Canopy processes, fluxes and microclimate in a pine forest

Interaction between forests and the atmosphere occurs by radiative and turbulent transport. The fluxes of energy and mass between surface and the atmosphere directly influence the properties of the lower atmosphere and in longer time scales the global climate. Boreal forest ecosystems are central in the global climate system, and its responses to human activities, because they are significant sources and sinks of greenhouse gases and of aerosol particles. The aim of the present work was to improve our understanding on the existing interplay between biologically active canopy, microenvironment and turbulent flow and quantify. In specific, the aim was to quantify the contribution of different canopy layers to whole forest fluxes. For this purpose, long-term micrometeorological and ecological measurements made in a Scots pine (Pinus sylvestris) forest at SMEAR II research station in Southern Finland were used. The properties of turbulent flow are strongly modified by the interaction between the canopy elements: momentum is efficiently absorbed in the upper layers of the canopy, mean wind speed and turbulence intensities decrease rapidly towards the forest floor and power spectra is modulated by spectral short-cut . In the relative open forest, diabatic stability above the canopy explained much of the changes in velocity statistics within the canopy except in strongly stable stratification. Large eddies, ranging from tens to hundred meters in size, were responsible for the major fraction of turbulent transport between a forest and the atmosphere. Because of this, the eddy-covariance (EC) method proved to be successful for measuring energy and mass exchange inside a forest canopy with exception of strongly stable conditions. Vertical variations of within canopy microclimate, light attenuation in particular, affect strongly the assimilation and transpiration rates. According to model simulations, assimilation rate decreases with height more rapidly than stomatal conductance (gs) and transpiration and, consequently, the vertical source-sink distributions for carbon dioxide (CO2) and water vapor (H2O) diverge. Upscaling from a shoot scale to canopy scale was found to be sensitive to chosen stomatal control description. The upscaled canopy level CO2 fluxes can vary as much as 15 % and H2O fluxes 30 % even if the gs models are calibrated against same leaf-level dataset. A pine forest has distinct overstory and understory layers, which both contribute significantly to canopy scale fluxes. The forest floor vegetation and soil accounted between 18 and 25 % of evapotranspiration and between 10 and 20 % of sensible heat exchange. Forest floor was also an important deposition surface for aerosol particles; between 10 and 35 % of dry deposition of particles within size range 10 30 nm occurred there. Because of the northern latitudes, seasonal cycle of climatic factors strongly influence the surface fluxes. Besides the seasonal constraints, partitioning of available energy to sensible and latent heat depends, through stomatal control, on the physiological state of the vegetation. In spring, available energy is consumed mainly as sensible heat and latent heat flux peaked about two months later, in July August. On the other hand, annual evapotranspiration remains rather stable over range of environmental conditions and thus any increase of accumulated radiation affects primarily the sensible heat exchange. Finally, autumn temperature had strong effect on ecosystem respiration but its influence on photosynthetic CO2 uptake was restricted by low radiation levels. Therefore, the projected autumn warming in the coming decades will presumably reduce the positive effects of earlier spring recovery in terms of carbon uptake potential of boreal forests.

AC transport at holographic quantum hall transitions

We compute AC electrical transport at quantum Hall critical points, as modeled by intersecting branes and gauge/gravity duality. We compare our results with a previous field theory computation by Sachdev, and find unexpectedly good agreement. We also give general results for DC Hall and longitudinal conductivities valid for a wide class of quantum Hall transitions, as well as (semi)analytical results for AC quantities in special limits. Our results exhibit a surprising degree of universality; for example, we find that the high frequency behavior, including subleading behavior, is identical for our entire class of theories.

TOTEM early measurements

The status of the TOTEM experiment is described as well as the prospects for the measurements in the early LHC runs. The primary goal of TOTEM is the measurement of the total p-p cross section, using a method independent of the luminosity. A final accuracy of 1% is ex- pected with dedicated β∗ = 1540 m runs, while at the beginning a 5% resolution is achievable with a β∗ = 90 m optics. Accordingly to the running scenarios TOTEM will be able to measure the elastic scattering in a wide range of t and to study the cross-sections and the topologies of diffractive events. In a later stage, physics studies will be extended to low-x and forward physics collaborating with CMS as a whole experimental apparatus.

