93 search hits
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Warten auf die Transkription: Die humanen Elongationsfaktoren NELF und DSIF
(2009)
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Sabine Wenzel
- Die humanen Elongationsfaktoren negative elongation factor (NELF) und 5,6-dichloro-1-beta-D-ribofuranosylbenzimidazole (DRB) sensitivity inducing factor (DSIF) sind Schlüsselfaktoren beim promotorproximalen Arretieren in der frühen Elongationsphase der Transkription. Dabei interagieren beide Faktoren direkt mit RNA-Polymerase II. NELF bindet über die RNA recognition motif (RRM)-Domäne der Untereinheit E zusätzlich an die naszierende RNA. Eine derartige Transkriptionsblockade tritt auch während der Transkription des Genoms des humanen Immunschwächevirus 1 (HIV-1) Provirus auf. NELF bindet hierbei an die Haarnadel-Struktur der naszierenden RNA (HIV-1 TAR RNA). Ein Ziel dieser Arbeit war die strukturelle Analyse der RRM-Domäne von NELF E sowie die Untersuchung einer möglichen sequenzspezifischen Bindung von NELF E RRM an HIV-1 TAR RNA. Außerdem galt es, die Struktur der kleinen Untereinheit von DSIF, hSpt4, zu bestimmen und ihre Interaktion mit der NGN-Domäne der großen DSIF-Untereinheit hSpt5 näher zu charakterisieren. Die Struktur der RRM-Domäne von NELF E wurde mittels magnetischer Kernresonanz (NMR) bestimmt (PDB: 2BZ2) und besteht aus einem viersträngigen Faltblatt mit zwei alpha-Helices, die gegen eine Seite des Faltblattes packen. An der RNA-Bindung sind die zentralen beta-Stränge mit den beiden aromatischen Resten Tyr43 und Phe77 aus dem Konsensussequenzmotiv ribonucleoprotein domain 2 (RNP2) bzw. RNP1 involviert. Der in der freien Form unstrukturierte C-Terminus der RRM bildet in der gebundenen Form (PDB: 2JX2) eine 3-10-Helix aus, die höchstwahrscheinlich direkt an der RNA-Bindung beteiligt ist. RRM-Domänen gelten als klassische einzelstrangbindende Domänen. Die prognostizierte Bindestelle von NELF E ist jedoch die doppelsträngige HIV-1 TAR RNA Stammregion. Die durchgeführten 15N-NMR-Titrationsexperimente und Fluoreszenzgleichgewichtstitrationen mit die HIV-1 TAR RNA umfassenden RNA Oligonukleotiden zeigten, dass NELF E RRM präferentiell einzelsträngige RNA bindet. Die ermittelten KD-Werte lagen dabei alle im niederen mikromolekularen Bereich. NELF E RRM zeigt somit im Vergleich zu anderen RRM-Domänen eine schwächere Affinität für RNA. Eine sequenzspezifische Bindung an die HIV-1 TAR RNA konnte, auch in Versuchen mit der gesamten NELF E Untereinheit, nicht beobachtet werden. Vieles deutet somit darauf hin, dass RNA-Bindung durch NELF E sequenzunabhängig stattfindet. Der Zeitpunkt der RNA-Bindung während der Elongation der Transkription wird womöglich durch andere Faktoren bzw. Ereignisse gesteuert. NELF übt seine inhibitorische Wirkung nur im Zusammenspiel mit dem Elongationsfaktor DSIF aus, über den es nur wenig strukturelle Informationen gibt. Die Koexpression von hSpt4 mit der NusG N-terminalen Homologie Domäne von hSpt5 (hSpt5-NGN) als dessen Interaktionspartner ermöglichte die Bestimmung der Kristallstruktur von hSpt4/hSpt5-NGN bis zu einer Auflösung von 1.55 Å (PDB: 3H7H). hSpt5-NGN besteht aus einem zentralen, viersträngigen Faltblatt und aus einzelnen Helices, die von beiden Seiten gegen das Faltblatt packen. hSpt4 verfügt über ein zwei- und ein dreisträngiges Faltblatt, die senkrecht zueinander angeordnet sind und sich auf der Interaktionsfläche