Effects of light conditions and plant density on growth and reproductive biology of Cascabela thevetia (L.) Lippold

Cascabela thevetia (L.) Lippold (Apocynaceae) is an invasive woody weed that has formed large infestations at several locations in northern Australia. Understanding the reproductive biology of C. thevetia is vital to its management. This paper reports results of a shade house experiment that determined the effects of light conditions (100% or 30% of natural light) and plant densities (one, two, four or eight plants per plot) on the growth, time to flowering and seed formation, and monthly pod production of two C. thevetia biotypes (peach and yellow). Shaded plants were significantly larger when they reached reproductive maturity than plants grown under natural light. However, plants grown under natural light flowered earlier (268 days compared with 369 days) and produced 488 more pods per pot (a 5-fold increase) over 3 years. The yellow biotype was slightly taller at reproductive maturity but significantly taller and with significantly greater aboveground biomass at the end of the study. Both biotypes flowered at a similar time under natural light and low plant densities but the yellow biotype was quicker to seed (478 versus 498 days), produced significantly more pods (364 versus 203 pods) and more shoot growth (577 g versus 550 g) than the peach biotype over 3 years. Higher densities of C. thevetia tended to significantly reduce the shoot and root growth by 981 g and 714 g per plant across all light conditions and biotypes over 3 years and increase the time taken to flower by 140 days and produce seeds by 184 days. For land managers trying to prevent establishment of C. thevetia or to control seedling regrowth once initial infestations have been treated, this study indicates that young plants have the potential to flower and produce seeds within 268 and 353 days, respectively. However, with plant growth and reproduction most likely to be slower under field conditions, annual surveillance and control activities should be sufficient to find and treat plants before they produce seeds and replenish soil seed banks. The most at-risk part of the landscape may be open areas that receive maximum sunlight, particularly within riparian habitats where plants would consistently have more favourable soil moisture conditions.

Remote sensing of boreal land cover: estimation of forest attributes and extent

Remote sensing provides methods to infer land cover information over large geographical areas at a variety of spatial and temporal resolutions. Land cover is input data for a range of environmental models and information on land cover dynamics is required for monitoring the implications of global change. Such data are also essential in support of environmental management and policymaking. Boreal forests are a key component of the global climate and a major sink of carbon. The northern latitudes are expected to experience a disproportionate and rapid warming, which can have a major impact on vegetation at forest limits. This thesis examines the use of optical remote sensing for estimating aboveground biomass, leaf area index (LAI), tree cover and tree height in the boreal forests and tundra taiga transition zone in Finland. The continuous fields of forest attributes are required, for example, to improve the mapping of forest extent. The thesis focus on studying the feasibility of satellite data at multiple spatial resolutions, assessing the potential of multispectral, -angular and -temporal information, and provides regional evaluation for global land cover data. Preprocessed ASTER, MISR and MODIS products are the principal satellite data. The reference data consist of field measurements, forest inventory data and fine resolution land cover maps. Fine resolution studies demonstrate how statistical relationships between biomass and satellite data are relatively strong in single species and low biomass mountain birch forests in comparison to higher biomass coniferous stands. The combination of forest stand data and fine resolution ASTER images provides a method for biomass estimation using medium resolution MODIS data. The multiangular data improve the accuracy of land cover mapping in the sparsely forested tundra taiga transition zone, particularly in mires. Similarly, multitemporal data improve the accuracy of coarse resolution tree cover estimates in comparison to single date data. Furthermore, the peak of the growing season is not necessarily the optimal time for land cover mapping in the northern boreal regions. The evaluated coarse resolution land cover data sets have considerable shortcomings in northernmost Finland and should be used with caution in similar regions. The quantitative reference data and upscaling methods for integrating multiresolution data are required for calibration of statistical models and evaluation of land cover data sets. The preprocessed image products have potential for wider use as they can considerably reduce the time and effort used for data processing.

Biomass for aviation fuel production in the Fitzroy Basin, Queensland: a preliminary assessment of native and plantation forest potential

SummaryThis scoping study assesses the contribution that woody biomass could make to feedstock supply for an aviation biofuel industry in Queensland. The inland 600?900 mm rainfall zone, including the Fitzroy Basin region, is identified as an area that is particularly worthy of closer study as it has potential for supply of woody biomass from existing native regrowth (brigalow and other species) as well as from new plantings. New analyses carried out for this study of Corymbia citriodora subsp. variegata trials suggest biomass plantings could produce harvestable yield of aboveground dry mass of about 85 t ha?1 over a 10-year rotation at relatively low-rainfall (600?750 mm mean annual precipitation) sites and about 115 t ha?1 at medium-rainfall (750?900 mm) sites. Estimates of productivity for native regrowth suggest potential productivity should be around 40 t ha?1 during the initial decade after clearing when systems are managed for bioenergy rather than grazing. In this paper, potential production systems are described, and sustainability issues are briefly considered. It is concluded that more detailed studies focused particularly on biomass production would be worthwhile, and further research requirements are briefly discussed.