Measurements of the top-quark mass using charged particle tracking

We present three measurements of the top-quark mass in the lepton plus jets channel with approximately 1.9 fb-1 of integrated luminosity collected with the CDF II detector using quantities with minimal dependence on the jet energy scale. One measurement exploits the transverse decay length of b-tagged jets to determine a top-quark mass of 166.9+9.5-8.5 (stat) +/- 2.9 (syst) GeV/c2, and another the transverse momentum of electrons and muons from W-boson decays to determine a top-quark mass of 173.5+8.8-8.9 (stat) +/- 3.8 (syst) GeV/c2. These quantities are combined in a third, simultaneous mass measurement to determine a top-quark mass of 170.7 +/- 6.3 (stat) +/- 2.6 (syst) GeV/c2.

Inhomogeneity of asteroid 2008 TC3 (Almahata Sitta meteorites) revealed through magnetic susceptibility measurements

Magnetic susceptibility measurements were performed on freshly fallen Almahata Sitta meteorites. Most recovered samples are polymict ureilites. Those found in the first four months since impact, before the meteorites were exposed to rain, have a magnetic susceptibility in the narrow range of 4.92 ± 0.08 log 10-9 Am2/kg close to the range of other ureilite falls 4.95 ± 0.14 log 10-9 Am2/kg reported by Rochette et al. (2009). The Almahata Sitta samples collected one year after the fall have similar values (4.90 ± 0.06 log 10-9 Am2/kg), revealing that the effect of one-year of terrestrial weathering was not severe yet. However, our reported values are higher than derived from polymict (brecciated) ureilites 4.38 ± 0.47 log 10-9 Am2/kg (Rochette et al. 2009) containing both falls and finds confirming that these are significantly weathered. Additionally other fresh-looking meteorites of non-ureilitic compositions were collected in the Almahata Sitta strewn field. Magnetic susceptibility measurements proved to be a convenient non-destructive method for identifying non-ureilitic meteorites among those collected in the Almahata Sitta strewn field, even among fully crusted. Three such meteorites, no. 16, 25, and 41, were analyzed and their composition determined as EH6, H5 and EL6 respectively (Zolensky et al., 2010). A high scatter of magnetic susceptibility values among small (< 5 g) samples revealed high inhomogeneity within the 2008 TC3 material at scales below 1-2 cm.