zugewandten Seite befinden. Zusätzliche helikale Elemente werden durch den ausgebildeten Zinkfinger fixiert. Die Komplexstruktur ist von einem zentralen sechssträngigen intermolekularen beta-Faltblatt geprägt. hSpt4/hSpt5-NGN zeigt sehr große Strukturähnlichkeit zur kürzlich publizierten homologen Komplexstruktur von Spt4/Spt5-NGN aus S. cerevisiae. Ein für die Bindung essentieller Glutamatrest (E338) von Spt5-NGN ist auch in hSpt5-NGN konserviert (E228). Der Komplex hSpt4/hSpt5-NGN(E228Q) konnte wegen der Unlöslichkeit von hSpt5-NGN(E228Q) nicht gereinigt werden. Dies deutet an, dass die Interaktion zwischen hSpt4 und hSpt5-NGN(E228Q) gestört ist. Somit ist E228 von hSpt5-NGN höchstwahrscheinlich ebenso essentiell für die Wechselwirkung mit hSpt4 wie im homologen Hefekomplex. hSpt5-NGN hat auch strukturelle Ähnlichkeiten zur N-terminalen Domäne (NTD) des bakteriellen Transkriptionsfaktors NusG. E. coli (Ec)NusG-NTD verfügt an der Position des Glutamatrestes über einen Glutaminrest (Q72). hSpt4-Bindung an EcNusG-NTD bzw. EcNusG-NTD(Q72E) wurde jedoch nicht beobachtet. Ein möglicher Grund dafür ist die unterschiedliche Ladungsverteilung an der hSpt4-Bindungsfläche. hSpt4 besitzt hingegen Ähnlichkeiten zum archaealen Transkriptionsfaktor RpoE‘‘. Überlagerung der Strukturen von hSpt4 und RpoE‘‘ aus P. furiosus (PDB: 1RYQ) zeigten, dass beide den Zinkfinger und die senkrecht zueinander angeordneten Faltblätter gemein haben. Strukturbestimmung sowie funktionelle Analyse von NELF E RRM und hSpt4/hSpt5-NGN repräsentieren somit einen weiteren Schritt auf dem Weg zur Entschlüsselung der komplexen zellulären Vorgänge während der eukaryontischen Transkription.
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Vegetative Vermehrung einheimischer Baumarten in Südecuador - Physiologische Grundlagen und deren Umsetzung
(2009)
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Sina Heppner
- Parallel zur fortschreitenden Vernichtung der megadiversen Bergwälder Ecuadors entstehen immer mehr Plantagen mit exotischen Pinus- und Eucalyptus-Arten. Eine ökologisch sinnvolle Wiederaufforstung kann jedoch nur unter Verwendung indigener Arten erreicht werden. Da die Samenernte problematisch ist, müsste das Pflanzmaterial vor allem durch vegetative Vermehrung erzeugt werden. Bisher fehlt es jedoch noch an Grundlagenwissen und effizienten Methoden zur Klonierung einheimischer Baumarten. • In der vorliegenden Arbeit wurde das vegetative Vermehrungspotential sowie der Kohlenhydrat- und Phytohormongehalt von sechs – in den Bergwäldern Südecuadors natürlich vorkommenden – Baumarten untersucht: Clethra revoluta (Clethraceae), Heliocarpus americanus (Tiliaceae), Isertia laevis (Rubiaceae), Myrica pubescens (Myriaceae), Piptocoma discolor (Asteraceae) und Tabebuia chrysantha (Bignoniaceae). • Trotz vielfältiger Variation der Anzuchtparameter über 1 ½ Jahre bewurzelten sich Sprossstecklinge adulter Bäume nicht. Die Pflanzen vertrockneten rasch. Messungen der stomatären Leitfähigkeit zeigen eine ungenügende Regulation der Stomata verbunden mit hohen Transpirationsverlusten. Die Gewinnung bewurzelter Sprossstecklinge von adulten Bäumen durch Abmoosen war zwar erfolgreich, aber bei der anschließenden Kultur in Erde vertrockneten auch diese Pflanzen. • Die Kohlenhydratmessungen ergaben bei mehreren Arten jährliche Schwankungen im Gehalt von Stärke und löslichen Zuckern, insbesondere der Raffinosefamilie (bei