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.

Carbon balance and component CO2 fluxes in boreal Scots pine stands

The forest vegetation takes up atmospheric carbon dioxide (CO2) in photosynthesis. Part of the fixed carbon is released back into the atmosphere during plant respiration but a substantial part is stored as plant biomass, especially in the stems of trees. Carbon also accumulates in the soil as litter and via the roots. CO2 is released into the atmosphere from these carbon stocks in the decomposition of dead biomass. Carbon balance of a forest stand is the difference between the CO2 uptake and CO2 efflux. This study quantifies and analyses the dynamics of carbon balance and component CO2 fluxes in four Southern Finnish Scots pine stands that covered the typical economic rotation time of 80 years. The study was based on direct flux measurements with chambers and eddy covariance (EC), and modelling of component CO2 fluxes. The net CO2 exchange of the stand was partitioned into component fluxes: photosynthesis of trees and ground vegetation, respiration of tree foliage and stems, and CO2 efflux from the soil. The relationships between the component fluxes and the environmental factors (light, temperature, atmospheric CO2, air humidity and soil moisture) were studied with mathematical modelling. The annual CO2 balance varied from a source of about 400 g C/m2 at a recently clearcut site to net CO2 uptake of 200 300 g C/m2 in a middle-aged (40-year-old) and a mature (75-year-old) stand. A 12-year-old sapling site was at the turning point from source to a sink of CO2. In the middle-aged stand, photosynthetic production was dominated by trees. Under closed pine canopies, ground vegetation accounted for 10 20% of stand photosynthesis whereas at the open sites the proportion and also the absolute photosynthesis of ground vegetation was much higher. The aboveground respiration was dominated by tree foliage which accounted for one third of the ecosystem respiration. Rate of wood respiration was in the order of 10% of total ecosystem respiration. CO2 efflux from the soil dominated the ecosystem respiratory fluxes in all phases of stand development. Instantaneous and delayed responses to the environmental driving factors could predict well within-year variability in photosynthetic production: In the short term and during the growing season photosynthesis follows primarily light while the seasonal variation is more strongly connected to temperature. The temperature relationship of the annual cycle of photosynthesis was found to be almost equal in the southern boreal zone and at the timberline in the northern boreal zone. The respiratory fluxes showed instantaneous and seasonal temperature relationships but they could also be connected to photosynthesis at an annual timescale.

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.

Jakojuuriviljelyn vaikutus kasvihuonekurkun (Cucumis sativus L.) kasvuun ja kehitykseen