USING FLUORESCENT AND NON-FLUORESCENT STEROLS TO STUDY RAFTS AND INTRACELLULAR CHOLESTEROL TRANSPORT IN MAMMALIAN CELLS

Cholesterol is an essential component in the membranes of most eukaryotic cells, in which it mediates many functions including membrane fluidity, permeability and the formation of ordered membrane domains. In this work a fluorescent and a non-fluorescent cholesterol analog were characterized as tools to study cholesterol. Next, these analogs were used to study two specific cell biological processes that involve cholesterol, i.e. the structure and function of ordered membrane domains/rafts and intracellular cholesterol transport. The most common method for studying ordered membrane domains is by disrupting them by cholesterol depletion. Because cholesterol depletion affects many cellular functions besides those mediated by membrane domains, this procedure is highly unspecific. The cellular exchange of cholesterol by desmosterol as a tool to study ordered membrane domains was characterized. It turned out that the ability of desmosterol to form and stabilize membrane domains in vitro was weaker compared to cholesterol. This result was reinforced by atomistic scale simulations that indicated that desmosterol has a lower ordering effect on phospholipid acyl chains. Three procedures were established for exchanging cellular cholesterol by desmosterol. In cells in which desmosterol was the main sterol, insulin signaling was attenuated. The results suggest that this was caused by desmosterol destabilizing membrane rafts. Contrary to its effect on ordered membrane domains it was found that replacing cholesterol by desmosterol does not change cell growth/viability, subcellular sterol distribution, Golgi integrity, secretory pathway, phospholipid composition and membrane fluidity. Together these results suggest that exchanging cellular cholesterol by desmosterol provides a selective tool for perturbing rafts. Next, the importance of cholesterol for the structure and function of caveolae was analyzed by exchanging the cellular cholesterol by desmosterol. The sterol exchange reduced the stability of caveolae as determined by detergent resistance of caveolin-1 and heat resistance of caveolin-1 oligomers. Also the sterol exchange led to aberrations in the caveolar structure; the morphology of caveolae was altered and there was a larger variation in the amount of caveolin-1 molecules per caveola. These results demonstrate that cholesterol is important for caveolar stability and structural homogeneity. In the second part of this work a fluorescent cholesterol analog was characterized as a tool to study cholesterol transport. Tight control of the intracellular cholesterol distribution is essential for many cellular processes. An important mechanism by which cells regulate their membrane cholesterol content is by cholesterol traffic, mostly from the plasma membrane to lipid droplets. The fluorescent sterol probe BODIPY-cholesterol was characterized as a tool to analyze cholesterol transport between the plasma membrane, the endoplasmic reticulum (ER) and lipid droplets. The behavior of BODIPY-cholesterol was compared to that of natural sterols, using both biochemical and live-cell microcopy assays. The results show that the transport kinetics of BODIPY-cholesterol between the plasma membrane, the ER and lipid droplets is similar to that of unesterified cholesterol. Next, BODIPY-cholesterol was utilized to analyze the importance of oxysterol binding protein related proteins (ORPs) for cholesterol transport between the plasma membrane, the ER, and lipid droplets in mammalian cells. By overexpressing all human ORPs it turned out that especially ORP1S and ORP2 enhanced sterol transport from the plasma membrane to lipid droplets. Our results suggest that the increased sterol transport takes place between the plasma membrane and ER and not between the ER and lipid droplets. Simultaneous knockdown of ORP1S and ORP2 resulted in a moderate but significant inhibition of sterol traffic from the plasma membrane to ER and lipid droplets, suggesting a physiological role for these ORPs in this process. The two phenylalanines in an acidic tract (FFAT) motif in ORPs, which mediates interaction with vesicle associated membrane protein associated proteins (VAPs) in the ER, was not necessary for mediating sterol transport. However, VAP silencing slowed down sterol transport, most likely by destabilizing ORPs containing a FFAT motif.

Disjunct eddy covariance measurements of volatile organic compound fluxes using proton transfer reaction mass spectrometry

Volatile organic compounds (VOCs) are emitted into the atmosphere from natural and anthropogenic sources, vegetation being the dominant source on a global scale. Some of these reactive compounds are deemed major contributors or inhibitors to aerosol particle formation and growth, thus making VOC measurements essential for current climate change research. This thesis discusses ecosystem scale VOC fluxes measured above a boreal Scots pine dominated forest in southern Finland. The flux measurements were performed using the micrometeorological disjunct eddy covariance (DEC) method combined with proton transfer reaction mass spectrometry (PTR-MS), which is an online technique for measuring VOC concentrations. The measurement, calibration, and calculation procedures developed in this work proved to be well suited to long-term VOC concentration and flux measurements with PTR-MS. A new averaging approach based on running averaged covariance functions improved the determination of the lag time between wind and concentration measurements, which is a common challenge in DEC when measuring fluxes near the detection limit. The ecosystem scale emissions of methanol, acetaldehyde, and acetone were substantial. These three oxygenated VOCs made up about half of the total emissions, with the rest comprised of monoterpenes. Contrary to the traditional assumption that monoterpene emissions from Scots pine originate mainly as evaporation from specialized storage pools, the DEC measurements indicated a significant contribution from de novo biosynthesis to the ecosystem scale monoterpene emissions. This thesis offers practical guidelines for long-term DEC measurements with PTR-MS. In particular, the new averaging approach to the lag time determination seems useful in the automation of DEC flux calculations. Seasonal variation in the monoterpene biosynthesis and the detailed structure of a revised hybrid algorithm, describing both de novo and pool emissions, should be determined in further studies to improve biological realism in the modelling of monoterpene emissions from Scots pine forests. The increasing number of DEC measurements of oxygenated VOCs will probably enable better estimates of the role of these compounds in plant physiology and tropospheric chemistry. Keywords: disjunct eddy covariance, lag time determination, long-term flux measurements, proton transfer reaction mass spectrometry, Scots pine forests, volatile organic compounds

Effect of soil tillage on erosion and nutrient transport in plough layer runoff

Yhteenveto: Viljelymenetelmien vaikutus eroosioon ja ravinteiden huuhtoutumiseen

Assessment of insect occurrence in boreal forests based on satellite imagery and field measurements.