Tabebuia) bzw. erhöhte Inosit-Gehalte während der trockneren Monate. Zwar zeigte der Gesamtkohlenhydratgehalt in den Blättern deutliche saisonale Schwankungen (bei Heliocarpus-, Myrica- und Tabebuia-Pflanzen), doch ließen sich in den Zweigen generell keine ausgeprägten Jahresgänge nachweisen. Während Heliocarpus- und Tabebuia-Bäume Reservekohlenhydrate mit einem Gehalt von 6 bzw. 9,5 % (besonders in Form von Stärke sowie als Glukose, Fruktose und Saccharose) einlagerten, könnte der ganzjährig geringe Kohlenhydratvorrat von Clethra- und Piptocoma-Zweigen einer erfolgreichen Adventivbewurzelung abträglich gewesen sein. • Die endogenen Cytokinin- (Initiation von Wurzelprimordien) und Auxingehalte (Wurzelwachstum) gelten als besonders wichtig für die Stecklingsbewurzelung. Der gemessene Phytohormonstatus fluktuierte jahreszeiten- und artabhängig. Ein für die Bewurzelung günstiger Hormonstatus, also ein hoher Auxin : Cytokinin-Quotient, fand sich bei Heliocarpus und Tabebuia in den niederschlagsreichsten Monaten. Clethra und Myrica enthielten hingegen fast kein Auxin, aber relativ viel Cytokinin. Bei diesen Arten scheint eine Bewurzelung deshalb besonders schwierig zu sein. • Da es nicht möglich ist, in den Hormonstatus adulter Bäume einzugreifen, ergab sich als Konsequenz die Strategie, Jungpflanzen als Donor-Pflanzen zu verwenden, deren Hormon- und Reservestoffstatus durch Stressbehandlung entsprechend verändert werden könnte. Insbesondere galt es, den Cytokininspiegel endogen abzusenken. Heliocarpus und Tabebuia wurden für diese Versuche ausgewählt und Wassermangel, Nährstoffmangel, Einengung des Wurzelraums und Kombinationen dieser Stressoren ausgesetzt. Zusätzlich wurden Wildlinge verwendet. • Diese Strategie erwies sich als erfolgreich, denn es bewurzelten sich über alle Behandlungen gemittelt 42 % der Heliocarpus- und 15 % der Tabebuia-Stecklinge. Am erfolgreichsten waren Stecklinge von dürregestressten Heliocarpus-Bäumchen mit 70 % Bewurzelung und von unter Nährstoffmangel gezogenen Tabebuia-Pflanzen (25 %). Bei dieser Art bewurzelten sich ebenso Wildlinge sehr effektiv (45 %). Noch bessere Ergebnisse wurden mit dem Abmoosen bei beiden Arten erzielt. • Die Vorbehandlungen führten bei Heliocarpus und Tabebuia zu einem Anstieg des Kohlenhydratgehalts (ausgenommen bei dürregestressten Tabebuia-Pflanzen). Es ergab sich allerdings keine statistisch signifikante Korrelation des Kohlenhydratgehalts der verschieden gestressten Donor-Pflanzen mit ihrer Bewurzelungsrate. • Wie angenommen, führte die Stressbehandlung der Donor-Pflanzen beider Arten zu einem geringeren Cytokininspiegel, wobei in Tabebuia die Bewurzelung von Stecklingen und der Cytokiningehalt der Donor-Pflanzen signifikant negativ miteinander korreliert waren (rs bis zu -0,93). • Die in dieser Arbeit erzielten Ergebnisse zeigen klar, dass auch junge Zweige von adulten tropischen Bäumen praktisch kein Bewurzelungspotential mehr besitzen. Bei Jungpflanzen lässt sich dieses Potential durch Stress aktivieren, wobei sich die Akkumulation von Reservestoffen und die Absenkung des Cytokininspiegels günstig auf die Bewurzelungsrate auswirkten. Diese Konstellation war bei Tabebuia stärker ausgeprägt als bei Heliocarpus. • Da die Möglichkeit besteht, aus einer Donor-Pflanze mindestens zwei Stecklinge zu schneiden, ist der Aufbau einer Anzucht geklonten Materials für die Wiederaufforstung zwar zeitaufwändig, aber durchaus realisierbar.