Kasvit ottavat vettä parhaiten kasteluravinneliuoksesta, jonka ravinnepitoisuus on pieni. Intensiivisessä kasvihuonetuotannossa käytetään silti kastelulannoituksessa usein korkeita ravinnepitoisuuksia ravinnepuutosten ja satotappioiden välttämiseksi. Jakojuuriviljelyssä kasvin juuriston annetaan kasvaa kahteen erilliseen kasvualustaosioon. Tällöin toiselle puolelle annetaan johtokyvyltään väkevää ja toiselle puolelle laimeaa ravinneliuosta. Erityisesti kasvihuonekurkun, joka on herkkä kasvualustan suolaisuuden aiheuttamille vedensaantiongelmille, on todettu hyötyvän tästä tekniikasta, mikä näkyy kasvaneina satoina. Tämän MTT Piikkiössä toteutetun kasvihuonekurkun jakojuuriviljelytutkimuksen tavoitteena oli tarkentaa tekniikkaa erityisesti kasteluliuosten johtokyvyn osalta. Yhtenäisjuuriviljelyn ja perinteisen jakojuuriviljelyn lisäksi kokeessa oli kaksi jakojuuriviljelykäsittelyä, joissa ravinneliuosväkevyyksiä vaihdettiin väliajoin juuriston toimintakyvyn parantamiseksi. Erillisessä osakokeessa tutkittiin erilaisten johtokyky-yhdistelmien vaikutusta kasvihuonekurkun vegetatiiviseen kasvuun maanpäällisten ja -alaisten kasvinosien välillä sekä juurten morfologiaan ja anatomiaan. Tulokset osoittivat, että jakojuuriviljely lisäsi kasvihuonekurkun sadontuottoa jopa 16 %, mutta ei vaikuttanut koko viljelykauden veden tai ravinteiden ottoon. Yhtenäisjuuriviljelyssä muodostui eniten piikkikärkisiä hedelmiä, mikä viittaa vedensaantiongelmiin haihdutustarpeen ollessa suurin. Viljelytekniikalla ei ollut vaikutusta kasvien vegetatiiviseen kasvuun tai kasvuston rakenteeseen. Lehtiruodeista tehdyt nitraatti- ja kaliummittaukset osoittivat, ettei kasteluliuosten ravinnepitoisuuksilla ollut vaikutusta juurten ravinteiden ottoon. Erilaisilla johtokyky-yhdistelmillä oli huomattavampi vaikutus kasvihuonekurkun juurten painoon kuin verson painoon tai varren pituuskasvuun. Lehtiruotianalyysit viittasivat ravinteiden erilaiseen allokointiin eri johtokyky-yhdistelmissä. Korkeiden johtokykyjen aiheuttama osmoottinen stressi johti muutoksiin juurten morfologiassa ja anatomiassa. Tulosten perusteella jakojuuriviljely paransi kehittyvien hedelmien kohdevahvuutta suhteessa muihin kohteisiin vaikuttamatta vegetatiiviseen kasvuun. Kun laimean ja väkevän ravinneliuoksen puolia vaihdettiin, juuristo otti joustavasti vettä ja ravinteita olosuhteiden määräämästä edullisemmasta johtokyvystä, jolloin kasvihuonekurkun viljelyssä saavutettiin merkittävä satoetu. Juuriston jakaminen vaikuttanee kasvien hormoniaineenvaihduntaan ja voi heikentää juuriston kasvua heikentämättä sen toimintakykyä, jolloin yhteyttämistuotteita kohdennetaan tehokkaammin maanpäällisten osien kasvuun.

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.

Local spatial structure of forest biomass and its consequences for remote sensing of carbon stocks

Advances in forest carbon mapping have the potential to greatly reduce uncertainties in the global carbon budget and to facilitate effective emissions mitigation strategies such as REDD+ (Reducing Emissions from Deforestation and Forest Degradation). Though broad-scale mapping is based primarily on remote sensing data, the accuracy of resulting forest carbon stock estimates depends critically on the quality of field measurements and calibration procedures. The mismatch in spatial scales between field inventory plots and larger pixels of current and planned remote sensing products for forest biomass mapping is of particular concern, as it has the potential to introduce errors, especially if forest biomass shows strong local spatial variation. Here, we used 30 large (8-50 ha) globally distributed permanent forest plots to quantify the spatial variability in aboveground biomass density (AGBD in Mgha(-1)) at spatial scales ranging from 5 to 250m (0.025-6.25 ha), and to evaluate the implications of this variability for calibrating remote sensing products using simulated remote sensing footprints. We found that local spatial variability in AGBD is large for standard plot sizes, averaging 46.3% for replicate 0.1 ha subplots within a single large plot, and 16.6% for 1 ha subplots. AGBD showed weak spatial autocorrelation at distances of 20-400 m, with autocorrelation higher in sites with higher topographic variability and statistically significant in half of the sites. We further show that when field calibration plots are smaller than the remote sensing pixels, the high local spatial variability in AGBD leads to a substantial ``dilution'' bias in calibration parameters, a bias that cannot be removed with standard statistical methods. Our results suggest that topography should be explicitly accounted for in future sampling strategies and that much care must be taken in designing calibration schemes if remote sensing of forest carbon is to achieve its promise.

Littoral vegetation of Lake Tohopekaliga: community descriptions prior to a large-scale fisheries habitat-enhancement project