The presence/absence data of twenty-seven forest insect taxa (e.g. Retinia resinella, Formica spp., Pissodes spp., several scolytids) and recorded environmental variation were used to investigate the applicability of modelling insect occurrence based on satellite imagery. The sampling was based on 1800 sample plots (25 m by 25 m) placed along the sides of 30 equilateral triangles (side 1 km) in a fragmented forest area (approximately 100 km2) in Evo, S Finland. The triangles were overlaid on land use maps interpreted from satellite images (Landsat TM 30 m multispectral scanner imagery 1991) and digitized geological maps. Insect occurrence was explained using either environmental variables measured in the field or those interpreted from the land use and geological maps. The fit of logistic regression models varied between species, possibly because some species may be associated with the characteristics of single trees while other species with stand characteristics. The occurrence of certain insect species at least, especially those associated with Scots pine, could be relatively accurately assessed indirectly on the basis of satellite imagery and geological maps. Models based on both remotely sensed and geological data better predicted the distribution of forest insects except in the case of Xylechinus pilosus, Dryocoetes sp. and Trypodendron lineatum, where the differences were relatively small in favour of the models based on field measurements. The number of species was related to habitat compartment size and distance from the habitat edge calculated from the land use maps, but logistic regressions suggested that other environmental variables in general masked the effect of these variables in species occurrence at the present scale.

Functional expression and subcellular localization of the Cl- cotransporters KCC2 and NKCC1 in rodent hippocampal and neocortical neurons

The work presented here has focused on the role of cation-chloride cotransporters (CCCs) in (1) the regulation of intracellular chloride concentration within postsynaptic neurons and (2) on the consequent effects on the actions of the neurotransmitter gamma-aminobutyric acid (GABA) mediated by GABAA receptors (GABAARs) during development and in pathophysiological conditions such as epilepsy. In addition, (3) we found that a member of the CCC family, the K-Cl cotransporter isoform 2 (KCC2), has a structural role in the development of dendritic spines during the differentiation of pyramidal neurons. Despite the large number of publications dedicated to regulation of intracellular Cl-, our understanding of the underlying mechanisms is not complete. Experiments on GABA actions under resting steady-state have shown that the effect of GABA shifts from depolarizing to hyperpolarizing during maturation of cortical neurons. However, it remains unclear, whether conclusions from these steady-state measurements can be extrapolated to the highly dynamic situation within an intact and active neuronal network. Indeed, GABAergic signaling in active neuronal networks results in a continuous Cl- load, which must be constantly removed by efficient Cl- extrusion mechanisms. Therefore, it seems plausible to suggest that key parameters are the efficacy and subcellular distribution of Cl- transporters rather than the polarity of steady-state GABA actions. A further related question is: what are the mechanisms of Cl- regulation and homeostasis during pathophysiological conditions such as epilepsy in adults and neonates? Here I present results that were obtained by means of a newly developed method of measurements of the efficacy of a K-Cl cotransport. In Study I, the developmental profile of KCC2 functionality during development was analyzed both in dissociated neuronal cultures and in acute hippocampal slices. A novel method of photolysis of caged GABA in combination with Cl- loading to the somata was used in this study to assess the extrusion efficacy of KCC2. We demonstrated that these two preparations exhibit a different temporal profile of functional KCC2 upregulation. In Study II, we reported an observation of highly distorted dendritic spines in neurons cultured from KCC2-/- embryos. During their development in the culture dish, KCC2-lacking neurons failed to develop mature, mushroom-shaped dendritic spines but instead maintained an immature phenotype of long, branching and extremely motile protrusions. It was shown that the role of KCC2 in spine maturation is not based on its transport activity, but is mediated by interactions with cytoskeletal proteins. Another important player in Cl- regulation, NKCC1 and its role in the induction and maintenance of native Cl- gradients between the axon initial segment (AIS) and soma was the subject of Study III. There we demonstrated that this transporter mediates accumulation of Cl- in the axon initial segment of neocortical and hippocampal principal neurons. The results suggest that the reversal potential of the GABAA response triggered by distinct populations of interneurons show large subcellular variations. Finally, a novel mechanism of fast post-translational upregulation of the membrane-inserted, functionally active KCC2 pool during in-vivo neonatal seizures and epileptiform-like activity in vitro was identified and characterized in Study IV. The seizure-induced KCC2 upregulation may act as an intrinsic antiepileptogenic mechanism.