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Vegetation ecology of springs: ecological, spatial and temporal patterns
(2009)
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Volker Audorff
- Acidification is a phenomenon, which affected the forested catchments of the northern hemisphere severely over recent decades. Acidic depositions depleted the buffering capacities of soil and groundwater, what lead to an impairment of forests, headwaters, and lakes. Even though the depositions were reduced considerably since the early 1990s, the recovery of catchments was found to occur time-delayed. The grade of recovery was found to vary significantly between regions. Biomonitoring is an appropriate tool to detect spatial and temporal patterns of ecosystem alterations, such as acidification and recovery. However, to know the interrelationships between organisms and their environment is an indispensable precondition for the identification of indicator species. The complexity of ecosystems and ecological processes hampers this quest oftentimes. Springs provide a natural setting that minimises such constraints. Compared to other habitat types, external factors are less relevant, which makes it easier to relate changes in species abundances to changes in their environment. Studying this species-environment relationship, here the response of plant species to the acidification of the spring waters was of particular interest. In a survey of five regions in Central Europe - taking spatial, hydrophysical as well as hydrochemical parameters of the springs into account - it was clearly shown that the species composition of springs is essentially determined by the spring water chemistry, and more precisely by the gradient of acidity and nutrient availability. This connection was reflected by spatial patterns within and between the regions. These patterns provide useful ecological information about spring water quality and in return about the acidity status of their forested catchments. Including catchment traits - like bedrock, climatic parameters, and forest vegetation - in the analyses, these emerged to be relevant for the species composition of springs, but less than the spring water chemistry. A path analysis showed that the catchments affect the vegetation of springs not directly, but indirectly via the determination of spring water quality. Hence, the catchments are a part of the functional chain, which is driven by the atmospheric depositions. The pH-value was found to represent the gradient of acidity and nutrient availability best. It can serve as a proxy measure that can be related to species occurrence and to species dynamics respectively, aiming to identify indicator species for assessing the status and alterations of spring water quality. With the aim to delineate niche optima and amplitudes, which in return can serve as indicator values, the realised niches of spring-inhabiting species were modelled with respect to pH. The niche attributes were found to be a matter of sampling scale. Larger plot sizes (grain) weakened the species-environment relationship, what consequently resulted in broader niche amplitudes. In contrast, the grain did not influence the species’ pH optima. Monitoring approaches that target to assess processes in time, such as acidification and recovery, are dependent on the response time of indicator species to changes in their environment. Investigating an interval of four consecutive years, inter-annual variability of the species composition could not be attributed to changes in the acidity of the spring waters. Looking at single species, bryophytes did not show a higher sensitivity to the inter-annual variability of the environment than vascular plants. Actually, only a minority of all species featured abundance changes which were significantly correlated to variations in spring water acidity. Our results suggest that the species inertia retards the vegetation dynamics of forest springs. A delayed or long-term integrating response of potential indicator species must be considered when evaluating their indicator suitability. In conclusion, the biomonitoring of spring water acidification or recovery is expedient only for longer time intervals. In a nutshell, the vegetation of springs is closely related to the hydrochemical traits of the spring waters, in particular to a gradient of acidity and nutrient availability. Individual species as well as whole plant communities are suitable indicators which allow for the monitoring of the acidity status of forested catchments. The results of this study contribute to a better understanding of the species-environment-relationships, and in return to an improvement of indicator systems.