An extreme dry-down and muck-removal project was conducted at Lake Tohopekaliga, Florida, in 2003-2004, to remove dense vegetation from inshore areas and improve habitat degraded by stabilized water levels. Vegetation was monitored from June 2002 to December 2003, to describe the pre-existing communities in terms of composition and distribution along the environmental gradients. Three study areas (Treatment-Selection Sites) were designed to test the efficacy of different treatments in enhancing inshore habitat, and five other study areas (Whole-Lake Monitoring Sites) were designed to monitor the responses of the emergent littoral vegetation as a whole. Five general community types were identified within the study areas by recording aboveground biomasses and stem densities of each species. These communities were distributed along water and soils gradients, with water depth and bulk density explaining most of the variation. The shallowest depths were dominated by a combination of Eleocharis spp., Luziola fluitans, and Panicum repens; while the deeper areas had communities of Nymphaea odorata and Nuphar luteum; Typha spp.; or Paspalidium geminatum and Hydrilla verticillata. Mineralized soils were common in both the shallow and deep-water communities, while the intermediate depths had high percentages of organic material in the soil. These intermediate depths (occurring just above and just below low pool stage) were dominated by Pontederia cordata, the main species targeted by the habitat enhancement project. This emergent community occurred in nearly monocultural bands around the lake (from roughly 60–120 cm in depth at high pool stage) often having more diverse floating mats along the deep-water edge. The organic barrier these mats create is believed to impede access of sport fish to shallow-water spawning areas, while the overall low diversity of the community is evidence of its competitive nature in stabilized waters. With continued monitoring of these study areas long-term effects of the restoration project can be assessed and predictive models may be created to determine the efficacy and legitimacy of such projects in the future.

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。

内蒙古高原成熟和退化羊草草原群落物种功能特性与土壤微生物量C、N、P对氮素添加响应

氮素是大多数陆地生态系统初级生产力的主要限制因子。由于人类的工业和农业生产活动不断加剧，导致全球性氮沉降增加，使大多数生态系统氮素的可获得性增强。从而降低或消除了氮素对生态系统的限制作用，加速了生态系统生物地球化学过程，对物种多样性和生态系统结构与功能产生了显著的影响。但由于成土母质、气候条件、地形地貌、植被组成等的差异，不同生态系统类型对氮素增加的响应也不尽相同。欧洲和北美一些发达国家地区对于草地生态系统对于全球性氮沉降增加响应进行了较全面的研究，对于分布广泛的欧亚大陆草原研究相对不足。 本文研究选择对于欧亚大陆草原较具代表性的成熟羊草草原群落及该群落的退化类型为研究对象，从1999年开始，在这两类群落中选取地形相对平缓均一，植被组成一致的地段设置了施肥小区并进行持续氮素添加实验。本文研究了成熟和退化羊草草原群落物种功能特性与土壤微生物量C、N、P对氮素添加响应。 羊草群落中6种主要植物的地上生物量、种群密度、比叶面积、叶氮和叶绿素含量对于氮素添加响应以及各指标之间相关关系的分析表明：比叶面积、基于质量的叶片含氮量和叶绿素含量、叶绿素a和叶绿素b的比值等叶片水平上物种功能特性间的相互作用，共同影响和决定了种群密度和地上生物量对氮素添加的响应。羊草通过提高比叶面积、叶片叶绿素含量和含氮量、种群密度及个体生物量等多重调节功能对氮素添加做出响应。西伯利亚羽茅主要通过提高比叶面积、单位质量叶片的叶绿素含量和含氮量，以及株丛生物量，使其在群落占据优势。大针茅和冰草在提高比叶面积、叶片叶绿素含量和含氮量的调节能力相对较低，种群密度沿氮素添加梯度显著降低。黄囊苔草只能通过提高叶片叶绿素含量和含氮量对氮素添加做出响应，其叶绿素a与叶绿素b的比值沿氮素添加梯度逐渐降低，种群密度和地上生物量也显著降低。糙隐子草的叶绿素a与叶绿素b比值沿氮素添加梯度显著降低，但由于糙隐子草具有较高的SLA，且对叶绿素、叶片含氮量的调节能力较强，氮素添加处理没有对其种群密度和地上生物量产生显著的影响。上述结果支持Tilman的光资源竞争假说和Knops等的物种替代假说。 成熟和退化羊草群落土壤微生物量、土壤有机碳、全氮、全磷、速效氮、pH以及凋落物碳、氮、磷含量的测定结果表明：（1）成熟羊草群落表层土壤微生物量碳、氮、磷含量均随氮素添加量的增加而降低;退化羊草群落表层土壤微生物量碳、氮、磷含量沿氮素梯度表现出先增加而后降低的趋势;相关分析的结果显示各群落土壤微生物量碳、氮、磷均与土壤pH呈显著的正相关。（2）微生物量碳、氮、磷含量均随土层深度的增加而下将;而对照的微生物量碳、氮、磷含量则与土壤有机质含量呈显著正相关。（3）年度间降水量差异对土壤微生物量碳、氮、磷具有较大影响。综合上述研究结果，我们认为成熟羊草群落土壤微生物生长不受氮素限制，但退化群落不同;氮素添加导致的土壤酸化作用可能是两类群落表层土壤微生物量下降的主要因素，且这种影响主要集中在0-10cm的表层土壤;表层土壤微生物量碳、氮、磷对氮素添加的响应可能还受到其它因子（如生长季降水量）的影响;深层土壤微生物量较低主要是由于土壤有机质含量较低的缘故。