Production process of Scots pine.

The accompanying collective research report is the result of the research project in 1986­90 between The Finnish Academy and the former Soviet Academy of Sciences. The project was organized around common field work in Finland and in the former Soviet Union and theoretical analyses of tree growth determining processes. Based on theoretical analyses, dynamic stand growth models were made and their parameters were determined utilizing the field results. Annual cycle affects the tree growth. Our theoretical approach was based on adaptation to local climate conditions from Lapland to South Russia. The initiation of growth was described as a simple low and high temperature accumulation driven model. Linking the theoretical model with long term temperature data allowed us to analyze what type of temperature response produced favorable outcome in different climates. Initiation of growth consumes the carbohydrate reserves in plants. We measured the dynamics of insoluble and soluble sugars in the very northern and Karelian conditions. Clear cyclical pattern was observed but the differences between locations were surprisingly small. Analysis of field measurements of CO2 exchange showed that irradiance is the dominating factor causing variation in photosynthetic rate in natural conditions during summer. The effect of other factors is so small that they can be omitted without any considerable loss of accuracy. A special experiment carried out in Hyytiälä showed that the needle living space, defined as the ratio between the shoot cylindric volume and needle surface area, correlates with the shoot photosynthesis. The penetration of irradiance into Scots pine canopy is a complicated phenomenon because of the movement of the sun on the sky and the complicated structure of branches and needles. A moderately simple but balanced forest radiation regime submodel was constructed. It consists of the tree crown and forest structure, the gap probability calculation and the consideration of spatial and temporal variation of radiation inside the forest. The common field excursions in different geographical regions resulted in a lot of experimental data of regularities of woody structures. The water transport seems to be a good common factor to analyse these properties of tree structure. There are evident regressions between cross-sectional areas measured at different locations along the water pathway from fine roots to needles. The observed regressions have clear geographical trends. For example, the same cross-sectional area can support three times higher needle mass in South Russia than in Lapland. Geographical trends can also be seen in shoot and needle structure. Analysis of data published by several Russian authors show, that one ton of needles transpire 42 ton of water a year. This annual amount of transpiration seems to be independent of geographical location, year and site conditions. The produced theoretical and experimental material is utilised in the development of stand growth model that describes the growth and development of Scots pine stands in Finland and the former Soviet Union. The core of the model is carbon and nutrient balances. This means that carbon obtained in photosynthesis is consumed for growth and maintenance and nutrients are taken according to the metabolic needs. The annual photosynthetic production by trees in the stand is determined as a function of irradiance and shading during the active period. The utilisation of the annual photosynthetic production to the growth of different components of trees is based on structural regularities. Since the fundamental metabolic processes are the same in all locations the same growth model structure can be applied in the large range of Scots pine. The annual photosynthetic production and structural regularities determining the allocation of resources have geographical features. The common field measurements enable the application of the model to the analysis of growth and development of stands growing on the five locations of experiments. The model enables the analysis of geographical differences in the growth of Scots pine. For example, the annual photosynthetic production of a 100-year-old stand at Voronez is 3.5 times higher than in Lapland. The share consumed to needle growth (30 %) and to growth of branches (5 %) seems to be the same in all locations. In contrast, the share of fine roots is decreasing when moving from north to south. It is 20 % in Lapland, 15 % in Hyytiälä Central Finland and Kentjärvi Karelia and 15 % in Voronez South Russia. The stem masses (115­113 ton/ha) are rather similar in Hyytiälä, Kentjärvi and Voronez, but rather low (50 ton/ha) in Lapland. In Voronez the height of the trees reach 29 m being in Hyytiälä and Kentjärvi 22 m and in Lapland only 14 m. The present approach enables utilization of structural and functional knowledge, gained in places of intensive research, in the analysis of growth and development of any stand. This opens new possibilities for growth research and also for applications in forestry practice.