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Transport Processes of Reactive Trace Gases in the Atmospheric Boundary Layer
(2009)
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Jens-Christopher Mayer
- Transport of trace gases within the atmospheric boundary layer plays a key role in feedback processes between the earth’s surface and the atmosphere and consequently in ecosystem budgets of carbon and nitrogen (among many more). For a correct quantification of the exchange between surface and atmosphere, it is crucial to understand the transport processes involved and to determine limitations of the presently available measurement techniques in order to apply the right technique with respect to the currently active transport processes. This dissertation focuses on three topics: (a) The analysis of effects of vertical transport mechanisms on surface measurements of trace gases, (b) the appropriate choice of an experimental setup to assess specific measurement errors of moving measurement systems and (c) the application of a series of measurement techniques for surface fluxes of reactive trace gases to determine their degree of agreement and to assess potential source of deviations. To study the impact of vertical transport mechanisms on surface trace gas measurements, this thesis presents a comprehensive set of measurements at the surface and within the atmospheric boundary layer (by tethered balloon). It enables the attribution of a recurrent negative excursion of ozone mixing ratios in the morning hours at a mountain summit to a very efficient vertical transport by free convection. It has been shown that, due to the rapid vertical transport, a layer of approximately 20 m thickness developed at the equilibrium height of the free convection, being located within the residual layer. It had a chemical composition similar to the air close to the ground while being surrounded by residual layer air masses. Hence, very strong gradients of the chemical composition were found within the residual layer. Evidence was found, that such a transport occurs rather frequently at this location, affecting at least 18 % of the days between April and September. To assess measurement errors introduced by the application of scanning methods as compared to gradient approaches, a higher temporal resolution of the vertical profiles was needed. Because of limitation inherent to a tethered balloon, an elevator based profiling system was installed, providing a temporal resolution of 10 minutes with a maximum ceiling of 100 m. Prior to the investigation of transport processes, the proper functioning of correction algorithms for the so-called dynamical error was investigated under real atmospheric conditions. This dynamical error is inherent to all moving measurement systems and arises from the non-zero response time of the deployed sensors. It has been shown that existing algorithms as well as one developed by the authors reliably balance the dynamical error. Furthermore it has been demonstrated, that the elevator data correlate with reference data at fixed levels with coefficients of determination being always greater than 0.992 at every level (10, 20, 40, 60, 80, 98 m). To evaluate the applicability of different flux measurement techniques for the determination of surface fluxes of reactive trace gases, three different approaches were compared. In order to determine surface fluxes of trace gases, a new modification of the modified Bowen ratio method was used. In this modification, the measurements of sensible heat flux and of the gradients were horizontally separated. This allowed the simultaneous measurement of the fluxes of various trace gases without creating errors due to flow distortion by bulky inlet systems. Surface emission fluxes of nitric oxide were found to be in the range 0.02 – 0.15 nmol m-2 s-1 (night/day), nitrogen dioxide fluxes varied around 0.1 nmol m-2 s-1 (deposition) with slightly positive values in the early afternoon, indicating emission. Ozone deposition fluxes ranged from close to zero to about 6 nmol m 2 s-1. A laboratory parameterization of biogenic soil emission fluxes of nitric oxide from incubated soil samples yielded values from 0.025 nmol m-2 s-1 to 0.12 nmol m-2 s-1 for environmental conditions encountered during the field campaign. This was in excellent agreement with the fluxes from field observations. Besides the comparison of field fluxes with laboratory data, a case study (1 night) comparison of carbon dioxide and ozone fluxes between two field methods was done. Results from the modified Bowen ratio method have been compared to fluxes derived from the integral boundary layer budget method. Both methods yielded similar mean carbon dioxide fluxes during the night. In contrast, ozone fluxes deviated between both methods. This deviation was attributed to chemical in-situ loss of ozone during night time within the profile being integrated by the budget method.
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Transport of oxygen in roots of rice (Oryza sativa L.) and of water in developing grape berries (Vitis vinifera L.)
(2009)
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Lukasz Kotula