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。

内蒙古典型草原地上净初级生产力估测及模型模拟的研究

地上净初级生产力（ANPP）是陆地生态系统碳循环的重要组成部分，但由于估测ANPP的方法不同使得对ANPP的估测值存在很大的不确定性。本文采用3种方法（群落中所有种群当年最大地上生物量之和（ANPPC1）、当年群落最大地上生物量（ANPPC2）以及每年固定日期（8月30日）的地上生物量（ANPPC3））在种群、功能群和群落水平上分别对连续19年（1980～1998）的内蒙古羊草和大针茅草原生态系统功能（如ANPP、植物多样性和水分利用效率（WUE））的动态变化进行了比较分析，同时探讨了不同估测方法下气候和放牧对ANPP的影响，在此基础上利用DNDC模型进行了ANPP的模拟和敏感性分析研究，主要结果包括： 在羊草和大针茅草原群落中，不同主要植物种群或功能群多年平均地上最大生物量出现的时间不同，而同一植物种群或功能群的年地上最大生物量出现的时间存在年际间的变化。采用当年最大地上生物量和8月30日固定日期的地上生物量作为群落ANPP的这两种方法高估了建群种或禾草功能群在群落中的作用。 羊草草原群落多年平均ANPPC1、ANPPC2和 ANPPC3分别是257.5、190.1和166.0 g.m-2;相应的大针茅草原多年平均ANPPC1、ANPPC2和 ANPPC3分别是180.4、132.8和122.5 g.m-2。就群落生产力而言，后两种常用的方法二和方法三分别低估了草原群落ANPP 14.2%～40.0%和15.5～59.0%。本文研究表明尽管ANPPC1与ANPPC2和ANPPC3之间存在显著的差异，但二者之间存在极显著的相关性:羊草草原的ANPPC1= 59.587+1.061×ANPPC2（r2＝0.865, p＜0.001），ANPPC1= 92.329+1.017×ANPPC3（r2＝0.569, p＜0.001）;大针茅草原的ANPPC1= 32.918+1.114×ANPPC2（r2＝0.814, p＜0.001），ANPPC1= 76.120+0.875×ANPPC3（r2＝0.499, p＝0.001）。 种群、功能群和群落地上净初级生产力与气候因子间的关系因不同的估测方法而异。羊草草原的建群种羊草种群仅ANPPS3与8月和11月份的平均温度间存在显著相关性，而大针茅草原群落的建群种大针茅种群的ANPPS1和ANPPS2与3月份平均最高气温间呈负相关关系。在羊草草原群落，杂类草功能群ANPPF1和ANPPF2与3月份气温呈负相关关系;而灌木半灌木功能群ANPPF1和ANPPF2与3月份降水呈负相关关系。在大针茅草原群落，禾草功能群ANPPF1与5月份最高平均气温呈显著负相关关系。羊草草原群落ANPPC1和ANPPC3分别与11月份最低温度和平均温度存在显著负相关关系;大针茅草原群落ANPPC1与2和6月份月降水量间相关性显著，ANPP2与5月和11月份月平均最低温度呈显著负相关关系，而4～9月份，1和4月份平均最低温度对群落ANPPC3起决定作用。 在羊草和大针茅草原群落，由方法一得到的群落水分利用效率、Shannon植物多样性指数和均匀度指数与方法二或方法三得到的相应的指数间存在显著的差异，方法二和方法三得到的值间差异不显著。群落地上净初级生产力与相应估测方法的植物Shannon多样性指数和均匀度指数之间相关性不显著。 放牧条件下羊草或大针茅草原群落中的建群种羊草和大针茅在群落中的相对地上生物量较围栏内的相应植物种群的降低，而糙隐子草种群在群落中的比例上升。不同的放牧管理条件下，群落中植物种群地上现存量的季节动态发生变化。群落的植物组成及植物种群地上生物量在群落中总生物量的比例发生了明显的变化。利用遥感来估测地上净初级生产力时，分别低估了羊草和大针茅草原ANPP 52％和27％。 DNDC模型可以很好地模拟内蒙古典型草原生态系统的地上生物量，通过敏感性分析表明降水是草原植物生长的主要限制因子，在降水量增加或降低至日降水的30%时，模拟的地上生物量显著地高于或低于实测地上生物量值。在多年平均降水量为347mm的情况下，随着土壤粘粒含量的增加，地上生物量逐渐降低，与北美草原一致。