Use of wetland buffer areas to reduce nitrogen transport from forested catchments: Retention capacity, emissions of N2O and CH4 and vegetation composition dynamics

The use of buffer areas in forested catchments has been actively researched during the last 15 years; but until now, the research has mainly concentrated on the reduction of sediment and phosphorus loads, instead of nitrogen (N). The aim of this thesis was to examine the use of wetland buffer areas to reduce the nitrogen transport in forested catchments and to investigate the environmental impacts involved in their use. Besides the retention capacity, particular attention was paid to the main factors contributing to the N retention, the potential for increased N2O emissions after large N loading, the effects of peatland restoration for use as buffer areas on CH4 emissions, as well as the vegetation composition dynamics induced by the use of peatlands as buffer areas. To study the capacity of buffer areas to reduce N transport in forested catchments, we first used large artificial loadings of N, and then studied the capacity of buffer areas to reduce ammonium (NH4-N) export originating from ditch network maintenance areas in forested catchments. The potential for increased N2O emissions were studied using the closed chamber technique and a large artificial N loading at five buffer areas. Sampling for CH4 emissions and methane-cycling microbial populations were done on three restored buffer areas and on three buffers constructed on natural peatlands. Vegetation composition dynamics was studied at three buffer areas between 1996 and 2009. Wetland buffer areas were efficient in retaining inorganic N from inflow. The key factors contributing to the retention were the size and the length of the buffer, the hydrological loading and the rate of nutrient loading. Our results show that although the N2O emissions may increase temporarily to very high levels after a large N loading into the buffer area, the buffer areas in forested catchments should be viewed as insignificant sources of N2O. CH4 fluxes were substantially higher from buffers constructed on natural peatlands than from the restored buffer areas, probably because of the slow recovery of methanogens after restoration. The use of peatlands as buffer areas was followed by clear changes in plant species composition and the largest changes occurred in the upstream parts of the buffer areas and the wet lawn-level surfaces, where the contact between the vegetation and the through-flow waters was closer than for the downstream parts and dry hummock sites. The changes in the plant species composition may be an undesired phenomenon especially in the case of the mires representing endangered mire site types, and therefore the construction of new buffer areas should be primarily directed into drained peatland areas.

Field intercomparison of four methane gas analysers suitable for eddy covariance flux measurements

Eddy covariance (EC)-flux measurement technique is based on measurement of turbulent motions of air with accurate and fast measurement devices. For instance, in order to measure methane flux a fast methane gas analyser is needed which measures methane concentration at least ten times in a second in addition to a sonic anemometer, which measures the three wind components with the same sampling interval. Previously measurement of methane flux was almost impossible to carry out with EC-technique due to lack of fast enough gas analysers. However during the last decade new instruments have been developed and thus methane EC-flux measurements have become more common. Performance of four methane gas analysers suitable for eddy covariance measurements are assessed in this thesis. The assessment and comparison was performed by analysing EC-data obtained during summer 2010 (1.4.-26.10.) at Siikaneva fen. The four participating methane gas analysers are TGA-100A (Campbell Scientific Inc., USA), RMT-200 (Los Gatos Research, USA), G1301-f (Picarro Inc., USA) and Prototype-7700 (LI-COR Biosciences, USA). RMT-200 functioned most reliably throughout the measurement campaign and the corresponding methane flux data had the smallest random error. In addition, methane fluxes calculated from data obtained from G1301-f and RMT-200 agree remarkably well throughout the measurement campaign. The calculated cospectra and power spectra agree well with corresponding temperature spectra. Prototype-7700 functioned only slightly over one month in the beginning of the measurement campaign and thus its accuracy and long-term performance is difficult to assess.