- 1. Radial oxygen flow in rice roots Measurements of ROL from rice roots grown in either aerated or deoxygenated conditions were combined with root histochemisrty and biochemistry. It was shown that the pattern of ROL from rice roots of stagnantly grown plants correlated with development of the apoplastic barriers in the root peripheral layers. Deoxygenated conditions induced early development of exodermal Casparian bands and suberin lamellae. In addition to suberization, early lignification of walls of densely packed sclerenchyma cells was found closer to root apex in stagnantly grown plants. Supporting these findings, biochemical analyses revealed that, when grown in stagnant solution, the amounts suberin and lignin in the OPR of all investigated zones were several folds greater than those of plants grown in aerated solution. Similarly, the highest rates of radial oxygen loss from plants grown in deoxygenated conditions were observed just behind the apex and decreased dramatically towards the base. By contrast, ROL from adventitious roots of plants grown in aerated solution was the highest at 30 mm from the apex. A new perfusion technique has been developed to measure the oxygen permeability coefficient of the outer part of root (OPR) of rice. Perfusion of aerenchyma of root segments with mixtures of O2/N2 of known oxygen concentration and at the same time measuring radial losses of oxygen allowed quantification of the permeability coefficient of the cell layers exterior to aerenchyma. The new method for measuring the O2 permeability of the outer part of roots (POPR) was applied to rice grown in either aerated or deoxygenated conditions. The results showed that POPR decreased along the root of plants grown in both conditions. However, when grown in deoxygenated medium, the O2 permeability across the OPR was lower by an order of magnitude at all tested distances compared with plants grown in aerated solution. The lower POPR in roots grown in stagnant solution as well as the reduction of POPR along the roots of plants from both conditions strongly correlated with the development of apoplastic barriers in the OPR. In order to estimate the contribution of apoplast and living cell for the overall movement of O2 across the OPR, the POPR was affected either by blocking the apoplastic pores in the OPR with salt precipitates or by killing the living cells with 0.1 N HCl. Treatment with CuSO4/K4Fe(CN)6 resulted in formation of brown precipitates only in roots of plants grown in aerated solution. This is strong evidence that well-developed apoplastic barriers impeded ion movement across the OPR in these plants. As a result of the formation of salt precipitates in the apoplastic pores of roots grown in aerated solution, the POPR decreased by 20-5%. This is in agreement with earlier findings of diffusional and bulk water flow of Ranathunge et al. (2005). The blockage of the apoplast with precipitates reduced the diffusional water permeability by about 20% and caused a massive 3- to 4- fold reduction of hydraulic conductivity. This suggested that the OPR of rice allow rather high water flow in the presence of relatively high resistance to O2. Killing of root segments by 0.1 N HCl increased the POPR by 20-55% of plants grown in both conditions. At least in part, these increases may result from vanishing the respiratory activity in the OPR. Overall, the physical resistance plays a dominant role in impeding O2 loss from rice roots, although effects due to respiration may contribute, namely, in the presence of low rates of radial oxygen loss. 2. The hydraulic properties in developing grape berries cv. Shiraz and Chardonnay Berries of Vitis vinifera cv. Shiraz can undergo weight loss during later stages of ripening. It was hypothesized that the reduction in net vascular flow of water into the berry, concomitant with transpirational loss, leads to berry weight loss. There may be also some back flow of water from the berry to the parent plant along the xylem. The present study provided first data on the hydraulic properties of the pedicel and components of the berries during development. Comparisons were made between Shiraz and Chardonnay, which is not known to undergo substantial weight loss. It was shown that the hydraulic conductance of single berries of both varieties declined during development. However, Shiraz berries had higher hydraulic conductance than Chardonnay for whole berries and all cut positions. The increase in hydraulic resistance was found in the proximal (brush) region and distal part of the berry, which was much larger in Chardonnay. For Shiraz, the increase in resistance was about half of that observed in Chardonnay. There was no evidence for changes in resistance in the pedicel or receptacle region of the berry. The reduction of hydraulic conductance could be due to combination of both: reduced aquaporin activity in the xylem parenchyma and restrictions in the xylem vessels.
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Theory and Practice of Reconciliation in Rwanda
(2009)
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Marcus Grohmann
- During recent years, scholars working on the peacebuilding process in Rwanda have often tended to single out specific aspects, for instance judicial responses to the genocide. Little research has been done, however, on the diversity of approaches that constitute the “reconciliation landscape” in Rwanda today. Basing itself on data from field research in 2006, this paper seeks to shed some light on the many programmes carried out in Rwanda related to reconciliation work. Emphasis is put on two case studies. While establishing a theoretical framework of the reconciliation process in the first part of the paper, the following chapters attempt to explain how this relates to the practice of reconciliation in the Rwandan context. The data collected suggest that in the face of political constraints, the Rwandan government must in part rely on civil society actors for the achievement of their goals of “unity and reconciliation”. The multitude of initiatives from actors with a wide range of motivations and approaches should be seen as complementary, while some may have to make up for the shortcomings and constraints of others.
