相对含水量范围内发生敏感变化。

土壤有机物添加对温带半干旱草地地下生物过程的影响

大气CO2 浓度和降水量增加有可能大幅提高中国北方部分草地生态系统净初级生产力，进而导致向土壤中输送的有机物相应增加。本研究以位于内蒙古自治区东乌珠穆沁旗内的半干旱草地生态系统为研究对象，通过向10−20 cm 土层添加不同质量和数量的植物凋枯物碎屑模拟有机物输入增加和喷灌模拟降水量提升，同时测定土壤微生物群落动态和植物生长指标，探讨在增加有机物输入和土壤水分的情况下土壤生物过程的变化及其对土壤碳排放和贮存的反馈作用。 研究结果表明，有机物添加可促进植物地上部分及根系的生长，并显著提高土壤中可溶性有机碳（能量）和氮（养分）的含量。土壤能量和养分水平的提高促进了土壤微生物的生长：在底物可利用性水平较高时，r−对策微生物（指具有生长迅速、C/N 值较低的微生物群组）在群落中占优势地位；随着底物水平的降低，土壤中K-对策微生物（指具有生长缓慢、C/N 值较高的微生物群组）在群落中逐渐占据优势地位。土壤微生物群落组成的改变进一步导致了微生物功能群代谢活性和特征的变化，具体表现为提高了有机物添加处理中土壤细菌群落的代谢潜能，并使细菌在群落水平上的生理剖面特征明显区别于未添加有机物的处理。 研究样地内土壤微生物主要受到底物中的能量（碳）限制，土壤活性有机质库作为可利用性较高的能量和养分的重要来源，对土壤微生物活性和土壤碳周转起着比水分因子更加重要的作用。土壤水分主要影响植物生长和根系活性，并增加了土壤微生物对底物响应的复杂性，但它对地下生物过程的作用程度以底物中能量和养分水平为前提。 利用稳定性13C 同位素示踪技术测量后发现，添加C4-植物凋枯物会加速C3 底物中碳的分解速度。结合有机物添加后土壤有机质库的变化，可以推测植物凋枯物（即能量物质）输入增加会导致土壤原生有机碳的正向激发效应。在此过程中，土壤微生物群落组成及功能群代谢活性的变化起着至关重要的作用。 不同光合途径（C3 和C4）的植物和同一植物不同器官组织（地上部分和根系）的凋枯物添加对地下生物过程，特别是土壤微生物群落代谢功能的影响是不同的。在添加C3-草本凋枯物的处理中，土壤细菌群落主要利用的碳源为氨基酸类化合物；而在添加C4-植物凋枯物的处理中，土壤细菌群落主要利用的碳源为羧酸类化合物。 本研究在野外自然条件下证明了在能量缺乏的中国北方草地生态系统中，土壤有机物输入增加不但不会提高土壤有机碳库的大小，而且可能导致土壤原生有机碳的激发效应。在利用土壤呼吸与环境因子（如温度）的关系进行模拟预测土壤碳排放时，需要考虑不同生态系统底物中的能量和养分水平，以及土壤微生物和植物根系等地下生物过程对底物水平的适应性。

黄土丘陵区退耕地先锋群落演替过程中细根特征的变化

采用空间序列取代时间序列的方法,对退耕时间分别为2、4、6和8年的退耕地群落细根特征进行了分析,以探讨退耕地植被演替过程中细根特征在土壤剖面上的变化及其在演替过程中的作用.结果表明:(1)群落细根根长密度和根面积密度随植被演替显著增加;比根长、比根面积和地下/地上生物量也有增加趋势;细根平均直径随植被演替波动,但有减少趋势;(2)在土壤剖面上根长密度、根面积密度和根系生物量均随土壤深度的增加而降低.其中超过63%的根长、61%的根面积和72%的生物量分布在0～20cm的表层土壤中;(3)根径级统计表明,多数细根直径在0～0.5和0.5～1.0mm之间,这两级细根长度占细根总长度的80%以上;(4)逐步回归分析表明,植被演替过程中细根特征的变化主要与土壤有效氮(第2年)、有效磷(第2～8年)和土壤水分(第8年)的含量有关,且随着植被演替,2～6年退耕地中细根特征与土壤资源正相关,而第8年中二者呈负相关.这可能与植物生长对资源的需求与土壤提供资源的能力之间的平衡有关.研究表明,退耕地植被演替过程对土壤资源有一定影响,尤其是土壤水分含量显著减少;而土壤水分等土壤资源的变化又对群落产生影响,导致茵陈蒿先锋群落向地带性...