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The Social System of the Round-Eared Sengi(Macroscelides proboscideus)
(2009)
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Melanie Schubert
- In the present study, I investigated the social system of the round-eared sengi (Macroscelides proboscideus), a small crepuscular mammal that lives in deserts and semi-deserts in Southern Africa. For studying the evolution of monogamy in this species, I determined the social organisation and male mate guarding in wild animals in the Goegap Nature Reserve, South Africa. Data were collected over three successive breeding seasons and one non-breeding season by radio-tracking and trapping over a period of 2.5 years. Additionally, extra-pair attempts of paired round-eared sengi males were studied under laboratory conditions using direct behavioural observations and morphological and physiological characteristics of males. In the field, the population was characterised by balanced adult sex ratios and by a lack in sexual dimorphism in body mass. Round-eared sengis lived in perennial territorial male-female-pairs. However, males maintained much larger areas than females that were sensitive to population density and the presence of neighbouring males. At higher density males used smaller areas than at lower population density, but the number of neighbouring males was fairly constant throughout the whole study. Male space use appeared to be primarily limited by the presence of neighbouring males. In contrast, females maintained smaller-sized territories despite changes in population density. Some paired males attempted to take over widowed females, but shifted back to their original home range following the intrusion of an un-paired male, possibly because of a low variation (about 10 %) in body mass. Female reproduced 2-3 litters during a long breeding season with an asynchronous birth interval between neighbouring females, favouring pre-copulatory and oestrus mate guarding. Males over-mark their females’ scent while following, possibly for concealing the females’ reproductive state and advertisement of the paired status, thereby decreasing the risk of intrusions by competitors. Mate guarding incurred costs, because, overall, males lost about 5 % of their body mass. On the individual level, male body mass loss was negatively related to the intensity of mate guarding during the pre-copulatory period. Furthermore, guarding was inversely correlated with male body mass in the pre-copulatory period and with the number of neighbouring males during oestrus, indicating that males vary their guarding effort in relation to their physical capabilities and the competitive environment. In addition, both sexes demonstrated promiscuous tendencies in the experimental study. Female sexual behaviour with male neighbours was positively related to the time neighbouring males spent in the females’ area and to male marking behaviour. Intruding males were attacked by resident males. Aggression experienced by intruding males was associated with body mass loss (about 4%) in these animals; the same was found for marking behaviour in the neighbouring area. Furthermore, glucocorticoid levels, determined from analyses of faeces and urine samples, positively correlated with male body mass loss and also with male marking behaviour of intruding males, indicating costs of this behaviour. In conclusion, pair-living is the predominant social organisation in round-eared sengis. Males suffered from a limited opportunity to monopolise more than a single female that may have resulted from females living solitarily in small exclusive territories, balanced adult sex ratios and a low variation in body mass between males. However, both sexes have promiscuous tendencies. Female reproduced asynchronously and advertised their reproductive status, so that males pursued two behavioural tactics: Males engage in mate guarding for ensuring paternity and also in extra-pair attempts with neighbouring females, possibly for increasing their reproductive success. Since both male tactics were energetic costly they may serve as honest signals of quality providing only higher quality males with the opportunity to enhance their reproductive success, which in turn may be advantageous for females in terms of genetic benefits.
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The evolution of Saharan dust input on Lanzarote (Canary Islands) – influenced by human activity in the Northwest Sahara during the early Holocene?
(2009)
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Hans von Suchodoletz
Hedi Oberhänsli
Dominik Faust
Markus Fuchs
Cécile Blanchet
Tobias Goldhammer
Ludwig Zöller
- An overall Holocene increase of Saharan dust input to the Canary Islands and to the North Canary Basin is accompanied by a strong coarsening of Saharan dust in loess-like sediments deposited on Lanzarote from ~7–8 ka. No similar coarsening events are indicated in investigations of the sedimentological record for the last 180 ka, a period showing several dramatic climate changes. Therefore a mobilisation of Holocene dust by anthropogenic activity in the northwest Sahara east of the Canary Islands is assumed. Although scarce archaeological data from the coastal area of that region does not point to strong anthropogenic activity during the early Holocene, a high density of unexplored archaeological remains is reported from the coastal hinterlands in the Western Sahara. Thus, the hypothesis of early anthropogenic activity cannot be excluded.
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Tailoring Fluorene-based Oligomers for micron and sub-micron sized Photopatterning
(2009)
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Esther Scheler
- This thesis describes the work on tailor-made synthesis, characterization and application of well-defined fluorene oligomers for photoptatterning. Two types of fluorene oligomers are presented: pure fluorene oligomers and fluorene cooligomers incorporating various comonomers for adjusting the conductive properties towards electron and hole conduction. Since possible applications for these materials feature organic light emitting diodes and organic field effect transistors we focused on the preparation of well-defined and defect free oligomers and the preservation of their electro-optical properties during photopatterning. Further on the requirements for material synthesis are easy procedures and large quantities. Therefore we developed an approach, which produces large quantities combined with the adjustment of the desired properties in one single step. The synthetic strategy throughout the thesis comprises the addition of an endcapping species in aryl-aryl polymerization reactions. The tailormade endcapper fulfils three tasks at once, the control of the molecular weight, the introduction of polymerizable acrylate moieties and the avoidance of undesired endgroups. As aryl-aryl coupling methods the nickel catalyzed Yamamoto and palladium catalyzed Suzuki condensations were applied. The completeness of the aryl-aryl coupling and the endcapping was proven with Maldi-ToF mass spectrometry. With this approach the properties of the oligomers can be easily adjusted in view of the optimization of their photopolymerization behaviour. The first oligomer series deals with the effect of the molecular weight on the properties and photopatterning behaviour of pure fluorene oligomers. The molecular weights were controlled by the amount of functionalized endcapper, which carried the polymerizable acrylate groups. As coupling method the Yamamoto coupling was applied. The molecular weights defined the temperature range of the nematic mesophases. An increase of the average chain length leads to higher transition temperatures (e.g. clearing temperatures) and to better film forming properties. The photopolymerization is usually performed in the nematic state to achieve a sufficient mobility of the acrylates. The irradiation conditions had major consequences on the preservation of the characteristic electro-optical properties of the fluorenes, the harsher the conditions the higher was the probability to destroy the chemical structure by photooxidation. Further on since each chain only carries two acrylate functionalities attached to the endcappers the total number of acrylates is different for high molecular weight and low molecular weight mixtures. The lowest molecular weight mixture contains the most acrylates and shows the lowest transition temperature, which leads to the best micron sized photopatterns. The second generation of pure fluorene oligomers demonstrate how different contents of polymerizable groups affect the photopolymerization behaviour. Here the molecular weights were kept constant around 5000 g/mol by equal amounts of endcapper and the acrylate groups were introduced by the fluorene monomers. The Yamamoto coupling was used and upon cooligomerization with a non-acrylate fluorene monomer the acrylate content was changed from 10% to 100%. The photopolymerization times strongly depend on the acrylate content, the 100% acrylate oligomer could be photopatterned in 30 seconds, whereas the 80% and 60% mixtures needed 2-5 minutes. In the best case crosslinking is 20 times faster than found for the preceding series, which ensures the preservation of the electro-optical properties. With the highest acrylate content a photocrosslinking even at room temperature became possible. The third oligomer series describes the incorporation of various comonomers such as TPD and bithiophene via Yamamoto reaction. Taking the knowledge of the two preceding generations into account we exploited the acrylate monomer from series two and introduced 30% comonomer. This ensured a sufficient content of acrylates for a fast photopatterning and enough comonomer for a shift of the electronic properties. We found that the electronic structure of the comonomer strongly affected the behaviour in the Yamamoto reaction. The HOMO and LUMO energy values were shifted towards hole or electron conduction. The photopatterning conditions were similar as found for the corresponding pure oligofluorenes with a 60% acrylate content. 2-5 minutes exposure produced highly emissive micro patterns. Thus a change of the electro-optical properties does not affect the photopolymerization behaviour and vice versa. Since we found differences in the incorporation of comonomers during Yamamoto coupling we applied the Suzuki coupling, which ensures an alternately linkage of monomers. Here protective groups had to be used since the Pd-catalyst does not tolerate acrylate functionalities. We found that the comonomers were incorporated quantitatively, but the major difficulty proved to be the polymeranalogous reactions following the polycondensation. The energy values were shifted towards electron and hole conduction and the photopolymerization behaviour was similar to the Yamamoto oligomers. An exposure time of 2-5 minutes produced patterns with a maximum resolution of 700 nm. To conclude the endcapping strategy combined with the Yamamoto coupling is a most effective tool for the adjustment of properties within one single step. The acrylate content as well as the molecular weight can be precisely tuned, which allows a good control of the photopatterning properties. In some cases e.g. with electron withdrawing comonomers the Suzuki cross coupling is the method of choice.
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Tagungsband zum Doctoral Consortium der WI2009
(2009)
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Torsten Eymann
- Bereits seit Anfang der 1990er Jahre wird jungen Wissenschaftlern im Vorfeld der Tagung "Wirtschaftsinformatik" ein Doctoral Consortium als unterstützendes Forum angeboten. Diese Einrichtung wurde auch zur größten internationalen Konferenz der Wirtschaftsinformatik, der WI 2009 in Wien fortgeführt. Dieser Band fasst die zum Vortrag ausgewählten Beiträge zusammen.
